_ Mammals
Е ol the
soviet Union
VOLUME I
Part 16
о Неро
М.Е Naumov
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, Part] — 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 1b) 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 1b
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. 1)
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. 1b 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. 1b. Carnivora (Weasels; Additional species)
vol. 2.—pt. 2. Carnivora (Hyaenas and cats).
1. Mammals— Soviet Union. I. Nasimovich, A.A. II. Bannikov,
Andrei Grigor’evich. III 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 1b
Family Mustelidae (Martens, Weasels, etc.)
Key for Identification of Genera in the
Family Mustelidae
Subfamily Mustelinae
Genus Martes Pinel, 1792 (Martens)
Key for Identification of Species in the
Genus Martes
Martes (Martes) zibellina (Linnaeus, 1758)
(Sable)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Martes (Martes) martes (Linnaeus, 1758)
(Forest Marten)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Martes (Martes) foina Erxleben, 1777
(Stone Marten)
Diagnosis
Description
Systematic Position
735
746
749
749
155
757
759
59
770
770
787
ПОТ
823
825
826
826
836
836
847
850
872
874
875
876
882
vi
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Martes (Charronia) flavigula Boddaert, 1785
(Kharza, or Yellow-throated Marten)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Genus Gulo Storr, 1780
Gulo gulo (Linnaeus, 1758) (Wolverine)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Genus Mustela (Linnaeus, 1758)
Key for Identification of Species in the Genus
Mustela
Mustela (Mustela) nivalis Linnaeus, 1766
(Weasel)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Mustela (Mustela) erminea Linnaeus, 1758
(Ermine)
Diagnosis
Description
Systematic Position
Geographic Distribution
883
891
895
904
905
905
905
910
910
912
914
919
920
925
926
926
931
932
940
943
953
954
962
965
966
967
972
972
975
985
995
995
997
997
1003
1003
Geographic Variation
Biology
Practical Significance
Mustela (Mustela) altaica Pallas, 1811
(Solongoi)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Mustela (Mustela) sibirica Pallas, 1773
(Kolonok or Siberian Weasel)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Mustela (Mustela) lutreola (Linnaeus, 1761)
(Russian, or European Mink)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Mustela (Putorius) putorius (Linnaeus, 1758)
(Forest, or Black, Polecat)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Vil
1008
1016
1029
1030
1031
1031
1036
1037
1043
1047
1051
1051
1052
1052
1058
1058
1065
1069
1078
1078
1080
1080
1086
1088
1094
1099
1106
1107
1108
1108
1115
1117
1124
1126
1132
Viil
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 J Pi)
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) 12
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
Subfamily Lutrinae
Genus Lutra Brisson, 1762 [=Briinnich, 1771]
Lutra lutra* (Linnaeus, 1758) ({Eurasian] Otter)
Diagnosis
Description
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Genus Enhydra Flemming, 1822
Enhydra lutris (Linnaeus, 1758) (Sea Otter,
or Kalan)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Supplement to the Order of Carnivores
Family Procyonidae (Raccoons, etc.)
Subfamily Procyoninae Gill, 1872
Genus Procyon Storr, 1780
Procyon (Procyon) lotor (Linnaeus, 1758)
(American Raccoon)
Diagnosis
Description
Systematic Position
Geographic Distribution
Geographic Variation
Biology
Practical Significance
Family Mustelidae
Mustela (Mustela) vision Schreber, 1777
(American Mink)
Diagnosis
Description
Systematic Position
*In Russian original, Lutra (Lutra)—Sci. Ed.
Geographic Distribution
Geographic Variation
Biology
Practical Significance
LITERATURE CITED
INDEX OF LATIN NAMES OF TAXA*
1399
1403
1405
1415
1416
1559
*The Russian original also included ап Index of Russian Names, omitted here.
491
492
493
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.
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 (Enhydra). 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. Та].
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
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 Ae II
mula is I 3 С ji Р я М 5 =38. This formula is not encountered in
all species of the family (developed in martens, Martes; wolverine,
Gulo; badgers s.s., Meles, Helictis, Arctonyx and others). A re-
duction in tooth number often occurs in the premolars and molars.
4 8 teh Sheil Ayig $
This occurs in various combinations ет Е aimee
Sel 3 1
(otters, Lutra); Е Р.М ain 34 (polecats, weasels, Mustela,
Eyra, Galictis; American badgers, Taxidea; skunks Spilogale,
1 |
Mephitis) and even [12+] Р ЭМО =32 (skunks, Conepatus) and
За (4
13с1 Р 3 M in 32 (ratel, Mellivora). In one case (sea otter, or
kalan, Enhydra), reduction occurs in the number of incisors re-
Si ant MRC S|
tained, and the formula takes the form I, CF ae М> =32. The great-
est reduction, however, is reached in the number of molars and
2
3 1 1
premolars 131 Р 2Мт= 28 (Lyncodon). In rare cases, all ог
some of the premolars are absent, and the dental formula signifi-
cantly varies individually, with age, as well as geographically—the
dental formula may be P - P - P Е P : = 38 = 36 = 34 (badger, Meles
meles)'.
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.
494
ЭЛ
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
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.
495
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
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.
497
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° М. 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
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).
‘romday ‘9’ ‘aeprlfaisnyy ‘[$12$еэм] зиэзтеш jo Апигеу Jo э8ие ‘731 ‘84 967
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,
498
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).
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; | 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, cr 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.
499
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-
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.
500
746
Key for Identification of Genera of the Family Mustelidae
16(2):
2b):
3 (4).
4 (3).
5 (6).
6 (5).
7 (8).
Digits of the forelimb conjoined, and entire paw герге-
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
о genus of sea otter or kalan, Enhydra (page 1330).
Digits of the fore and hind limbs are not conjoined and
movable (sometimes perhaps connected by membrane),
hind not forming flippers and middle digit [III] is longer
than others. In lower jaw, 3 incisors on each side, molars
and posterior premolars with well developed pointed tu-
bercles or with sharp cusps.
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*;(Figureil’85) sta ines Li eee ee ee
eh oe tela genus of honey-badgers, Mellivora (page 1207).
External ear conch present, color of one sort or another.
Molars and premolars 5—6 in lower jaw.
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).
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.
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.
‘NY SSS
ПУ
i
We
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
8 (7).
9 (10).
501
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)
Soe ae sad oct ее: genus of otters, Lutra (page 1283).
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.
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
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
10 (9).
11 (12).
genus of otters, Lutra. Sketch Бу М.М. Kondakov.
passing along sides and across croup from shoulder to
Shoulder ........-..:..:5 genus of wolverines, Gulo (page 920).
Dimensions smaller—body length less than 65 cm,
condylobasal length of skull less than 115 mm. Tail length
variable; coloration otherwise.
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)
о ЗИ genus of martens, Martes (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.
a (g wd I)
In upper jaw behind canine, 4 teeth, in lower—5S.
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) ........
НЯ cle genus of marbled polecat, Vormela (page 1176).
14 (13). Coloration otherwise, without spots or stripes. Processes
of pterygoids do not reach tympanic bullae ..............
genus of ermines and polecats, Mustela (page 954) (V.H.).
749
Fig.188. Position of pterygoid processes in genus Vormela (joined with tympanic
bullae) and in the other genera of the weasel family (not joined with tympanic
bullae—black [common] polecat, Mustela putorius L.). Sketch by N.N. Kondakov.
502 Subfamily of Weasels
Subfamilia Mustelinae Gill, 1872
Genus of Martens
Genus Martes Pinel, 1792
1775. Martes. Frisch. Natur-Syst. d. vierfiiss. 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. Kaup. 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.
503
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.
I 2 С Р ы М a2 38
Dental formula ii. Sana
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.
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, М. 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
505
152
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.
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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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 genera 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, М. 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).
506
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
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) ...............6.
Sry yellow-throated marten, М. (Ch.) flavigula (р. 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
PARA OOM RE) RAE seen 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
507
756
5 (6).
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.
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),
On(S) se
Martes (Martes) zibellina L., pine marten, Martes (Martes) martes L., and stone
marten, Martes (Martes) foina Erxl. Sketch by М.М. Kondakov.
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
И 8 Ot. 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)”—$с1. Ed.
1758:
1840.
1844.
1855’.
1855.
1855.
1855.
1855.
1855.
1855.
1918.
757
Subgenus of True Martens
Subgenus Martes Pinel, 1792
SABLE
Martes (Martes) zibellina Linnaeus, 1758
Mustela zibellina. Linnaeus. Syst. Nat. Ed. X. 1. p. 46.
Northern Siberia. According to Ognev (1931) and other
authors, more precisely: western Siberia north of Tobol’sk.
Mustela melampus. Wagner. Schreb. Saugeth. Suppl., 2,
p. 229. Japan.
Mustela brachyura Temmink. Siebolds Fauna Japon.
Mamm., p. 33. Japan, Hokkaido.
Mustela zibellina var. asiatica. Brandt. Mém. mathem.,
phys. et natur., 7, р. 6, 23, Taf. 1, Fig. 1. Kamchatka.’
M. zibellina var. alba. Brandt. Ibidem, p. 14, Taf. 2. Fig.
5.
M. zibellina var. fusco-flavescens. Brandt. Ibidem, p. 14,
Rafio2Figs6) 'ТаЕ. ‘ЗЕ. 7:
М. zibellina var. осйгасеа seu ferruginea. Brandt. Ibi-
dem, p. 14. Taf. 3, Fig. 7.
M. zibellina var. maculata. Brandt. Ibidem, p. 14, Taf. 3,
Fig. 9.
Mustela zibellina var. asiatica rupestris. Brandt. Ibidem,
afM2wRige2!
Mustela zibellina var. asiatica sylvestris. Brandt.
Ibidem, Taf. 2, Fig. 3.
*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.
п 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 (“уаг. 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.
508
758
Т922.
1922.
1922.
1922.
1.923.
1925.
1925:
1925:
1927.
1927.
1941.
1941.
1941.
1943.
1947.
Mustela zibellina princeps Birula. Byalynitskii-Birula.
Ezheg. Zoologich. muzeya. Ross. ak. nauk 1917-1921,
22, p. 8. Barguzin range.
Mustela zibellina var. kamtschatica. Dybowski. Arch.
Tow. Nauk. Lwowe, 1, р. 349, Nomen nudum. Kamchatka.
Mustela zibellina var. baicalensis. Dybowski. Ibidem, p.
349. Nomen nudum. Vic. Baikal.
Mustela zibellina var. amurensis. Dybowski. Ibidem, p.
349, Amur, Ussuri.
Mustela melampus coreensis*. Kuroda et Mori. Journ. Mam-
mal. 4, p. 27. Tenan, South[ern]. Chusei distr[ict]. Korea.
Martes zibellina yeniseensis. Ognev. Journ. Mamm. 6,
No. 4, р. 277. Krasnoyarsk u[yesd], Yenisei guber[naya],
taiga plain.
Martes zibellina sajanensis. Ognev. Ibidem, p. 278.
Orzybai [=Orsyba] river, northern Sayan [mountains].
Martes zibellina sahalinensis. Ognev. Ibidem, p. 279.
Sakhalin, Vedernikovo. [=Saghalien, Wedernikovo].
Martes zibellina correensis. Kishida. Choju Chosahokoku,
4, p. 130. Korea.
Martes zibellina hamgyensis. Kishida. Dobuts Zasshi, 39,
р: 509.**
Marites zibellina tungussensis. Kusnetzov. Tr. Mosk.
zootekhnich. inst., I, p. 116. Lower Tunguskaya [river].
M.(artes) z.(ibellina) sahalinensis arsenjevi. Kusnetzov.
Ibidem, p. 122. Samarga River, Ussuri Territory.
M.(artes) z.(ibellina) sahalinensis schantaricus. Kusnetzov.
Ibidem, p. 122. Bolshoi [Great] Shantar Island, Okhotsk Sea.
°%Martes zibellina averini. Bashanov. Bazhanov.
Kazakhskii fil. Ak. N. Souza SSR in 1942, p. 13.
Katon-Karagaiskii region, Vostochno-Kazakhst. obl[ast],
southern Altai.
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.
Туре (Zoological museum, Moscow university) from left bank of middle course
of Bystrukha river, upper Katun [river]; southwestern Altai (V.H.).
509
759
1955. Martes zibellina tomensis. Timofejev et Nadejev.
Timofeev et Nadeev. Sobol’, 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. Маме; zibellina vitimensis. Timofejev et Nadejev.
Timofeev et Nadeev. Ibidem., 47. Valley of the Mama
river, right tributary of the Vitim [river], Yakutiya.
1955. — Манеу zibellina obscura. Timofejev et Nadejev. Timofeev
et Nadeev. Ibidem., р. 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, р. 185. Yakutiya, Aldan Valley!’ (V.H.).
Diagnosis
Tail length with terminal hairs not more than half the body length.
Color uniform, on throat and chest a yellow spot of irregular form;
sometimes absent. Head usually lighter than back, sometimes
whitish. Bony auditory bullae relatively elongated and adjacent.
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 Бу С.А. 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
М. (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.
510
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 сп? 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.
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
512
Sid
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.'”
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
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.
514
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.
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
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513 Fig. 191. Skull of the sable, Martes (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; В) Age group II—1 year, 8-10
months; С) Age group III—2 years, 8-10 months; D) Age group IV—about 3
years, 8-10 months and older.
515
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 (М 1.06); in females, 0.78-1.14 (М 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 (М 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 (М 1.20);
in females, 1.08-1.33 (М 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 (М
1.30); in females, 1.17-1.43 (М 1.24).
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)3. 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
С
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-М448—630.0% of the
length of the carcass without the hide (365-М422—490 mm). Heart
weight is 5.23-M8.82-15.20 gm, ог 9.0-M12.2-21.5%o (п 117);
lung weight (without trachea) 8.10-М14.9-52.50 gm; liver weight
10.32-M14.90-45.7 gm; kidney weight 1.40-М2.76-5.35 gm;
spleen weight 0.62-М1.65-3.90 эта“ (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-М1472-1810, of females (21), 820-М1132-1560. Weight in
July in males, 1200-М1433-1810; in females, 870-М980-1300
(Kuznetsov, 1941). Optimal weight of breeding sables before the
“Тре 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).
516
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.).
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 (“‘kidas”'*), 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.
517
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
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.
519
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’®.
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°
М. 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
'6Неге 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.
518 Fig. 194. Data on the past and present distribution of the sable, Маме (Martes)
zibellina L. in Europe. V.G. Нершег: 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.
"In 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).
520
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 Ропто* 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
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.
'8References 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.
WAT
region of 55—56° М. 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° М.
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”!.
'SSabaneev’s assumption (1875, page 200) that it extended to 52° “perhaps 51°
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°, 15 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 51°, 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.
21 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).
521
778
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 П’теп” 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
22Middendorf (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.
522
779
sources of the Lyapin at approximately the latitude 63° 30° М. 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-
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° М. (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 Smirnov, 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
524
781
reached the Barabinsk forest-steppe. Here the range boundary went
to the mouth of the Om, apparently, north of Lake Chana 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
UI’ba basins, almost reaching the Irtysh. In the south, the bound-
ary included the Kurchum and the region of Lake Markakol’
(Ognov, 1931; Yanushevich and Blagoveshchenskii, 1952;
Afanas’ev et. al., 1953; V.G. Heptner). At Markakol’, the bound-
ary of the range reached the state Богаег”. 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.
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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).
ЕЕ rH
Sa
$
%
ll
nfm
5 Novosibirsk)"
\
5 Sih т
Boundries of Ys ss т ge
“Centers” of
ccurrence of зе < 9
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
ь.-
A
и
Е:
ГИ
aa i
<
=:
ng the period from 1927-1951 (after Nadeev and Timofeev,
1955, with modifications).
ation duri
1. Sites where native form was released, 2. Sites where Vitim sable was released into region of occurrence of another form, 3. Same for Buryat
of sable for the entire range
525 Fig. 197. Principal release sites
sable, 4. Same for Kachatka sable, 5. Same for Bureinsk sable
785
QZ?
A
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.
ad Sable distribution
К | ® in 1935
ан g Sable distribution
aa in 1941/1942
---| @ Sable distribution
in 1950
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, II’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 Q) 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
(CG
7
Е
@ Distribution in 1935 ф 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).
“Аг 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.
0 30 60 90 120
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.
530
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
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
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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-М
85.7-90.6 mm, females (87) 73.8-M 78.9-83.8 mm; zygomatic
531
792
width of males, 44.1-М 48.0-54.8 mm, females 39.0-М 43.6—47.2
mm; height of male skull 29.5-М 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 рр. 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).
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 оне
border of taiga southwards to Chulym river.
Absent outside USSR.
7193
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.) =. 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.
532
794
Body length of males, 394-М 415-420 mm, females, 360-М
382—410 шт; tail length of males, 122-М 144-155 mm, females,
120-М 129-145 mm; condylobasal length of males, 73.9-М 77.9-—
81.0 mm, females, 68.4-М 73.1-79.1 mm; zygomatic width of
males, 39.6-М 41.6-44.5 mm, females, 37.7 М-39.7-41.8 mm;
height of male skull, 29.6-М 30.5-32.3 mm, females, 28.5-М 29.7—
31.4 mm.
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.) =. 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).
533
195
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. Shantar sable, M. (M.) z. schantaricus Kusnetzov, 1941.
Dimensions as in Ussuri sable. Color darker than Sakhalin and
Ussuri. Pelage quite fluffy.
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-М 490-580 mm, females, 415, М
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, М 90.4—94.6 mm (М 88.5 mm-93), females, 79-4, М 82.1-
796
84.4 шт (М 80.8 mm-—76); zygomatic width of males, 51.3, М
53.5—56.2 шт (М 50.2 шт-81), females, 43.6, М 46.3-52.8 mm
(М 44.9 тт-74); skull height of males, 31.6, М 33.5-35.1 mm (М
33.3 ит-94), females, 29.8, М 31.5-39.6 mm (М 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, М 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, М 1239-1600
g, females (25) 800, M905-1130 g, January males (22) 880, M
1193-1500g, females (25) 740, М 916-1120 g; Kamchatka sa-
ble—males (4) 800, М about 1500-1869 g, females (5) 600, М
about 1100-1140 g.
Outside the limits of the USSR, the following forms are noted
(sometimes, several of them are related to “М. театри5”): 1) М.
z. brachyura Temminck, 1844—in Hokkaido, Japan; 2) М. (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 et Mori,
1923 (synonym hamgyensis, Korean Peninsula) (V.H.).
534
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
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 рег 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).
535
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
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 [A. 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 соп-
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.
536
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-
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 the Yenisei, habitat of sable. “Stolby”
preserve south of Krasnoyarsk. Photograph by G.D. Dul’keit.
537
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.
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 TES 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.
539
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 51.0 36.8 73.5 64.0
Siberian chipmunk — 3.9 355) 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 Tell 19.7 13.0 2.0
Birds 23 1.0 6.9 25.0
Tetraonids ТА 3:9 1.2 No information
Hazel hen dst — 1.2 No information
Bird eggs 3.8 1.3 0.6 0.6
Insects 53.8 13.1 Sofi 15.0
Pine “nuts” 3.8 36.8 37.6 34.0
Berries 44.5 131 35.7 66.0
*The Russian word senostavka (“haystacker”) is used, rather than the more сот-
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
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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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 |2
Willow and rock ptarmigan 1.5 1.2
Small birds 125 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 Zl
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
0 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
540
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
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); а 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.
541
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 (О.К. 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 (bonité) 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
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 O[.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, У.[егпоп], 1957—Sci. Ed.
542
808
did not exceed а circle 4 km in diameter (about 12.6 Кт?), 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
km’.
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.
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.
543
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%— т 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
Baikal, the diurnal prevails, and in the Trans-Urals—crepuscular-
noctural. On B[ol’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
543 Fig. 207. Sable in natural habitat. Stolby preserve south of Krasnoyarsk. Photograph
by СР. 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; in 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
544
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,
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 (“атап5”) 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.
545
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 (Dul’keit,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%—т the third year; the remaining animals—
in the fourth year or they do not reproduce at all (Nadeev and
Timofeev, 1955). Termination of sexual activity is observed there
at 13 years; individual females remain fertile until 15 years old. At
16-17 years, non-productive attempts at reproduction still occur.
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 16-29 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.
546
814
General duration of pregnancy is determined as 273-275
(Starkov, 1947) оп 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%— т
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);
in Barguzin sable, with the calculation of mortality before the be-
ginning of harvest—0.7 (Z.F. Svatosh); in Kondo-Sos’vinsk за-
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), 15 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, т
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-уеаг 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.
547
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).
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 (Manteifel’, 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
548
817
was observed in the West Siberian lowland (Raevskii, 1947), т
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.
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
549
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 17.69%, 32.0% § 251% 1" 14.5900: 59611 2B oe
About 2 years 30.9 20.8 25.1 211.5 12.5 27.
Older than 2 years 26.7 18.2 24.3 12 14.6 2:3
Particularly old 24.8 29.0 24.5 47.0 62.4 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%).
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 3.2 39.4 28:5
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—S0% 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 Seo
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
1958/59 227 2.01 45.6 81.5
1959/60 30.4 0.51 1515 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
550
820
Population dynamics. Changes in their numbers is а 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
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
*In Russian original, literally specific weight—Sci. Ed.
551
821
fluctuations, depending оп а 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.
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 5 x 9 to 8 x 15 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 т; 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.
552
823
The sable foot sinks into the snow for 5-10 cm. Weight loading is
equal to 12-14 g per 1 ст? (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
а 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.).
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
553
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, “уогоюуауа”).
The greatest efforts should be directed towards the develop-
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 MARTEN?**
Martes (Martes) 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.
554
826
1941. Маме; martes uralensis. Kusnetzov. Ibidem, р. 126. Vic.
Miass, southern Urals.
1947. Martes martes sabaneevi. Jurgenson. (Yurgenson). Tr.
Pechoro-lIlychsk. zapov., 5, р. 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
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.
27Some 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.
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Fig. 208. Pine marten, ог zheltodushka [yellow-breast], Martes (Martes) martes L.
Sketch by A.N. Komarov.
555
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
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
556
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
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).
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
8Particularly 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
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558
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
and southern Urals and Bashkiriya have 15—20 (п 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 (п 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.
559
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, М 42.3 mm; weight, 0.20 to 0.35 gm, М 0.29 gm.
In young, length is 35.1-38.3 mm, М 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)*°.
Kidas.*' 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
560
559
560
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)
are as follows: body length of та!ез—390-М 452-485 mm, fe-
males—365—M 406—450 mm; tail length of males—170-—M 187-
240 mm, females—110—M167-180 mm; hind-foot length of
та!ез—75-М 86-98 mm, females—70-M 73-77 mm. Length of
os penis 36.7-М 40.6-—44.7 mm.
Weight of males 600-М 754—980 gm, females 410-М 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-М 81.5-88.2 mm; zygomatic
width 52.1-М 55.6-58.1 mm; skull height 36.1-М 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.
* 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
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 сагсаззез?).
561
836
А marten-like postorbital constriction is present т 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.).
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 Iokan’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.
563
838
Following this parallel, in general, or а 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’*.
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° М. 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.
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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 ХХ 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 Tara 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
Balty region (former Baltskii co.; 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.
565
842
From Tambov ог 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,
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
566
844
changes, аз 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 th® 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
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 Tara. The marten existed not only in Tobolsk and Tara
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
п 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, Маме; (Martes) martes L. V.G. Heptner.
847
567 (Sowerby, 1923) is deprived of any foundation. The reference
568
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
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-М 86.26 + 0.46-
88.0 mm, of females (9) is 77.0-М 79.00 + 0.70-83.0 mm; zygomatic
width of males 46.0-М 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-М 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-М 433-452 (and to 500
mm), length of tail 195-M 218-260 mm (Yurgenson, 1947).
Condylobasal length of male skull (32) 75.3-М 80.19 + 0.42—
84.8 mm, of females (18) 71.1-М 74.71 + 0.55-77.3 mm; zygomatic
width of males 41.4-М 44.36 + 0.36-49.0 mm, females 38.2-
М 42.22 + 0.32—-44.3 mm; skull height of males 27.4-М 31.07 +
0.14—33.3 mm, females 27.0-М 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.
569
849
Dimensions similar to preceding form. Body length of martens
of upper Pechora (67) 380-М 433—460 mm; tail length 170-М
198-235 mm. Weight (42) in winter 580-М 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-М 83.57 + 0.78-
87.0 mm, female (2) is 74.9-75.0 mm; zygomatic width of male
45.0-М 47.14 + 0.43-49.1 mm, female 43.0-М 43.6-44.2 mm;
skull height of male 31.8-М 33.0 + 0.41-34.9 mm, female 29.4—
М 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
570
850
light-orange ог 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-М 82.36 + 0.62
mm (85.3 mm)*—53.0,* females (14) is 74.6-М 75.43 + 0.45 mm
(78.3 mm)—81.2; zygomatic width of males 43.1, М 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-М 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) М.
(M.) т. notialis Cavazza, 1912—South Italy south of Abruzzi (У.Н.).
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%— т 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
571
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
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.
6Occurrence in food of 20% or more.
572
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.
Seasonal variation of food is also well illustrated. For exampie
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) Lapiand—587 (Nasimovich, 1948);
574
854
2) upper Pechora—593 (Е.М. Teplova and Р.В. 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 Middle Zhiguli Northern
Pechora zone 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 1:5 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, %
Minimum Maximum
Voles 7.8 29.4 23.3
Squirrel 1357, 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.
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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 (Héglund, 1960)*, 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 All
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 12.5 715.5 45.1
Squirrel 0 17.5 7.1
White hare 0 92 5.0
Insectivores 0 6.1 2.0
Birds 0 32.5 21.5
Including tetraonids 2.8 22.8 13.6
Bird eggs 0 10.5 3.4
Frogs 0 9.2 3.5
Berries 0 43.1 11.2
Mountain ash 0 17.0 3.2
*Not in Lit. Cite—Sci. Ed.
575
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-
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.
576
859
Shrews in marten food are either an accidental element or ап
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-
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
(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 Кп?. In
the spruce forest massifs of the southwestern part of Kalinin
578
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 (Р.В. 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 т; in the north-
ern Caucasus—at a height of 10-20 т and more), but in certain
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
579
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.
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.
580
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
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.
581
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.
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.
582
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 [Вшео 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 т
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
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.
583
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
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
9-11 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.
584
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.
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 оп
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.
Formozovy, 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.
585
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
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 MARTEN*’®
Martes (Martes) foina Erxleben,** 1777
1977. Mustela foina. Erxleben, Syst. Regni Anim., 1, p. 458.
Germany.
*Special sorts of traps—Sci. Ed.
38Also, belodushka (white-breast), “kunitsa-belodushka” (white-breasted marten)
and among furriers, КипИза gorskaya (mountain marten).
**Misprinted in small lower-case type in Russian original—Sci. Ed.
586
1792
1801.
1869.
1873.
1879.
1906.
1911.
1914.
ТЭД,
1946.
875
Mustela domestica. Pinel. Actes Soc. H.N. Paris, 1, р. 55.
France.
Mustela foina alba. Bechstein. Gemeinn. Naturg.
Deutschlands, 1, p. 759. Thuringia, Germany.
Mustela martes var. fagorum. Fatio. Faune vertebr. Suisse,
1, p. 318. Switzerland.
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’.
Mustela leucolachnea. Blanford. Second Yarkand Mission,
Mammalia, p. 26. Yarkand, northwestern China.
Mustela foina nehringi. Satunin. Izv. Kavk. muzeya, 2, p.
120, tab. 2. Tbilisi. Transcaucasus.
Martes foina bosniaca. Brass. Aus dem Reiche der Pelze, p.
468. Bosnia, Yugoslavia.
Martes foina altaica. Satunin. Opredel. Mlekopit. Ross. Imp.
key for mammals of the Russian Empire. 1, p. 111. Altai.
Martes rosanovi. V. et E. Martino. Zap. Krymsk. obshch.
estestvoisp. lyubit. prirody, 7, 1 (reprint). Northwestern slope
of Chatyr-Dag, Crimea (Crimean preserve; У.Н.).
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.).
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
»
ri
iy
=
Я
{
586 Fig. 216. Stone marten, ог white-breast, Martes (Martes) foina Егх!. 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.
587
588
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
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-
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
AN
ИА
uy hs cn “
И ИИ iy’
CHAN :
И NV ) и г
| sith
Le
587 Fig. 217. Variations in form of throat patch of stone marten, М. (М.) foina Erxl. Upper
row—European marten, М. (М.) Х foina Erxl. (Middle Europe); middle row—first and third
from left—Caucasian stone marten, М. (M.) Е nehringi Sat. (Vladikavkaz and Talysh),
second and fourth from the left—Crimean stone marten, М. (M.) Л rosanovi У. et Е. Mart.
(Crimean preserve); lower row—Middle Asia stone marten, М. (М.) Г 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.
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
7 уха
222
ua
ESN 3
S-
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Wy «+
Ах К <
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<S К А YS
a
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\ Wii? SS SS8 =
в. 3 _ ANN “as №
< = > @ У Sy d
Rs a ft
i
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589 Fig. 218. Skull of stone marten, Martes (Martes) foina Erxl.
590
881
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
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-М 1325-
1950 gm, of females 865-М 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-
М 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-М 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] (М. flavigula) and il’ka [fisher] (М. 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 а 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 (Ezel’ [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).
593
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 Novosil’ (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.
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
3 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.
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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).
594
595
888
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
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, Мате; (Martes) Гота Егх1. (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, PriKosogol’
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;
49No 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.
596
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, [’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.
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)**.
*Ргезепсе 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 М. (M.) } 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-М 82.00 + 0.65-
84.6 mm; of females (5) 77.6-М 78.8—80.0 mm; zygomatic width
of males 47.9-М 51.7 + 0.75-52.8 mm, of females 46.2-М 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.
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893
2. Crimean stone marten, М. (M.) Ё гозапом У. et Е. Martino,
LONG:
Dimensions somewhat smaller than in preceding form. Skull
small, not massive.
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-М 43.8—
48.1 mm, of females 42.6-М 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, М. (M.) } 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 skull*? (13) 81.0-М 83.9 + 0.29
(84.3)-86.0 mm, of females (15) 76.1-М 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).
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894
mm; zygomatic width of the male skull 50.3-М 52.9 + 0.47 (51.0)-
56.4 mm, of females 46.2, М 48.6 + 0.40 (44.9)-50.6 mm; height
of male skull 30.1-М 32.1 + 0.26 mm (31.7 mm)-33.1 mm, of
females 29.1-М 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, М. (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,
underfur almost white, tail dark brown; throat patch very variable,
sometimes completely undefined.
Condylobasal length of male skull (5) 81.2-М 83.2-84.6 mm,
of females (4) 77.7-М77.9-78.0 mm; zygomatic width of males 49.9—
М 51.4-52.0 mm, of females 46.5-М 47.0-47.4 mm; height of male
skull 30.5-М 31.6-32.5 mm, of females 29.8—М 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) М. (M.) f.
mediterranea Barrett-Hamilton, 1898—Pyrenees [Iberian] Penin-
sula; 2) М. (M.) f. bosniaca Brass, 1911—Yugoslavia; 3) М. (M.)
Е milleri Festa, 1914—Island of Rhodes; 4) М. (M.) f. bunites
Bate, 1906—Crete; 5) М. (M.) Ё syriaca Nehring, 1902—Syria; 6)
М. (M.) f. toufoeus Hodgson, 1842—Tibet, Lhasa; 7) М. (M.) /.
kozlovi Ognev, 1931—Kam, eastern Tibet. The last two forms ap-
parently are synonyms and possibly belong to intermedia (У.Н.).
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
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permanent and is uneven. These are unforested or almost unforested
mountains, ravines, rock slides and breccias with, characteristi-
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 т. 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 рег 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.
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897
Sparrows 34.1 Sunflower seed 11.8
Lizards 7.0 Common cherry fruit 815
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 оссиггепсез— 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 0 to 75%; in summer from 20 to
75%, in autumn from 11 to 33%, in winter from 0 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%).
The occurrence of birds in the marten diet varied in summer
from 0 to 33%, in autumn—from 0 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 0 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 0 to 70%,
in autumn—from 9.17 to 100%, and in winter—from 0 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 65.5 Grapes 10.8
Birds 5.1 Dogwood 16.1
Reptiles 355 Myrobalan plum 3.5
Insects 84.0 Blackthorn 4.3
Acorns 2.4 Pear 315
Blackberry 23:3 Nuts 1.0
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 (М.Г. 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
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899
Кое 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
(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) ог 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
602
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 т, and up to 1700-2400 т, 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
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.
602
901
Я.
Fig. 221. Stone marten, Martes (Martes) рота Егх1, 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
603
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 оп
the roofs of houses. It was observed early in the morning and on
moonlit nights. The period of pregnancy is also the same as in pine
marten, i.e. 236-237 days (Manteifel’, 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 1961 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.
603
604
903
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-
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.).
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905
Subgenus of Himalayan Martens, ог Kharza
Subgenus Charronia Gray, 1865*°
KHARZA“
Martes (Charronia) flavigula Boddaert, 1785
1785. Mustela flavigula. Boddaert. Elench. Anim. p. 88. Nepal.
1811. Mustela aterrima. Pallas. Zoographia Roso-asiatica, 1, p.
81. Between the Ud and Amur rivers.
1862. Mustela (Martes) flavigula var. borealis. Radde. Reisen Siid.
Ost. Sibirien, 1, p. 19, 24. Bureinsk mountains (V.H.).
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.
606
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
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
608
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-М612-719 mm, of
females (12) is 500-М575-620 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-М116.5-136 mm, of females (9) 100-
М112.0-123 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-М100.6-109.4 mm; zygomatic width of
males (5) 54.4-M60.7-67.9 mm, of females (5) 53.7-М57.2-59.4
mm; skull height of males (7) 38.0-М42.8—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-M3317-5748 gm, of females (10 specimens between August
and February) 1155-М2765-3827 gm (Bromlei, 1956) (V.H).
909
ANS
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in Ss
A И А
chs ЗАМ , =
| И: \ i | я №:
A м
t ) a ) \ “AWS TEN
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607 Fig. 224. Skull of kharza, Martes (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
609
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)
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 [r.] (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),
4SRange 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 (У.Н.).
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 (Charronia) flavigula
Bodd. in the USSR. V.G. Heptner.
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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. Нершег.
61
—
914
It is distinguished from the nominal form Бу 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.
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, koreana) 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) М. (Ch.) Е flavigula Г. 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) М. (Ch.) Е chrisospila Swinhoe, 1866—Taiwan;
4) М. (Ch.) f. peninsularis Bonhote, 1901—southern Tenasserim
and Malacca; 5) М. (Ch.) f. indochinensis Kloss, 1916—Indochinese
Peninsula, northern Tenasserim; 6) М. (Ch.) Ё 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
612
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
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, 45.5 63.1 44.9 31.5 42.3
goral, moose calves,
wapiti, wild pig (suckling)
Small exploited species 27.1 29:7 26.6 20.0 32.2
(squirrel, hare, sable,
Siberian weasel)
Mouse-like rodents, pikas 522 — 6.1 11.6 3.4
Hazelhen 12.6 Why 18.3 17.6 6.8
Small birds 3.4 — 4.1 787 Пот
Fish (migratory salmon ) 21 — — — 8.5
Molluscs 1.4 -— — 5.9 —
Insects 0.5 — — 1.9 —
Nuts and fruits 2.2 — — 3.8 5.1
613
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.
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 home 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 Ритог’е). 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.
615
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.
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
616
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:
ем = 33.
Shunde entre: Bo hd PP
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
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 т. 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.
618
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.
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
(С. 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).
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1758.
1780.
1780.
1792.
1816.
1820.
1829.
1829.
1918.
1918.
1922.
1948.
925
WOLVERINE [ROSOMAKHA]*
Gulo gulo (Linnaeus, 1758)*
Mustela gulo. Linnaeus. Syst. Nat., ed. X, 1, p. 45. Northern
Scandinavia (Lapland).
(Ursus) luscus. Linnaeus. Syst. Nat., ed. X. 1, p. 47. North
America, Hudson Bay.
Gulo sibiricus. Pallas. Spicil. Zool. 14, p. 35, tab. 2. Upper
Ob’ river. Requires confirmation—Altai.
Ursus gulo albus. Kerr. Anim. Kingd. Syst. Cat. No. 381,
190. Kamchatka.
Gulo vulgaris. Oken. Lehrb. Naturg. 3, 2, p. 1004. Renam-
ing of Gulo.
Gulo borealis. Nilsson. Skand. Fauna. Dagg. Djur. 1, p. 95.
Renaming of Gulo.
Gulo arcticus. Desmarest. Mammalogie, 174. Renaming of
Gulo.
Gulo arctos. Kaup. Entw. Gesch. Nat. Syst. Europ. Thierw.
1, p. 68. Renaming of Gulo.
Gulo biedermanni. Matschie. Sitz-Ber. Ges. naturf. Freunde.
Berlin, p. 147. Mountains south of Telets Lake, Altai.
Gulo wachei. Matschie. Ibidem, p. 147. Sources of Katun’
river north of Belukha, Altai.
Gulo kamtschaticus. Dybowski. Arch. Tow. Nauk. Lwow, 1,
p. 349. Nomen nudum.
Gulo gulo kamtschaticus. Averin. Trudy Kronotsk. gos.
zapovednika, 1, р. 145. Nec 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.
619
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.
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.
АА
№ Ni 1
Fig. 230. Wolverine, Gulo вшо L. Sketch by А.М. Komarov.
620
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.
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 по
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.
621
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.
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)
1$ 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[oological]
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.
98
Systematic Position
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623
932
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
625
933
transgressions; however, а 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 72°40° N. lat. mouth of the Khur, left tributary of the
Pyasina (73° N. lat.); Novaya and Bol’shaya Balakhnya rivers (about
73°20° М. lat.); (Yakovlev, 1930) and Taimyr Lake at 74° М. 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
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.
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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 Chana
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.
626
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.
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 Radomyshl’ (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’ М. 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
1Places 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.
627
938
Tara (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
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° М. lat.) and at approximately 60° М. 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 Chana (Chanov,
Kuibyshev and Vengerovsk regions of Novosibirsk district, 1948—
1951), to Chagly Lake north of Kokchetav! (1949; Sludskii, 1953,
1953a), to Borovom southeast of Kokchetav (Mikhel’, 1934), to
Zlatoust оп 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.
628
940
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’ М. lat.) and the PriKosogol’
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
629
941
reverse, and in the whole intervening extent is а 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.
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 (Anady1
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 (Kurtén and Rausch, 1959) that
this Eurasian species is represented by one form only, G. g. luscus
Linnaeus, 1758.
630
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-
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, а 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
631
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.
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 а 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
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 а 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
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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
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
635
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 (6’ 0)—83 gm (9 9) with
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.
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-уеаг 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
636
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.).
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.
637
954
АП 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. vierfiiss. Thiere, p. 11.
1817. Putorius. Cuvier. Régne anim., 1, p. 147. Mustela putorius
Linnaeus.
1829. Arctogale. Kaup. Entw. Gesch. Nat. Syst. Europ. Thierw. 1,
p. 30. Mustela erminea Linnaeus.
1829. Ictis. Каир. Ibidem. 1, р. 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, р. 234. Mustela
vulgaris Erxleben = Mustela nivalis Linnaeus.
1841. Lutreola. Wagner. Ibidem. 2, p. 239. Viverra lutreola Linnaeus.
1871. Mustelina Bogdanov. Birds and animals of the chernozem
belt of Povolozh’e (Tr. Obshch. estestvoisp. pri. Imp. Kazansk.
Universit., 1, section 1, р. 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
1$ correct).
955
1871. Hydromustela. Bogdanov. Ibidem, р. 167. Mustela lutreola
Linnaeus.
1899. Eumustela. Acloque. Faune de France, Mamm., р. 62. Mustela
vulgaris Erxleben (= Mustela nivalis Linnaeus) et Mustela
erminea Linnaeus.
1911. 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
638
956
of species, several of them have evident differences in skull struc-
ture, though in close species, they are not great (see later).
Sanh Я
Dental formula I 3 С i P 3 M ar 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
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.
640
958
(40-75 gm the smallest species of the carnivore order), and larg-
est species (М. 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
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).
EX ~ <
Arar р;
АЙ
ey, У
641
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,
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 Бе 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
?Ап 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.
642
962
or steppe polecat, М. 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. (M.) nivalis Linnaeus, 1758; 2) ermine, M. (M.)
erminea Linnaeus, 1758; 3) solongoi М. (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.).
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
3(2).
4(1).
5(8).
6(7).
7 (6).
skull is up to 53 mm, of female—up to 46 mm ................
PONE Meh eet rites ermine, M. (M.) erminea (page 995).
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).
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.
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.
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 .............
И black polecat, М. (P.) putorius (page 1107).
Color of upper body light with yellow color predominant,
because light underfur is weakly covered by dark-brown
guard hairs; venter light, yellowish. Only distal half of tail
black; basal part has light color of back. Black leg color
sharply contrasting with color of trunk. Postorbital part
of braincase with well-defined isthmus, and its edges
form angled lines, corners directed towards each other
. steppe (white) polecat, M. (P.) eversmanni (page
1133).
3An exception is present in some races of solongoi [M. altaica]; see 14 (13) and
text.
643
964
8(5).
9112):
10 (11).
11 (10).
12 (9).
13 (14).
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).
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.
Skull moderately elongated—distance between mastoid proc-
esses constitutes about half of condylobasal length of skull.
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 ...........иниииииниинниниии
... Russian (European) mink, М. (M.) lutreola (page 1078).
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
ОО American mink, М. (M.) vison (page 1391).
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.
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) ...............:sssceeeeeceeeeeeees
sata eae ke cat Siberian weasel, M. (M.) sibirica (Page 1051).
644
643
965
14 (13). Dark “mask” оп muzzle absent. Postorbital constriction
1766.
1777.
1800.
1811.
1877.
1896.
1899.
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)
А solongoi М. (М.) altaica (page 1030) (V.H.).
Subgenus of Weasels
Subgenus Mustela Linnaeus, 1758
WEASEL!
Mustela (Mustela) nivalis Linnaeus, 1766
Mustela nivalis Linnaeus. Syst. Nat., ed. XII, 1, p. 69.
Westerbotten province, Sweden’.
Mustela vulgaris Erxleben. Syst. Regni Anim., 1, p. 471.
Umerennaya Evropa [Temperate Europe]. After Ognev
(1935)—Vic. Leipzig.
Mustela boccamela. Bechstein. Pennant. Ubers vierf. Thiere,
2, р. 395. Sardinia.
Mustela gale. Pallas. Zoogr. Rosso-Asiat. 3, p. 94. Renam-
ing of vulgaris.
Mustela stoliczkana. Blanford. J. Asiat. Soc. Bengal, 46, 2,
p. 260. Yarkand.
Mustela rixosa Bangs. Proc. Biol. Soc. Washington, 10, p.
21. Saskachewan, Canada.
Mustela nikolskii. Semenov. Zap. Akad. nauk, ser. 8, vol. 6,
р. 14. Vic. Simferopol, Crimea’.
'The correct spelled name for this species is undoubtedly, /azka, and is derived from
the verb /azat’ [=ю 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° М. 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.
1900.
1901.
1903.
1907.
1911.
1921.
1922.
1926.
1927.
1936.
1936.
195i:
1952.
1953:
Putorius nivalis pallidus. Barrett-Hamilton. Ann. Mag. Nat.
Hist. 5, p. 48. Kokand, Fergana.
Putorius nivalis caucasicus. Barrett-Hamilton. Ibidem, p. 48.
Caucasus, Gochal (?) mountains, 12000’.
Mustela (Ictis) dombrowskii. Matschie. Sitzungsber. Ges.
naturf.-Freunde Berlin, p. 231. Siul’nita. Romania.
Putorius (Arctogale) pygmaeus. J. Allen. Bull. Amer. Mus.
Nat. Hist., 19, p. 176. Gizhiga, northern shore of Okhotsk
Sea.
Putorius nivalis dinniki. Satunin. Izv. Kavkazsk. muzeya, 3,
p. 105. Stavropol (Caucaskii).
Ictis boccamela nikolskii. Satunin. Izv. Kavkazskovo muzeya,
5, р. 257. Crimea’.
Mustela rixosa namiyei. Kuroda. Journ. Mamm., 2, p. 209,
Aomori, northern part of Honshu (Hondo) island, Japan.
Ictis nivalis var. kamtschatica. Dybowski. Arch. Tov. Nauk.
Lwow, 1, р. 349. Nomen nudum.
Mustela punctata. Domaniewski. Ann. Mus. Zool. Polonici.
N.H. 5, p. 55. Darasun; southern Trans-Baikaliya.
Mustela nivalis mosanensis Mori. J. Chosen. N.H. Soc., 5,
p. 28. Iengan near Mosan, Korean Peninsula.
Mustela pygmaea yesoidsuna. Kishida. Dobuts Zasshi, 48,
4, p. 177. Hokkaido, Japan.
Mustela pygmaea karaftensis. Kishida. Ibidem, р. 177.
Sakhalin.
Mustela trettaui. Kleinschmidt. Falco, 33, p. 11. “Germany”.
Mustela nivalis kerulenica. Bannikov. Byull. Mosk. Obsch.
Isp. Prirody, 57, No. 2, p. 40. Vic. Undurkhan on Kerulen
[r.], Mongolian Republic.
Mustela nivalis heptneri. Morosova-Turova. 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.
645
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.).
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.
645
646
968
Fig. 234. Weasel, Mustela (Mustela) nivalis L. Sketch by А.М. 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
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 а 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
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 1$ 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
647
a7"
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.
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ое SSeS А
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Fig. 235. Skull of weasel, Mustela (Mustela) nivalis L.
972
Weight of males is 36-250 gm, of females, 29.5-117 тб
(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.
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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.
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 (М. 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
652
976
existence of а 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
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 (Ellerman 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.
‘1эщ34эн “OA “Т 5Ираам (Djaisnp) vjaisny ‘Jaseam ay} JO adues $912э4$ [55 “BIg 159
653
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-М160-172 mm; tail length,
12-М22.6-28 mm; hind foot length, 16-М21-25 mm.
Condylobasal length of male skull, 22.8-M32.7—34.8 mm; mas-
toid width, 11.0-М15.2-16.8 mm; interorbital width, 5.0-M7.2-—
8.2 mm; postorbital width, 7.0-М8.0-8.5 mm; zygomatic width,
13.8—М16.5-19.8 mm.
Weight of males (14), 37-М52.6-68 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
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-М166.6-185 mm, tail length
(11),* М20.2-28 mm; length of hind foot, 12-M18.2—23 mm}.
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-М14.3-16.0 mm; interorbital width of males, 6.6-М7.2-8.5
mm, of females, 5.5-М6.3-7.5 mm; postorbital width of males,
6.5-М7.4—8.0 mm, of females, 6.2-M6.9-8.0 mm.
Weight of males (9), 36-М48.6-84 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 sex of which is not shown: 30-М46.8-84 gm.
Weight of female with 11 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.
654
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.
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-М226.5-260 mm; tail length,
55-М72.3-85 mm; length of hind foot, 22-M39.5—42 mm.
Condylobasal length of male skull, 41.0-М42.1-46.2 mm; of
females (6), 29.8-M31.2—36.0 mm; mastoid width of males, 20.0-—
М20.2-22.4 mm; of females, 14.0-М15.1-16.8 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-М8.8-8.4 mm; of females, 6.0-
М7.1-8.0 mm; zygomatic width of males, 21.5-М22.6-24.0 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,
655
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-М234.1-242 mm; tail length,
55—M68.2-87 mm; length of hind foot, 27-МЗ3.2-35 mm.
Condylobasal length of male skull, 40.5-М43.1-45.5 mm;
mastoid width, 21.0-М21.3-22.0 mm; interorbital width, 9.8—
М10.3-10.8 mm; postorbital width, 7.8-М8.3-9.0 mm; zygomatic
width, 23.4-М24.4-26.0 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.
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 B 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-М186-213 mm; of females
(9), 145-М165-185 mm; tail length of males, 28-М40.5-66 mm;
of females, 20-M33.6—42 mm; length of hind foot of males, 22—
М27.3-30 mm; of females, 19-М22.1-25 mm.
Condylobasal length of male skull (13), 30.5-М35.3-40.2 mm;
of females (9), 29.0-М 32.0-36.8 mm; mastoid width of males,
15.0-М17.3-19.6 mm; of females, 12.8-М15.0-17.5 mm; interor-
bital width of males, 6.0-М7.7-9.5 mm; of females, 5.0-М6.9-7.0
mm; postorbital width of males, 5.0-М8.2-9.3 mm; of females,
5.3-M7.1-9.0 mm; zygomatic width of males, 11.0-М17.9-22.0
mm; of females, 15.3-М16.2-18.5 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.
656
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 М. (M.) п. nivalis. Тай
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-М212.6-243 mm; of females
(6), 155-М181.3-212 mm; tail length of males, 45-М57.9-75.5
mm; of females, 45-М54.1-64 mm; length of hind foot of males,
21-МЗ0.8—38 mm; of females, 29-М25.0-31 mm.
Condylobasal length of skull of males, 31.5-М38.8—43.0 mm,
of females (6), 25.8-M33.2—36.0 mm; mastoid width of males,
13.5-М18.4—20.5 mm; of females, 14.0-М15.1-16.8 mm; interor-
bital width of males, 6.5-М8.5-10.2 mm; of females, 6.2-М
7.1-7.5 mm; postorbital width of males, 5.0-М8.4-10.2 mm; of
females, 6.0-М7.1-8.0 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.)
п. pygmaea as identical to the Middle European “minuta” and an
extreme variant (“minus-variant”) of M. (M.) п. 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-М192-215 mm, tail length,
37-М47-60 mm; length of hind foot, 25-М27-31 mm.
Condylobasal length of the male skull, 31.8-M35.4—40.0 mm;
mastoid width, 15.0-М16.6-19.0 mm; interorbital width, 6.5—M7.7—
8.4 mm; postorbital width, 6.5-M7.5-8.5 mm; zygomatic width,
18.0-М19.7-22.0 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.
657
985
Ognev, 1935; Novikov, 1956; Stroganov, 1962). The question of
weasels in the European part of the country and Caucasus is par-
ticularly complicated.
* * *
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) М. (М.) п. numidica Pucheran,
1855—Morocco, Algeria, Malta, Azores Islands? Corsica; 3) M.
(М.) п. stoliczkana Blanford, 1877—Kashgaria’®; 4) М. (M.) п.
siberica Barrett-Hamilton, 1900—Pyrenees [Iberian] Peninsula and
Balearic Islands; 5) M. (M.) n. galinthias Bate, 1906—Crete; 6) M.
(М.) п. russeliana Thomas, 1911—-Sichuan, southern China; 7) М.
(М.) п. namiyei Kuroda, 1921—Japanese Islands; 8) М. (M.) п.
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.)
п. 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.
658
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
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, 0.7-12.8 9.7 0.2-8.6 0.9
Kalinin district
Zhiguli 9.7-122.8 36.2 0-13.3 2-1
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structures within а 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.
659
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,
[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
659
660
989
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
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 “Tep”., 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 agama is between the teeth of the
weasel. 26 April 1963. Photograph by Yu.K. Gorelov.
661
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-
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
662
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’év 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
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 (Р.В. 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 (Р.В. Yurgenson). In the Mongo-
lian Republic, between 1944-45, there was noted a nine-fold т-
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 па) 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
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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 А.М. Formozoy, nat. size.
0
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 А.М. 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, р. 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.). |
664
996
1895.
1896.
1912.
19133
1914.
1922:
1922.
1923.
1928.
1928.
1929.
193.
1932:
1935.
1936.
1938.
Putorius ermineus ferghanae. Thomas. Ann. Mag. Nat. Hist.,
15, p. 452. Kara-Karyk mountains, Fergana. (?V.G.).
Putorius arcticus. Merriam. North Amer. Fauna, 11, p. 15.
Point Barrow, Alaska.
Mustela lymani. Hollister. Smiths. Misc. Coll., 60, 14, p. 5.
Tapucha*, on the Chuisk Trail, southern Altai.
Mustela nippon. Cabrera. Biol. Soc. Espan., 13, p. 392.
Sinano, on Hondo (Honshu) Island, Japan.
Mustela kanei. G. Allen. Proc. New Engl. Zool. Club, 5, p.
58. Nizhne-Kolymsk.
Putorius erminea var. kamstschatica. Dybowski. Arch. Tow.
Nauk. Lwow, 1, p. 349. Nom. nudum.
Putorius. erminea var. sibirica. Dybowski. Ibidem, p. 349.
Nom. nudum. Nec Pallas, 1773.
Arctogale erminea tobolica. Ognev. Biologich. izvestiya, I,
р. 112. Tara, Former Tobol’sk governance.
Mustela erminea transbaikalica. Ognev. Memuary zool. otd.
Obshch. lyubit. estestvozn., antrop. i etnogr., 2. Sosnovka,
Barguzinsk Preserve.
Mustela erminea orientalis. Ognev. Ibidem, р. 15.
Pokhodskoe $. [village] on the Kolyma [river], 69°04’ М.
lat., 160°55° Е. long.
Mustela erminea baturini. Ognev. Izv. Tikhookeanskoi
nauchno-prom. stantsii, 2, No. 5, p. 9. Great Shantar Island
in Okhotsk Sea, lower Anaur river.
Mustela erminea birulai. Martino. Ezhegodn. Zool. muzeya
АМ SSSR, 31, 1930, р. 208. Aktyubinsk. Nomen
praeoccupatum—Kolonokus alpinus birulai Ogn., 1928.
Mustela erminea ognevi. Jurgenson. Zool. Anz., 98, p. 11.
Mouth of Taz river, western Siberia.
Mustela erminea schnitnikovi. Ognev. Zveri SSSR 1
prilezhashchikh stran, 3, p. 37. Former Kopal’sk Co.,
Semirechensk governance.
Mustela erminea karaginensis. Jurgenson. Byull. Mosk.
Obshch. ispyt. prirody, 45, No. 3, p. 240. Karaginsk Island
near eastern shore of Kamchatka.
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. Sc., 23,
No. 37, p. 559. Kamchatka.
1951. Mustela erminea martinoi. Ellerman 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,
р. 123. Chegem river gorge, northern slope of Great Cauca-
sus (У.Н.).
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 M11.3;
guard hairs in category III in winter to 13 mm (M11.9), in summer
M10.5; guard hairs in category IV to 12 mm (M11.5), in summer
M9.9; length of underfur in winter M9.1 mm, in summer 8.1. The
total number of hairs in 0.25 ст? т winter comprises 4808, of
665
998
which two* are directional hairs while underfur hairs number
4,637—1 for every 27 underhairs; in summer, оп 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.
667
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
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
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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-М2.47-2.7 gm; that
of females (70) is 1.3-M1.54-1.9 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—
M48.39-51.8 mm, that of females is 40.3-М43.39-45.8 mm (У.
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.
668
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.
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-М25.5-
27.7 mm; weight is 0.033-МО.041-0.048 gm; in immature (young
and subadult) its length is 22.2-М23.5-24.3 mm, weight is 0.016-
МО.025-0.32 gm (У. 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, 37.0-47.6 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.).
'8According 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).
670
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
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 Цигир (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.
‘1э34эН `9`Л “USSN э41 ш ‘1 раипииа (2]2151И/) DJaIsny ‘эп JO UONNQINSIP JO з1эрзоЯ ‘фус “314 699
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-
зегуе!?. 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.
671
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 [Lake] 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 Alakol’ 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.
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.
tn 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, II’ 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,
п 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.
673
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. n.*
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
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) (У.Н.).
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.
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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. Оп
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-М248.7-278 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-М46.2—55
mm.
Condylobasal length of skull of males (76), 41.0-М46.8—52
mm, of females (20), 39.8—М42.0-45.2 mm; zygomatic width of
males, 21.5-М26.0-27.8 mm, of females, 21.2—-M22.5—25.3 mm;
interorbital width of males, :10.0-М12.0-13.0 mm, of females,
M10.2 mm*; mastoid width of male skull, 19.0-М22.9-26.2 mm,
of females, 18.0-М21.1-24.5 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.
674
1011
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-М57.7-58.1% of the condylobasal length of its skull.
Body length of males (43), 200-М260-300 mm, of females,
(35) is 200-М230-270 mm, tail length”? of males is 50-М90-120
mm, of females 50-М70-100 mm; length of hind foot of males
40-М45-48 mm, of females, 34-М37-42 mm; height of ear of
males 18—M20-—22 mm; of females, 14-М16-18 mm.
Condylobasal length of male skull (45), 43.4-М48.3-50.1 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**-М12.0-13.2 mm, of females 9.2-M10.8-
12.0 mm, postorbital width of males, 9.6-М11.2-12.1 mm, of
females, 8.2-М9.2—11.1 mm; mastoid width of males, 21.2—М23.6-
25.2 mm, of females, 19.3-М21.0-22.2 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.
675
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-М258-295 mm, of females,
185-М220-260 mm; tail length of males, 45-M75-105 mm, of
females, 40-М60-90 mm; length of hind foot of males, 37=М44—
48 mm, of females, 32-M36—42 mm.
Condylobasal length of male skull (14), 44.0-М48.4—49.4 mm,
of females, 41.2-M43.7-45.2 mm; zygomatic width of males, 24.4—
М27.3-28.7 mm, of females, 20.8-M24.4—26.2 mm; interorbital
width of males, 11.8-М12.2-12.6 mm, of females, 9.9-М11.3-
12.0 mm; postorbital width of males, 10.0-М11.4-12.8 mm, of
females; 9.0-М10.2-11.7 mm; mastoid width of males, 21.3-
M23.6—25.3 mm, of females, 19.2-М21.1-23.0 mm.
Weight of males (5), 145-М191-247 gm, of females (4), 106—
149 gm (Altai preserve, winter; material from ММО).
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-М260-325 mm, of females
(7), 176-М212-222 mm; tail length of males, 70-М86-100 mm,
1013
of females, 67-М72-77 mm; length of hind foot of males, 40—
M43-48 mm; of females, 33-M38-—43 mm; height of ear in males,
20-М21-22 mm, of females, 18-M19-20 mm.
Condylobasal length of male skull (24), 44.0-M46.0-49.2 mm,
of females (23), 38.9-M40.9—44.3 mm; zygomatic width of males,
24.0-M26.2—28.8 mm, of females, 20.0-М22.3-25.2 mm; inter-
orbital width of males, 10.1-М11.4-13.2 mm, of females, 8.8—
М9.5-10.0 mm; postorbital width of males, 10.0-М11.0-12.5 mm,
of females, 8.7-M9.7-10.9 mm; mastoid width of males, 20.1-
М22.5-24.7 mm, of females, 18.0-М19.2-20.2 mm.
Weight of male (8), 102-M166—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-М42.7-43.7 mm;
zygomatic width, 21.6-М23.0-23.7 mm; interorbital width, 9.8—
M10.1—11.0 mm; mastoid width, 19.3-М20.3-21.6 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-М85.7-—
90 mm, length of hind foot, 31-М40-45 mm.
676
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,
21.0-М23.2-24.5 mm, of females, 18.2-М19.5-23.7 mm; inter-
orbital width of males, 9.1-M10.8—11.6 mm, of females, 7.9-M8.7—
10.0 mm.
Postorbital width of males, 9.3-М11.0-12.0 mm, of females,
8.3-М10.1-10.7 mm; mastoid width of males, 19.1-М21.0-22.1
mm, of females, 16.8—М18.1-21.8 mm.
Weight of male (9), 101-М134-174 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-М259-300 mm; tail length 70-
M75-82 mm; length of hind foot, 40-М41-42 mm.
Condylobasal length of male skulls (9), 40-М43-45 mm;
zygomatic width, 22.0-М23.3-25.5 шт; interorbital width, 9.5-—
M10.4—-11.0 mm; mastoid width, 19.0-М20.6-22.0 mm.
Weight of males in summer (12), 121-М145-211 gm, of
females (10), 60-М72.5-86 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.
677
1015
It is a well differentiated form, to which the ermine of the
Gobi Altai and adjacent parts of the Mongolian Republic (mongolica,
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: 1) M. (M.) e. hibernica. Thomas et
Barrett-Hamilton, 1895—Ireland; 2) М. (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) М. (M.) e. ricinae Miller, 1907—islands of the
Hebrides; 5) M. (M.) e. minima Cavazza, 1912—Switzerland; 6)
М. (M.) e. nippon Cabrera, 1913—Japan; 7) М. (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) М. (M.) e. semplei
Sutton et Hamilton, 1932—Baffin Land and adjacent part of main-
land; 11) M. (M.) 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) М.
(M.) е. bangsi Hall, 1944—region west of the Great Lakes; 14) М.
(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.
(M.) 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) М. (М.) e. alascensis Merriam, 1896; 23) М. (M.) e. initis Hall,
1944; 24) М. (M.) e. salva Hall, 1944; 25) М. (M.) e. celenda Hall,
1944; 26) М. (M.) е. seclusa Hall, 1944; 27) М. (M.) е. 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’) Бу 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.
678
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
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
680
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.
agrestis] predominate; in forest areas, various species of red-backed
voles; in tundras, lemmings also. Ermine rarely attack the commercial
*Local word?—Sci. Ed.
1019
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1020
animals and birds—even more rarely with success. However, cases
of ermine feeding on muskrats, 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.
Table 64. Nutrition of ermine in the winter period in Pechora-Ilych preserve over
a 12-year period (% of occurrence)
Type of food Range of fluctuation Average for
in occurrence 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—3.05 1.358
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
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, ес. 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.-1 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 рег
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.
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 рег 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.
683
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.
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 compet.tors. 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
684
1026
уеагз— гот 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
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
685
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 т
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).
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 рег 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.
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.
686
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 т.
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
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. Mémoires 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 longstaffi. 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. 1 etnogr., 2, р. 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.
687
1031
Diagnosis
Color from bright ocher-reddish to straw-ocher or from nut-brown-
ish to brown. Lighter below than above, transition from dorsal
color to ventral color lower part gradual or border sharp. Tail
monotone, without black terminal half. Dark coffee-brown field
(“тазК”) 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
688
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
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 а 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 1.1. Sokolov, А.М. Formozov and А.Р. 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.
689
1034
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Fig. 247. Skull of solongoi Mustela (Mustela) altaica Pall.
.690
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
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).
691
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.
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 (М. 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
693
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 western 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 Ili 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-
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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In the literature, very general references to the occurrence of
the solongoi “т 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 /apsus—Sci. Ed.
694
695
693
694
1041
perhaps somewhat beyond it, i.e. to 60° М. 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° Е. 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)”.
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
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
5In the description of the solongoi range east of Baikal, besides the sources
mentioned, unpublished information offered by S. Anaskin (Bur[yat] ASSR), P. Zimin
(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).
Тре 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).
7Range 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.
694 Fig. 249. Species range of the solongoi, Mustela (Mustela) altaica Pall. (V.G. Нершег).
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.
696
1044
50101201 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.
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 тт;
ear height of males is 16-20 mm.
Weight of males (winter) is 217-255 в, of females, 122—135 g
(lower Ili, 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
697
1045
(“birulei”’) is highly probable, but needs confirmation. Оп 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-М241-260 mm, of females (5),
217-М223-227 mm; tail length of males, 110-M129-145 mm, of
females, 104-М108-—113 mm; length of hind foot of males, 41.0-
№М44.1-47.0 mm, of females, 35.5-М37.2-38.4 mm; ear height of
males, 20—M20.8-21.6 mm, of females, 15.7-М16.8-18.2 mm.
Condylobasal length of male skull (7), 51.2-М52-53.0 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-М24.4-25.7 mm; interorbital
width of males, 10.3-М11.3-12.6 mm, of females, 9.3-М 9.6-10.2
mm; mastoid width of males, 22.8-M23.3—24.0 mm; of females,
22.0-М22.4—23.2 mm.
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 М. (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-М224-226 mm; tail length of males, 108-М129-144 mm, of
females, 100-М106-117 mm; length of hind foot of males, 40.8—
М42.3-44.2 mm, of females, 33.2-M36.9—43.8 mm; ear height of
males, 17.4-М18.3-26.6 mm, of females, 16.0-М17.3-20.2 mm.
Condylobasal length of male skull (6), 49.5-М50.5-52.3 mm,
of females (7), 43.0-M44.2—46.2 mm; zygomatic width of males,
25.7-М 26.6-27.3 mm, of females, 20.0-М22.6-24.5 mm; interor-
bital width of males, 9.3-M10.3-11.8 mm, of females, 8.1-М9.2-
10.0 mm; mastoid width of males, 20.2-М22.5-23.8 mm, of
females, 18.7-М19.0-19.4 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
698
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
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 daurica] 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*
Daurian pika 34.2 21:2
Small birds 13.4 0.9
Brandt’s vole [Microtus brandti] 9.3 27.2)
Narrow-skulled vole [M. gregalis] 9.3 36.5
Daurian partridge[Pardix 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] 21 9.1
Fish 1.0 —
*Summer food data after material from southeastern Trans-Baikaliya (Daurian
steppes).
699
1049
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 66.6
Rodents 61.0 66.6
Voles, mice 24.2 DED:
Muskrat 14.7 13.9
Tamarisk jird. [Meriones tamariscinus] 10.5 5:5
Midday gerbil [M. meridianus] 1.0 —
Water vole [Arvicola terrestris] 5.8 87
House mouse [Mus musculus] — ied
Tolai hare [Lepus tolai] — 8.3
Birds 92. В.М
Pheasant [Phasianus colchicus] — 2.1
Moorhen [Gallinula chloropus] — 55
Ducks — Dil
Small birds Dal 2:7
Lizards — 22:5
Frogs — 2:7
Fish 3.1 —
Insects — 8.3
Coagulated blood 32.6 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
ermine.
700
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.
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, пит-
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
Ili 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 а 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,
pl. 61, Fig. 1. Pekin [Beijing], China.
1904. Mustela sibirica miles. Barret-Hamilton. Ann. Mag. Nat.
Hist. 13, p. 391. Dauriya, Trans-Baikaliya.
1911. Mustela manchurica. Brass. Aus dem reiche der Pelze, p.
490. Manchuria. |
70
—
1052
1911. Kolonokus sibiricus australis. Satunin. Izv. Kavkazskoyo
muyzeya, 5, р. 23. Zhiryakovo, Antropovsk. voll[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, а 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.
702
1053
=
i ai м Е
IANA CON =
) ПО 4 Ma WS =
as
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.
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.
ve LADD
№ a al
pb) ay nt
WF
fi NN
mie к
pa Ne
)
|. у
AV И i
| Ny i in р
My, I TNS
il |
i
AT \ Wy
ve NA у
| (т
>
= SS
NN А:
Ay | &
/)
pr. fat we .
ь ле М А
Wo м г № yw hy) Le &
uN, wi) \ Wy И
703 Fig. 251. Skull of the kolonok, Mustela (Mustela) sibirica Pall.
704
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
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
705
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
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
310-430 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.).
Geographic 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
707
1059
below 63° М. 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’ М. 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*?.
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 63° 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.
3
weasel, Mustela (Mustela)
a Pall., in the USSR (V.G. Heptner): 1—long distance transgressions
climatization; 3—place where the Japanese
itatsi Temm., was introduced on Sakhalin.
introduced for ac
sibirica
th in Yakutiya; 2—place where kolonok were
706 Fig. 252. Borders of distribution of the kolonok, Mustela (Mustela) sibiric
to the nor
1061
УПуш at 65° М. 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 mouth 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’ М. lat.; У.Р. 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 Markakol’ 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.
708
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
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 Chana, 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)3? 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).
31This 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 urochishche, meaning “small village’—Sci. Ed.
710
1065
northern montane region—Pri-Kosogol’, Khangai and Kentei; оп
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**.
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.
71
—
1066
cant even by furriers (Kuznetsov, 1941). Several East Asiatic т-
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.
australis, miles).
Measurements comparatively small. Color of winter fur not
vey bright, yellowish-red.
Condylobasal length of male skull (33) is 58.0-М61.7-63.5
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-М12.5-13.2 mm, of females,
10.5-М11.0-12.2 mm; mastoid width of males is 26.8-M27.5—
28.7 mm, of females, 23.0-М24.3-26.1 mm.33
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.
Condylobasal length of male skull (16) is 63.8—М65.4—67.3
mm, of females, 57.2-M61.2-62.7 mm; zygomatic width of
males 1$ 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-М12.7-13.0 mm; postorbital width of males is 13.2-M13.6—
14.0 mm, of females, 12.0-М12.7-13.3 mm; mastoidal width of
males is 28.2-М29.5-31.0 mm; of females, 24.0-М25.0-27.0 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*‘, М. (M.) 5. 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).
“И 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 по
reason at all to litter the Russian language with a word completely alien to it,
especially a non-declinable one.
712
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] Mfoscow]
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
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:
ОМ. (M.) s. subhemachalana Hodgson, 1837—Himalayas from
Nepal to Bhutan; 2) M. (M.) s. canigula Hodgson, 1942—Tibet; 3)
М. (M.) 5. 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) М. (M.) 5. manchurica Brass, 1911—Manchuria; 8) М.
(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) М. (M.) 5.
азай 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
а 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 т coastal oak forests. It is met with from the sea coast
714
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
Table 67. Geographic variation in foods of kolonok (% of occurrence)
Sos 5 =
Pile MS 22 = =
я Е рев
ве т Ни aga a =
Type of food Nash Op Saale я А fe
Ma INN OLA urs use bi нее Ее
а toy cra а
а jaan tate
И Е
а я BUS ile o
Water vole [Arvicola 36.6 56.1 46.0 1.8 7.0 — —
terrestris |
Small voles and писе 15.0 15.6 15.0 50.1 20.0 24.6 68.7
Altai zokor — — — — 21.0 — —
[М. myospalax]
Great jerboa —- — — — 140 — —
[Allactaga major]
Siberian chipmunk — — — 0.9 — 9.6 —
[Tamias sibiricus]
Pikas — — — 27 — 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 — 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
А.С. 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, О. арта], 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.
714
TAS)
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
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 Лапе-
August, and fish—in September—October (migration of salmonids).
The role of plant food (not above 4.9%) increases in August—
February. Amphibians, reptiles and invertebrates are foods of the
warm 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 0
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
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
716 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.
ТА
718
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,
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.
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 а montane creek in а
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 (Jxodes 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 10. 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.
719
1879
1912.
1912.
ТО.
1912.
1912.
1912.
1932.
1932:
1939.
1939.
1939.
1944.
1079
Lutreola europaea Homeyer. Zoolog. Garten, 20, p. 184.
Renaming of lutreola Linnaeus.
Mustela (Lutreola) lutreola wyborgensis Matschie.
Sitzungsber. Ges. naturforsch. Freunde Berlin. 1912, p. 347.
Vyborg, north of Leningrad.
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.).
Mustela (Lutreola) lutreola budina. Matschie. Ibidem, p.
349. “Ortelsburg, East Prussia” (Shchitno southeast of
Ol’shtyn [Olsztyn], northern Poland; V.H.).
Mustela (Lutreola) lutreola varina. Matschie. Ibidem, p. 351.
Vic. of Shverin, Mecklenburg.
Mustela (Lutreola) lutreola albica. Matschie. Ibidem, p. 351.
Levitts [Lewitz] river—tributary of the Elbe. Mecklenburg.
Mustela (Mustela) lutreola glogeri. Matschie. Ibidem, p. 354.
“Brig [Brzeg], Silesia” (Bzheg, southeast of Vrotslav
[Wroclaw], Poland; V.H.).
Mustela lutreola transsylvanica. Ehik. Allat. Kézlem, 29, р.
142. “Komitat Turoch, Austro-Hungary” (district south of
Beskid, Czechoslovakia—about 150 km southwest of
Krakow; У.Н.).
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.
Mustela (Lutreola) lutreola turovi. Kuznetzov. In: Novikov.
European mink, p. 47. Caucasus.
Lutreola lutreola borealis. Novikov. Ibidem, p. 63. Milet
village, former Bogorodsk Co. Moscow governance. Nomen
praeoccupatum—Mustela flavigula var. borealis Radde,
1862.
Lutreola lutreola caucasica. Novikov. Ibidem, р. 63.
Prokhladnaya Station, northern Caucasus. Nomen ргаеос-
cupatum—Mustela nivalis caucasicus Barrett-Hamilton,
1900.
“L. (utreola) 1. (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.
720
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
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
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720 Fig. 258. Russian mink, Mustela (Mustela) lutreola L. Sketch by А.М. 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.
721
1082
body, and in particularly dark individuals and races, а 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
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
723
1084
edge of the carnassial tooth, but it does not lie in the groove on
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-М150-180 mm;
length of hind foot of males is 54-M59-64 mm, of females, 50-
М52-53 mm; height of ear in males is 20-М23-25 mm. Male tail
constitutes 35-M41-52% of body length, of females, 37-М40-
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-
М13.6-15.4 mm, of females, 11.1-М12.4-14.0 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-М29.4—31.7 mm (Novikov, 1939).
1085
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724
1086
Dimensions of mink (18) from Volga-Kama Territory: body
length of males is 310-M375-—406 mm, of females, 300-М327-
360 mm; tail length of males is M145 mm,* of females, 120-
M128—142 mm; length of hind foot of males is 52-M56.2—61 mm,
of females, 45-М50.0-58 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-МО.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 (“Kolonocus’”) and to polecats
(“Риюопти5”), 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.
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.
*6For 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 (“Риюоти5”) 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
725
1088
represent species parallel to the black (М. putorius) and white (М.
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,
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
Tara 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 Tara, it arches
to the Ural ranges, somewhere approximately at the latitude of
Chelyabinsk. South-southwest of Tara 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 (Kopa settlements; Sludskii, 1953). Infor-
mation concerning the occurrence of mink in Alakol’(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° М. 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.
727
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
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
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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
shown by the above-mentioned transgressions of mink to Kokchetav
and the Irgiz. Beyond the Irtysh (upper Tara), however, the mink
was still absent in 1950 and 1960 (for details, see Novikov, 1939).
*8Range 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 а 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)*.
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.
729
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,
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.).
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, М. (M.) 1. 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-М365-380 ти; tail
length is 124-М137-150 mm; length of hind foot is 54-М56-58
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—
М13.4-15.0 mm, of females, 11.1-М12.8-14.0 mm; zygomatic
width of males 15 32.4-M34.1-36.9 mm, of females, 28.9-МЗ2.1-
37.0 mm; mastoid width of males is 27.0-М29.7-32.2 mm, of
females, 26.5-М28.2—31.7 mm.
*In Russian original, mistakenly given as “mm”, rather than %—Sci. Ed.
730
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.
2. Middle Russian mink, М. (M.) 1. 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-М12.3-13.9 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-M31.2—33.2 mm, of
females, 28.0-М29.2-30.3 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, М. (M.) 1. 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.
731
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-М56.3-59.4 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-М36.8—40.0 mm, of females, 32.0-M33.3-
35.8 mm; mastoid width of males is 29.4-M31.8-34.9 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
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, М. (M.) 1. lutreola and the Caucasian, М. (M.) I. 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, М. (М.) l. novikovi is actually a
transitional population between М. (M.) lI. lutreola and М. (M.) 1.
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, М. (M.) I. cylipena Matschie (syn.
budina, varina, albica, glogeri, hungarica)”.
Dimensions very large, only slightly inferior to Caucasian mink,
М. (M.) 1. turovi. Color quite dark, apparently, corresponding to
color of Middle Russian mink М. (M.) 1. 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-М60.6-63.1 mm, of
females, 53.3-М54.0-54.9 mm; interorbital width of males is 13.3-
М14.2-15.2 mm, of females, 11.8—12.0 mm; zygomatic width of
males is 35.5-М37.3-38.4 mm, of females, 32.4-32.6 mm; mas-
toid width of males is 30.8-МЗ1.9-33.3 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.
732
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, М. (M.) 1. transsylvanica 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.
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, М. (М.) 1. biedermanni Matschie, 1912—France (У.Н.).
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.
733
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 (У. 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
of water—1.0-1.5 т wide and not more than 1 т 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
734
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
surface decreases to 100-150 т 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 т from the water’s edge. It is of simple construc-
tion: 1-2 passages 8-10 cm in diameter and 1.40-1.50 т 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 т. 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
735
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).
Reproduction. In Moscow Zoo, estrus was observed on 22-26
April".
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—3l1st 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
“‘j.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%.
736
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).
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 шт (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.).
*Local word for trap/Inet type—Sci. Ed.
737
1758.
WSS!
1827.
1843.
1851.
1926.
1929.
1932.
1948.
1952:
1955:
1107
Subgenus of Polecats
Subgenus Putorius Cuvier*, 1817
FOREST, OR BLACK, РОГЕСАТ“
Mustela (Putorius) putorius Linnaeus, 1758
Mustela putorius Linnaeus. Syst. Nat. Ed. X, 1, p. 46. Upsala,
Sweden.
Mustela furo. Linnaeus. Syst. Nat. Ed. X, 1, p. 46. Africa,
Morocco.
Putorius vulgaris. Griffith, Cuvier’s Animal Kingd., 5, p.
120. Substitute for putorius Linnaeus.
Putorius foetidus Gray. List Spec. Mamm. Brit. Mus. Sub-
stitute for putorius Linnaeus.
Putorius verus. Brandt. In: Semashko. Russian Fauna, 2, p.
357. Substitute for Mustela putorius Linnaeus.
Putorius putorius stantschinskii. Melander. Nauch. izvest.
Smolensk. gos. univers., p. 137. Roslavl’sk co., Smolensk
gov. Described as a color “variety”.
Putorius putorius orientalis. Brauner. Ukrainsk. mislivets
ta. Ribalka, No. 2-3, р. 9, Ukraina. Exact type locality not
indicated, but apparently, northern part of former Khersonsk
gov. Nom. praeocc.—Mustela erminea orientalis Ognev,
1928.
Mustela putorius rothschildi. Pocock. Scot. Nat. Edinb.
Dobrudza River, Romania.
Putorius putorius L. f. piriformis. Kostron. Prace Mor.-slez.
Akad. prir. ved., 20, p. 52. Czechoslovakia.
Putorius putorius ognevi. Kratochvil. Sborn. Vysoke skoly
zemed. a lesnecke v Brne, No. 1, p. 8. Middle European
part of USSR. Nomen praeoccupatum—Mustela erminea
ognevi Jurgenson.
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. Мотшеп praeoccupatum—Mustela erminea
orientalis Ognev, 1928; Putorius putorius orientalis Brauner,
1929.
1965. Mustela (Putorius) putorius mosquensis. Heptner. Zool. Anz.,
176, H.1, р. 2. Заууто 20 km east of Moscow (У.Н.).
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 сп? on back 8,500-
9,000; ratio, one guard hair to 19-20 underhairs, and on abdomen
average number of hairs per 1 сп? 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.
738
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.
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.
739
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
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
ИИ
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. р. 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.
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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
741
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
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.
Carnassial 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.
742
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
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) 15 33.5=МЗ6.5-38.2 mm, of young
(17), 31.0-M34.0-37.0 mm; weight of penis bone is 0.280-МО0.337-
0.470 gm, of young, 0.080-МО.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.
“7After 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.
743
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.
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 “М. 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.
**In English, “ferret”’—Sci. Ed.
7
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 (У.Н.).
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’ М. 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’,
744
745
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
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 647 М. lat. Thence,
the northern border of the range goes on, to the upper Vychegda
(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 Tara 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
119
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 Поп“. 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° М. 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; Korneev, 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.
746
1121
The range of black polecat to the south of the line Zhitomir-
Kiev-Orel-Voronezh-Gor’ kii-Kirov-Ufa-southern Urals (approxi-
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 по 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° М. 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.).
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.
748
1124
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
749
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
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. stantschinskii” 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, М. (Р.). р. 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) М. (P.) р. aureola Barrett-Hamilton, 1904—
Spain; 3) M. (P.) p. rothschildi Pocock, 1936—Dobruja, Romania;
4) М. (P.) р. angliae Pocock, 1936—England; 5) М. (P.) р.
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
751
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).
Table 68. Geographic variation in foods of the forest polecat (in percentage of
occurrence)
Type of food Volga-Kama Territory (Grigor’ev Middle zone
and Teplov, 1939) (Lavrov, 1935)
Summer Winter 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 111
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 Cy 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 dimensians 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
752
1130
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
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.
753
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 (Manteifel’, 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.
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.
754
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 evérsmanni Lesson var. michnoi. Kastschenko.
Kashchenko. Ezhegodn. Zool. muzeya Ak. Nauk, 15, p. 271.
Kiran river 20 km from Troitskosavsk (Kyakhta), Trans-
Baikaliya.
1912. Putorius eversmanni sibiricus Kastschenko. Kashchenko.
Ezhegodn. Zoolog. muzeya Ak. Nauk, 17, p. 395. Aginsk
steppe, Trans-Baikaliya. Nomen nudum. Nec 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, р. 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,
р. 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 Chana.
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.** Ili 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.
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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.).
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.
“SS
‘ INN
А"
ANY
~
Fig. 270. White, or steppe, polecat, Mustela (Putorius) eversmanni Less. Sketch by
A.N. Komarov.
756
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
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, summer 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.
ПЭ
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
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
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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).
760
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 (У.
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).
One giant polecat had a body length without tail of about
75-80 cm, although normal dimensions of animals from this ге-
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 а 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.)
eversmanni>>.
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).
76
—
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
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
762
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).
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.).
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 т 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
764
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,
1931)°°.
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
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 (58°40° М. lat. and a little east of 52° Е. 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 published 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.
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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 Tara (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).
765
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-
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, 5.
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
57Ро]еса{$ were found at some points on the Ob’ between the mouth of Tym
(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.
1451
the frontier (Favorskii, 1936; Р. 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
766
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
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 both 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 Khodzheil’ 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).
767
1153
Farther, polecats exist, though sporadically distributed арраг-
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
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), М.М. Bogdanov 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.
768
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— т Austria where it rapidly spread
and its number strongly increased (Bauer, 1952; Kratochvil, 1962;
О. Wettstein)*.
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).
т 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 а chromist black polecat, the so-called “P. р. stantschinskii” was
the basis for this mistake. Only a short time ago, the white polecat had slightly
penetrated into Poland from Rava-Russkaya region.
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. Нершег.
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
769
770
1157
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
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 (“Р.
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 рам, 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.
771
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
(“talassica”) 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).
772
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 aurea).
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.
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. р. 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
ТИ
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 1$ 16.2—24.2
mm; females, 16.6-19.4 mm; postorbital width of males is 12.0-
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;
774
1165
and to very characteristic structure of winter fur. Although as
pointed out, some characteristics of this form are revealed in gen-
eral 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
775
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° М. 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.
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
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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
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
775
1168
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 TSS 4.9 7.4 — eg 3.0 2-2
Harvested birds 3.27 — 0.8 — 1.16 — —
Соттоп 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 — — 23:1 — 0.97 — —
Insects — a — — — — 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
ТИТ
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
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” т 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
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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
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
200, mating (7 cases) was observed from 9 April to 9 June. Ежег-
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).
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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
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.
719
780
1174
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
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 ржа [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.
п an active season, one ground squirrel eats about 6 kg food, 25% of
which consists of grains.
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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
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.
2177
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
See
Dental formula I 3 С 7 P a M Aa 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
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 Blas., and reconstructed range of
marbled polecat, Vormela peregusna Gueld. V.G. Heptner.
783
079
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).
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.
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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 Gildenstaedt, 1770
1770. Mustela peregusna. Gildenstaedt. 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, р. 60.
Tedzhensk oasis, Turkmeniya.
1910. Vormela sarmatica alpherakyi. Birula. Izhegodn. Zoolog.
muzeya. Ak. Nauk, 15, p. 333. Near Ashkhabad.
1910. Vormela negans. Miller. Proc. U.S. Nat. Mus., 38, p. 385,
plate 17. Ordos desert, about 100 miles north of Yulinfu
(Yuilin’).
1935. Vormela peregusna peregusna natio intermedia. Ognev. Zveri
SSSR 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, р. 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 co.
1948. Vormela peregusna obscura. Stroganov. Ibidem, p. 131.
Vakhsh valley, Tadzhikistan (V.H.).
Diagnosis
Only species in its genus.
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 А.М. Komarov.
1182
785 Fig. 278. Marbled polecat Vormela peregusna 019. 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 Ъе-
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**
*In Russian original, reads “summer,” a lapsus—Sci. Ed.
**In Russian, chebrak, from Turkic shabrak, or saddle blanket—Sci. Ed.
786
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
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
паре 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.
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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
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
789
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 118-205 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]).
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.).
788
See,
Fig. 279. Skull of marbled polecat, Vormela peregusna Giild.
EAS SERA OS
CS г. ony ws ~S
< =
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1187
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 т
above sea level. It is found along the entire Caspian shoreline to
Derbent, and farther south, to Baku.
791
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”
М. lat.) and somewhat north of it—to Gukasyan region (41° 00° М.
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-
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 П’изК 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
Fig. 280. Reconstructed range of marbled polecat, У. peregusna Giild. in USSR V.G. Нершег.
790
1191
Tadzhikistan along river уаШез 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 пт.
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.
792
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.’
In the interfluve between the Dnepr and Voiga, the northern
border, begins near Voronezh, descends somewhat southward, and
then, passing slightly north of 50° М. 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
3Sokur (1960) demonstrated its occurrence also in Nikolaevsk district (Vladimirsk
region); however, this place was not shown in the map and no details are reported.
193
marbled polecat is already only sporadically distributed, rare ог
even very rare everywhere west of the Don; in many regions
it appears occasionally and is obviously on its Mey 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; Korneev, 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.
795
794
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-
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.).
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
SShanxi 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.
ПЭ
Nn
1196
adjacént 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 (“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
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
“соПаг”). Color of lateral occipital stripes usually yellowish. Scapu-
lar stripes very distinct; of thigh, weakly defined and bearing dark
796
1197
spots. White frontal band not divided medially and has по соппес-
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 rearbled 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
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
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 У. р. koshevnikovi Sat.
(on left) and Semirech’e marbled polecat, У. р. 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
797
Fig. 283. Marbled polecat surveys its surroundings before coming out of Из 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 has 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).
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 (“соПаг” 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
(“alpherakyi’) to the Central Asiatic (Ordos) V. р. 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,
798
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 еихта, see above, and ornata): У. р. 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.
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.).
799
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—gerbiis,
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
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
799
800
Fig. 285. Marbled polecat in colony of great gerbil, Rhombomys opimus. 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
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
к
800 Fig. 286. Tracks of marbled polecat on wet sand in а 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
801
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.
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
Р.И.)
1207
Genus of Honeybadger ог Ratel*
Genus Mellivora Storr, 1780
1780. Mellivora. Storr. Prodr. Meth. Mamm., р. 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.
Dental f ] tcl РМ 22 ae
ental formula 1 СТР Му=32.
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.
34 tl ncude to]
**In Russian original, erroneously given as I = C=] PAM = 32 а Ed:
Тан
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).
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
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 (p.1179) 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, 15. 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 т
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.
804
80
Nn
1211
RATEL OR HONEYBADGER?
Mellivora capensis Schreber, 1776
1776. Viverra capensis Schreber. Sdéugeth. pl. 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
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.
3“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
“it-ail”), which means “dog-bear” (it—dog).
Description given mainly from material in ZMMU.
1212
805 Fig. 289. Ratel, ог honeybadger, Mellivora capensis Schreb. Kushka—Kashan
interfluve, southern Turkmenia, 1960. Photograph by Yu.K. Gorelov.
806
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
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-
шге“. 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.
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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
808
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
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 sharp 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”.
809
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.
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, У. 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 ratel 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.
810
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
(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; Jolbarsli well (north of 38° М. 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
811
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
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. Zhernovov, I.V. Svyatyi, V.P. Kostin, and collec-
tions of Z[oological] M[useum of] M[oscow] U[niversity].
№221
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 (10’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
812
222
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.
Within the boundaries of the USSR, there is only one form,
apparently, not differing in any substantive way from the Indian.
Indian honeybadger, M. c. 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. c. 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. c. leuconota
Sclater, 1867—West Africa northwards to southern Morocco; 5) М.
c. signata Pocock, 1909—Sierra Leone; 6) M. c. consica Thomas et
Wroughton 1907—Lake Chad; 7) M. c. buchanani Thomas, 1925—
Air plateau (Asben), Sahara; 8) M. c. abyssinica Hollister, 1910—
Ethiopia; 9) M.c. broekmani Wroughton, 1920—Somalia; 10) M. c.
cottoni Lydekker, 1906—Ituri region, Congo; 11) М. с. makwelli
Thomas, 1923—Kenya; 12) M. c. sagulata Hollister, 1910—Tangan-
yika; 13) M. c. vernayi Roberts, 1932 Bechuanaland; 14) M. c. 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).
813
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-
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.
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.
815
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 м. The diameter ranges from 17—18 to 30 cm, more often
25—30 cm. The distribution of burrow depth ranges from 25 cm
to 1.5 т. 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
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
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.
814 Fig. 294. Hilly landscape in Badkhyz preserve, southern Turkmenia. Habitat of
ratel or honeybadger. May, 1961. Photograph by Yu.K. Gorelov.
1227
NS . К
А
\
AY US SSN
К
Fig. 295. Tracks of honeybadger. Sketch by V.P. Kostin.
816
1228
31 May; she gave birth to one cub. In the first days of Мау, а
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.
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.
817
1229.
1795. Taxus. Cuvier et Geoffroy. 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.
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.
Дани 1
3
Complete dental formula is I 3 C 7 P я М от 38. However,
individual first premolars ог 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 as regards specialization of the dentition. The southern Asiatic
Arctonyx, also closely related to Meles, is usually considered a
*This is now the accepted name for the genus of ferret badgers—Sci. Ed.
1231
818 Fig. 296. Range of the genus of badgers, Meles Briss., and species range of the
common badger, Meles meles L. Borders of the range in Afghanistan, China and, in
part, in the Indochinese Peninsula are to a certain extent given approximately. Dotted
lines separate the regions of the distribution of the main race groups of badgers—
European meles, Siberian—Chinese “sandy” badgers, arenarius—leptorhynchus, and
Far East badger, amurensis—anakama. A question mark in the region of the watershed
of Kama and Pechora points to the unclear limit between the two race groups.
V.G. Heptner.
more specialized form (Pocock, 1941), although, apparently, with-
out sufficient basis.
Very primitive representatives of the subfamily, or forms close
to it (Broiliana, Stromeriella) are known from the early Miocene
of Europe and Asia. A series of genera which indicate the great
818
1232
variety of forms of the group are known from the upper Pliocene
of the Old World. The majority of them, however, as regards the
line leading to the genus Meles, represent lateral branches with
various specialization (Palaeomeles, Plesiomeles, Taxodon and
others). Aside from them, Parameles (Odessa), Melodon and
Parataxidea stand closest of all to the genera Meles and Taxidea.
Parataxidea is known from the upper Miocene and lower Pliocene
of Asia and is widely distributed, itself representing a separate,
more specialized, group. Melodon must be considered the source
of the genus Meles.
The genus Meles itself appeared in the upper Pliocene from
whence are known M. gennevauxi, M. taxipater, and M. thordi and
from which through M. m. atavus the line leads to modern M.
meles. This is, apparently, the typical Asiatic form, developing
without close connection to the American badger (Taxidea) which
is known only from America (upper Pliocene). It is, apparently,
related to the Asiatic Parataxidea.
The genus includes one species: Meles meles Linnaeus, 1758.
In practice, the forms of badger are of little positive impor-
tance (fur, destruction of some pests).
In the USSR it is distributed throughout the greater part of the
European territory of the country, through the southern half of
Siberia and the Far East, and in Middle Asia (V.H.).
BADGER
Meles meles Linnaeus, 1758
1758. Ursus meles. Linnaeus, Syst. Nat. Ed. X, 1, p. 48. Uppsala,
Sweden.
1785. Meles taxus. Boddaert. Elench. Animal, 1, p. 80. Europe.
1816. Meles europaeus. Demarest. Nouv. Dict. Hist. Nat. Substi-
tute for meles.
1844. Meles anakuma. Temminck. Fauna Japonica, Mamm. p. 30,
pl. 6. Vic. Nagasaki, Japan.
1847. Taxidea leucurus. Hodgson. J. Asiat. Soc. Bengal, 16, p.
763, pl. 29. Lhasa, Tibet.
1859. Meles taxus amurensis. Schrenck. Reisen und Forsch. im
Amur-Lande, p. 17, pl. 1, Fig. 1. Amur near mouth of Ussuri
[rivers].
1867.
1875.
1891.
1895.
1899.
1900.
1901.
1901.
1905.
1907.
1910.
OS:
1916.
1926.
1926.
193i:
1931.
1940.
1233
Meles leptorhynchus. Milne-Edwards. Ann. Sc. Nat. Zool.,
8, p. 374. Vic. Peking [Beijing], China.
Meles canescens. Blanford. Ann. Mag. Nat. Hist., 16, p.
310. Abadekh, between Shiraz and Isfahan, Iran.
Meles schrenkii. Nehring. Sitzungsber. Ges. Naturf. Freunde
Berlin, p. 103. Substitute for amurensis Schrenck.
Meles taxus arenarius. Satunin. Arch. Naturg., 1, р. Ш. Ryn-
peski, between lower Volga and Ural [rivers].
Meles meles typicus. Barrett-Hamilton. Ann. Mag. Nat. Hist.,
4, p. 384. Substitute for meles Linnaeus.
Meles taxus sibiricus. Kastschenko. Kashchenko. Opredelitel’
Mlekop. Zhiv. Tomskogo kraya, plate 15. Former Tomsk
governance. Restricted to (Stroganov, 1962) Kolyvan’, about
400 km north of Novosibirsk.
Meles amurensis altaicus. Kastschenko. Kashchenko.
Ezhegodn. Zool. muzeya I. Ak. nauk, 6, p. 613. Southern
end of Teletsk Lake, Altai.
Meles amurensis raddei. Kastschenko. Kashchenko. Ibidem.,
p. 613. Steppes of Trans-Baikaliya.
Meles meles minor. Satunin. “Priroda i okhota’, No. 2.
Borzhom, Trans-Caucasus.
Meles blanfordi. Matschie. Wiss. Ergebn. Filchner Exp. nach
China, 10, 1, p. 143. Kashgariya.
Meles tianschanensis. Hoyningen-Huene. Zur Biol. d. estland.
Dachases, p. 63. Tyan’-Shan’ [Tien Shan].
Meles melanogenys. J. Allen. Bull. Amer. Mus. Nat. Hist.,
32. p. 433. Musan on Tumangan river, northern part of
Korean Peninsula.
Meles meles ponticus. Blackler. Ann. Mag. Nat. Hist., 18, p.
75. Skalita near Trabzon (Trapezund), [Trebizon], Asia Minor.
Meles meles caucasicus. Ognev. Uchen. zap. Sev-Kasvk. inst.
kraevedeniya, 1, p. 50. Vic. Vladikavkaz.
Meles meles tauricus. Ognev. Ibidem. 1, p. 51. Beshuisk
forestry camp in Chatyr-Dag, Crimea (Crimean preserve).
Meles leptorhynchus talassicus. Ognev. Zveri Vost. Evropy
i Sev. Azii, 2, p. 478. Southern slope of Talassk Alatau,
western Tyan’-Shan’ [Tien Shan].
Meles meles heptneri. Ognev. Ibidem., p. 775. Aleksandro-
Nevskaya north-west of Kizlyar, eastern Cis-Caucasus.
Meles meles severzovi. Heptner. Zeitschr. Г. Séugetierk., 15,
p. 224. Arkit, south of Syry-Chilek Lake, south of the north-
820
82
—
1234
eastern extremity of the Chatkal range, Tyan’-Shan’ [Tien
Shan]. —
1962. Meles meles aberrans. Stroganov. Zveri Sibiri, 2, p. 167.
Akmolinskaya_ district, Bogembai village, northern
Kazakhstan (V.H.).
Diagnosis
The only species of the genus.
Description
General body constitution heavy and massive, especially in full
winter pelage. Entire body seems to possess a wedge-shaped form—
the broad and convex posterior half suddenly narrows anteriorly
and, through the short ill-defined neck, passes to the relatively
small, narrow, obviously elongated head. From the ear region, the
head, continuing the general outline of the body, the wedge pro-
ceeds to the noticeably elongated, narrow facial portion with the
somewhat extended, quite movable nose.
Limbs short and massive, plantigrade with naked lower surface
of feet. Claws strong, elongated (on middle digit of fore limb, 30-
35 mm along curve, 22-26 mm in straight line) and obtuse at end,
adapted to digging. Tail short, approximately equal to head length.
Ear pinna relatively very small, rounded. Bare tip of nose quite
large, separated from mouth margin by narrow band of fur. Eyes
are small.
The animal usually moves slowly and heavily, as if listless,
lowering its head. The posterior part of its trunk appears higher
than the shoulder and the general appearance is depressed. The
badger does not flex the back as martens, polecats and wolverine
do, and does not stand erect like the honeybadger. However, it can
run quickly at a gallop.
In winter fur, pelage on back and sides is long and coarse,
consisting of coarse, even bristly guard hairs and sparse quite soft
undercoat. The venter is covered with short, sparse hairs and skin
is usually visible in the inguinal region. Length of guard hairs on
the middle of the back in winter 1$ 75-80 mm.
Color of the Middle Russian badger in its winter fur coat, i.e.
late in autumn before retiring for the winter is as follows. Throat,
lower neck, chest and legs black, venter black with light brownish
1235
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Fig. 297. Badger, Meles meles L. Sketch by А.М. Komarov.
820
tints, inguinal regions brownish
dorsum and sides is a beautiful and pure light silvery-gray, as if
coated with translucent black ripples. On the sides, there are straw-
colored highlights, more in the lower part. Such coloration is
determined by differently colored guard hairs which have a broad
(35-38 mm) dirty-whitish or straw-whitish basal part, and after it
a wide (22—25 mm) deep black band and a white or slightly grayish-
white tip 10-12 mm long. On the sides, bases of hairs have more
intensive straw tone, the black band is somewhat narrower, and
1236
ends of the hairs is \ВИе'. The tail is covered by long coarse
hairs, quite fluffy, and generally of the same color as the back.
The head is white, with white covering lower lips and chin
also, where it is sharply bordered by the black throat. Two pure
black bands pass along the head, sharply differentiated from the
white background. Each of these begins on the upper lip at the
mouth opening anterior to the corner of the mouth, and initially
passes directly upwards. Somewhat below the level of the eye, the
bands pass backwards at a right angle and, rising somewhat
upwards and getting wider, pass through the eyes and, getting still
wider, cover the whole base of the ear and even a part of the
parietal region. In this region, its width is greatest.
Behind the ear, the band, sometimes widening somewhat again,
extends along the dorsolateral part of the neck sometimes to its
middle (sometimes noticed somewhat farther back) and here disap-
pears, gradually merging with the color of the upper body. Some-
times, it slants downwards between the color of the side of the
neck (corresponds with back color) and the white field below. The
width of the band anteriorly is about 15 mm, and in the ear region
it is 45-55 mm. The black tone of the band is more intense in front
of the eye, between eye and ear, around the ear and directly behind
it. In the anterior section, the band frequently has a dirty or grayish
tone and its outline is not sharp. Most frequently, the initial
vertical portion of the band is weakly defined or almost undefined.
The black facial bands outline a wide white band with almost
parallel sides which extends from the nose tip through the [midline
of the] forehead and crown. This band covers the occiput and the
anterior part of the neck and sometimes almost the whole neck,
gradually merging with the color of the upper body. Sometimes it
does not lose its breadth and sometimes it tapers to a point and is
quite well outlined.
The white fields of the ventro-lateral surface of the head ex-
tend backwards as a wide band of quite even width, between the
black color of the lower head and neck and the black head band.
Posteriorly, these [lateral] bands usually extend to the same dis-
tance as the upper [mid-dorsal] white band, and sometimes more—
they occur notably on the greater part of the neck’s length between
‘Sometimes it is stated that along the badger’s spine, gradually spreading out
posteriorly, runs a brown band. This is a misundertstanding—such a pattern is absent
in our badgers.
822
1237
the gray color above and the black below. Posteriorly, they gradu-
ally merge with the color of the upper body. The ears are black
with wide (to 17-18 mm) white edges along the upper margins.
This edging is well marked against the black area of the facial
band. Claws are dark-horn with yellowish cast.
Summer fur is much shorter, coarser and sparser. Length of
guard hairs along the back is more than 50 mm. On the venter,
hairs are particularly sparse and in places, the skin is visible through
them. Color of badgers in summer is darker and more dirty dorsally,
with yellowish or ocherous tinge; black tones ventrally are not as
deep, but brownish. Yellowish tinge of the fur is determined by the
corresponding color of the guard hair tips.
In the badger, there is one molt, extending the whole summer.
New hairs attain their full growth only in hibernation. Therefore,
summer fur is actually old being replaced and newly growing.
General tone depends in part on wear and dirtiness of the fur.
Individual variation in badger color is quite obvious. Equally
with several predominantly moderate “gray” types, there occur
lighter and even very light animals generally of an almost silvery
color. More frequently observed is a more or less significant dark-
ening of general color tone of the dorsum, or the appearance on it
of yellowish tones, giving the whole skin clayey highlights. Changes
in intensity of color of the black bands on the head occur, and also
the clarity of the white tone, which may be darkened. A brown tint
may appear along the black ventral fields. According to some data,
in each locality two color types are present—lighter and darker.
Mention of color dimorphism in this species (Stroganov, 1962) is,
however, without foundation. It is ordinary fluctuating variation.
Among sharp mutational changes, melanistic, albino and eryth-
ristic forms are known, and badgers of yellow color (Neal, 1948).
Sexual differences in color are absent. Newborn animals are
covered with short white fur; in slightly older animals, fur is lighter
than in adults. By the first hibernation, young animals don adult
pelage. With increasing age, white portions of the hair increase in
badgers, they lighten, and may even become very light.
Geographic variation in color is considerable and, it appears
that relative to the described color types, it lies in somewhat weak
lightening on one hand, and in strong darkening on the other. In
the latter case, color may attain a brown tone, while light fields of
the head become so dark that facial bands become poorly visible.
1238
The extent, form and location of black facial bands change also.
In some forms, they do not cover the whole base of the ear, but
pass below it, usually only touching the inner margin of the ear
conch base, and usually are narrower.
The skull (Middle Russian badgers) is quite massive and heavy,
noticeably elongated and, if crests are disregarded, quite even in
outline, not angular and moderately high.
The line of the dorsal skull profile, if the sagittal crest is not
considered, is convex, with the highest point in the interorbital
region. From this point, the line of the facial part descends at an
obvious angle. Even steeper is the profile of the nasal opening.
Posterior from the interorbital constriction, the profile line forms
a very gentle arch, but descending more strongly in the occiput.
The braincase is moderately developed, not swollen and not elon-
gated, rounded-oval in outline, while the facial part of the skull is
elongated and relatively narrow.
Zygomatic arches are narrow anteriorly and widely separated
posteriorly, with their greatest width at the level of the articular
condyles. Zygomatic width is considerably more than mastoid. The
arches themselves are strong and massive, curved upwards in their
posterior parts. The postorbital constriction of the skull is well-
developed, but short and not sharply defined, and the skull width
here is slightly less, equal or almost, equal to interorbital width. In
very old individuals, a fairly sharp intersection is formed here.
Supraorbital processes are short, but well developed, sharply out-
lining the orbit, the dimensions of which are relatively very small.
Its greatest diameter constitutes slightly less than half the interor-
bital width. The orbit opens posteriorly, but bone borders more
than half its circumstance.
The hard palate, especially that part lying behind the toothrow,
is long and narrow. Width of the palate between the inner margins
of the molars is approximately 3.5 times less than its length from
the anterior edge of the sphenopalatine notch to the alveoli of the
middle incisors. The interpterygoid groove limiting the palate
posteriorly is short and wide. Its length is only slightly more than
the distance between the ends of the hook-shaped processes. The
tympanic bullae are of irregularly angular (triangular) form, slightly
swollen medially and moderately compressed (flattened) laterally.
They do not contact the articular process. Mastoid processes are
strongly developed and form massive outgrowths behind the
823
1239
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Fig. 298. Skull of the badger, Meles meles L.
auditory tubes which open independently of them. Paroccipital proc-
esses are short, triangular in form, their bases in contact with the
posterior parts of the auditory bullae. The longitudinal diameter of
the nasal opening is greater than the transverse. Infraorbital
foramina are very large, irregular oval-rounded in outline. Their
vertical diameter is equal to the diameter of the canine or greater
1240
than it: its greatest diameter is approximately half the diameter of
the orbit.
Sagittal (arrow-shaped) and occipital crests are strongly devel-
oped—in old males, the height of the sagittal crest may be equal
to half the interorbital width and even more. This crest reaches its
greatest height in the middle of the parietals, arching downward
posteriorly, and its end overhangs the occipital crest and occipital
region in the form of a hook.
The lower jaw is relatively light and elongated; its ramus is
almost straight; jaw height in the middle part constitutes about
one-third the height in the region of the coronoid process. Poste-
rior to the dental region, the lower line of the jaw profile rises
upwards quite abruptly. The angular process is thick and short, the
coronoid, broad and high—its height above the level of the articular
process is almost equal to its width at this same level. The posterior
region of the lower jaw has a low longitudinal crest ventrally.
Sexual dimorphism in the skull is insignificant and, besides
dimensions, is manifested in somewhat less development of crests
and generally somewhat “infantile” appearance of females. In young
animals, the braincase is more swollen, postorbital constriction
less marked—its width not less, but equal to or even more than
interorbital width (in the very young animals, it is distinguishable),
protuberances, etc. of skull are weakly defined, crests are not
developed or only beginning to form. Complete development of
the skull is, apparently, attained in the second year, although fu-
sion of sutures begins earlier.
Geographic variation in the skull, not counting the size, is
either weakly manifested or almost absent. There is some change
in predominating form of the infraorbital foramen, in part (sculp-
turing) of the skull.
The upper carnassial tooth is relatively small, in plan, triangu-
lar outline (not pointed)—its length approximately equal to its width.
Along the outer side, it bears one tall cutting cusp. The first molar
is strongly enlarged in dimensions—its area is not less than the
second, and greater than the area of the carnassial. It is rhomboid
in form with parallel outer and inner sides, the main diameter lying
approximately along the line of the toothrow; its inner side is
greater [longer] than the outer. The tooth carries some low cusps,
from which the two main ones are located along its outer edge and
one lies more or less in the middle of its cup-shaped crown.
825
1241
The lower carnassial tooth is large, strongly pointed, with three
well developed apices in the anterior part and with an elongated
flat and concave posterior part (“heel”); in length it exceeds the
anterior part. The small premolars (one upper and two lower) are
of the cutting type. The very small first upper premolar is easily
lost and is frequently absent in adults, its alveolus is also not
evident. The first lower premolar may also disappear and the
4 Знай
Bahk 1 ЗЕ
26=P—M—= = 6 p= Me
dental formula I17C7P7M> 38 becomes Laer aMs 36
or even тс 1 P ; M : = 34 or perhaps asymmetrical. With aging,
the upper molars may be more flattened as a result of wear of its
apices.
Loss of the first premolar in different parts of the range occurs
in different percentages of individuals (geographic variation). The
same is true with the lower premolar; in a portion of the races, it
has two roots, and in another—one.
The os penis is almost straight, with the dorsal side slightly
concave or somewhat curved in an arc, and oval in cross section.
At its tip, it is strongly widened in the form of a spoon and pierced
in the middle with a small foramen elongated in the longitudinal
direction. The sharp hook-shaped bend of the anterior end charac-
teristic of the subfamily of martens, Mustelinae, is not formed.
Body length is 600-900 mn,,’ tail length 120-240 mm, length
of hind foot, 75-130 mm, height of the ear is 35-70 mm.
Condylobasal length of male skull is 113.0-144.0 mm, of fe-
males, 100.0-140.0 mm; zygomatic width of males is 68.0-90.0
mm, of females, 62.0-85.5 mm; interorbital width of males is
25.2—34.5 mm, of females, 22.0-33.0 mm; postorbital width of
males is 19.9-27.5 mm, of females, 18.1-26.2 mm; mastoid width
of males is 55.9-71.9 mm, of females, 52.3-67.0 mm (from ap-
proximately 300 males and females of the Z[oological] M[useum
of] M[oscow] U[niversity] and Stroganov, 1962, Siberia).
Os penis length is 72-76 mm.
*Body length of “about 90 cm” was stated by Ognev (1931) as the average length
of old male badgers in Moscow province. The majority of authors refer to this length
as maximal. Only Kuznetsov (1952) refers to greatest length of European badger as
105 cm, while Kashchenko (1900, 1902) stated that the greatest length of the
Transbaikal badger is more than 100 cm. These data are, apparently, somewhat exag-
gerated, although the length of 90 cm is possibly not maximum.
1242
Weight of badgers at the time of their active life changes very
greatly, growing from spring to autumn, and reaching its maximum
entry into hibernation. The badger accumulates very much fat, and
relative to its general dimensions, its weight is disproportionally
great. Average weight in autumn of badgers in the European part
of the country, the northern Caucasus and Siberia is about 16—17 kg;
a few large and particularly fattened animals reach 20-24 kg
in weight; reports of badgers of 30 kg (Siberia; Stroganov, 1962)
and 30 and 34 kg (Moscow and Ryazan districts; Ognev, 1931)
are not reliable.
Weight of the Middle European badger in summer is 7-13 kg,
in autumn 15—17 kg and as an exception—20 and perhaps even 25
kg (Gaffrey, 1961). Average weight of English male badgers is
12.2 kg, females somewhat less. The maximum established weight of
males is 19 and 19.5 kg and of females is 17.2 kg. There exist unveri-
fied reports of animals of 20.4 and 22.6 kg in weight (Neal, 1948).
Females are somewhat smaller and lighter than males. Dimen-
sions and weight of badgers vary somewhat geographically. Al-
though these changes are not very sharp, extreme forms (Middle
Russian—Amur, for instance) differ considerably (V.H.).
Systematic Position
Only species in the genus.
Geographic Distribution
Europe, Near and Middle Asia, southern and middle zones of Si-
beria, the southern Far East, China, and Central Asia.
Geographic Range in the Soviet Union
This constitutes a considerable part of the species range—its north-
ern half and part of the western, and occupies the greater part of
the country.
In the northwest, the northern border of the range begins in
“the southwestern corner of Russian Laplandia” (Pleske, 1887):
here, apparently the following is understood as districts to the west
of Kandalaksha on the south, including Kuusamo (presently—
Kusamo, in Finland). This is apparently the most northernly habi-
tat of the species in Europe, lying at about 67° N. lat. On the Kola
827 Peninsula, the badger is absent, but to the south of Kandalaksha,
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it is encountered all over Karelia and in particular, recorded at
Ruvozer (Marvin, 1951, 1959). Thence eastward to Arkhangel’sk,
the southern coast of the White Sea forms the northern border of
the range. On Solovets Islands the badger is absent, and it is not
known whether it occupies the northern parts of Onezhsk
Peninsula.
On the eastern coast of the White Sea (Zimnii [winter] coast),
the northern border of the badger’s range begins north of
Arkhangel’sk, slightly above 65° N. lat. (Shil’dlog village or far-
ther north). Thence, it passes eastwards along this latitude to Kuloi
and farther to Mezen’ [still] on 65° N. lat. (Leshukonskoe village).
Between Kuloi and Mezen’, the border gives a sharp sag to the
south—a narrow extension—descending to Karpogar on the Pineg
[river]. From Leshukon on Mezen’, the range border directs itself
to the northeast along an irregular line and reaches the region of
the source of Pesha river, flowing into Cheshsk gulf. This [point]
is only 50-70 km south of the Arctic circle.
From here, the range border turns sharply to the southeast and,
proceeding somewhat to the southwest of Ust’-Tsyl’ma on the
Pechora, it goes to the middle course of the left tributary of the
Pechora—the Izhma river south (about 70—80 km) of Izhma city.
Thence, rising somewhat to the north, the border passes to the[main]
Pechora at 65° N. lat. (data of V.Ya. Parovshchikov for 1962).
Farther, the border goes to the Urals, apparently along this same
latitude (Narodnaya mountain) or a little to the south—to Lyapinsk
Urals (Flerov, 1933).
It is possible that, in the European part of the country, the
range of the badger extends in some places even more to the north.
Thus, a badger was caught at Kuloi near Kar’epol’e (50-60 km
more south of Mezen’ city; Zhitkov, 1904) and near Mezun’ city
(southern border of forest-tundra; Parovshchikov, 1959) and even
on the coast of Cheshsk gulf near the mouth of the Pesha (forest-
tundra; Parovshchikov, 1959). In these places, the character of the
occurrence of this animal is not entirely clear, and the available
data require more precision. However, it is evident that this far
penetration to the north is connected with river valleys.
In western Siberia the badger was recorded near Salekhard on
the Ob’ at the Arctic circle (Stroganov, 1962; the most northern
point of occurrence in Siberia). The border sinks sharply to the
east and passes through the extreme upper reaches of the Pur and
828
1245
Taz (about 63° М. lat.; Yanushevich and Blagoveshchenskii, 1952*).
It is possible that the actual northern border in the Trans-Ural lies
somewhat farther to the south—near Berezov on a little lower
(about 64°; Yanushevich and Blagoveshchenskii, 1952**) and cor-
responds more closely to the situation of the border in the Urals
and eastward from the lower Ob’. The Berezov—upper Pur and
Taz is nearer to the permafrost border. It is possible that its dis-
tribution to Salekhard is a distribution just along the Ob’ valley.
However, on the other hand the animal was noted at Khal’ mersede
north of the Arctic circle (Leble, 1953).
From the region of Taz sources, the border, going eastward,
descends to the south and crosses Yenisei, somewhere at about 61°
N. lat. At this latitude, or a little north, it goes into the basin of
Podkamennaya Tunguska where the badger is met with at Baikit—
at the mouth of Chunya in Tunguska (appr. 61° 35°: М. Naumov,
1934) and a little eastward on the Mutorya river (Podarevskii,
1936). It is entirely possible that the actual border of the range on
the right bank of the Yenisei proceeds more to the north—the
animal was noted (apparently transient) at the mouth of the Taimura
river in Nizhnaya [lower] Tunguska (N. Naumov, 1934; this same
place is sometimes referred to as the mouth of the Vivi river—both
rivers flow into the Tunguska at nearly the same place).
Farther to the east, information on the limits of badger distri-
bution are very few. Apparently, the border first directs itself
towards a left [bank] tributary of the Lena—the Ichera, then ex-
tends somewhat northeastward (perhaps along the Lena valley) to
the mouth of the Vitim (about 59° 30° М. lat.; Maak, 1851; О.
Ivanov) and even to a point about 150 km farther south of Suntar
on the Vilyui (О. Ivanov, about 60° 40° М. lat.). This place lies on
the Lena or very close to it. In the latter case it is, very likely,
always found as a transient since it is very distant—to Kudu-Kyuel’
village on the Olekma is approximately 115 km in a straight line
*“1958” in Russian original—Sci. Ed.
**“1953” in Russian original—Sci. Ed.
3The border in western Siberia is generally poorly clarified, and information is,
in part contradictory. Thus, the earlier data (Slovtsov, 1892; Chugunov, 1915) speaks
of its distribution in “Surgut Territory” and not north of 62° N. lat. in Surgut region
on the Ob’. According to I.P. Laptev (1958), the northern border of the range, begin-
ning at Berezov, embraces*** Kazym and passes somewhat to the south of the sources
of Pur and Taz, reaching the Yenisei river at the mouth of Podkamennaya Tunguska.
***Misspelled “okhvatyvaet”, instead of “obkhvatyvaet” in Russian original
—Sci. Ed.
1246
from its confluence with the Lena (О. Ivanov). Transgressions to
the two above-mentioned places and in part to the mouth of the
Vitim are very rare—3 cases during 40 years (D. Ivanov).
From the mouth of the Vitim, the border curves, apparently
arching a little towards the east, including the Bodaibo region, and
turns back towards the southwest, apparently passing to the Kirenga
river (to the west of Baikal) without reaching the northern extrem-
ity of Baikal and somewhere here, it reaches the shore of the Lake
(from data of V.V. Timofeev)’.
In Trans-Baikaliya, the outline of the range no less compli-
cated than to the west and north of [Lake] Baikal. The border in
this region does not, judging from everything, extend so far to the
north. Starting, apparently, on the shore of Baikal somewhere around
the mouth of the Barguzin [river], it passes along the eastern slope
of the Barguzin range on the northeast, crossing the Barguzin [river]
at approximately 55° N. lat. and turns abruptly back to the south
or southeast, including the Ikatsk range lying along the left bank
of the Barguzin and just fails to reach the sources of the Ok’
(tributary of the Uda—about 53° 30° М. lat.).
Thence, the border bends sharply to the east and northeast,
crosses the Vitim [river], proceeds through the region of the Amalat
headwater—a tributary of the Vitim—(in its bend) and extending’
northeastward, passes somewhere south of the Kalar—in all events
including the sources of the Olekma and its tributary, the Tungir;
i.e., nearly reaches 56° N. lat. (V.V. Timofeev, S. Anashkin).
Therefore, apparently, the range beyond Baikal envelops the Vitim
plateau, or at least a part of it.
From the above description, it follows that that part of the
range located in the upper Lena north of Baikal (Bodaibo) is not
directly united with Trans-Baikal. Not excluded is the possibility
that such a complex picture of the northern range border in the
Baikal region is formed on account of a deficiency of accurate
data. However, it is most probable that this is the actual situation,
and it is explained by the fact that the badger bypasses to the north
and south of the northern Baikal and Stanovoi highlands, the south-
ern and northern Muisk ranges and several ridges connected with
them (Kodar, Udokan).
4On the map of Stroganov (1962, p. 162), the occurrence of badger
northeast of Baikit on the Podkamennaya Tungaska was mentioned. This 1s,
apparently, an incorrectly plotted habitat on the Mutorga river.
829
1247
In Yakutiya, the badger is apparently not a permanent member
of the fauna. It is known only from previously mentioned
transgressions to the mouth of the Vitim south of Suntar village,
and at Kudu-Kyuel’ village on the Olekma river.
From that place on the upper Olekma in Trans-Baikaliya at
about 56° N. lat. the border goes southeast, south of the Gilyui
(Gassovskii, 1921) approximately through Solov’evsk to Zeya city
and then to the middle course of the Selemdzha. Thence, it de-
scends still more steeply to the southeast—to upper Burei, a little
below the mouth of the Niman and in the source of a tributary of
the Burei—the Tyrma. Farther along the Evoronsk lowland the
border ascends towards the northeast, to the middle course of the
Amgun’, and then proceeds again to the southeast, crossing the
Amur and reaching the ocean at 52° N. lat. (from materials of
A. Samsonov and V.P. Sysoev).
On Sakhalin, the badger is absent.
To the south of the above-mentioned line in the European part
of the USSR, the badger is encountered to the state frontier and
beyond it. It is also all over Siberia. In Middle Asia, the badger is
met with throughout the mountains and is only absent in individual
localities, mainly in extreme high-montane regions. Thus, the ani-
mal is apparently absent in the high-montane Pamirs, most notably
in the western [Pamirs]. Badgers do not, however, avoid deserts.
In the Kyzyl-Kum, they are encountered at least along the edge;
they are also recorded in the Ust’ yurt, at least the eastern [Ust’ yurt]
(Bazhanov, 1951) and along the lower Amu-Dar’ya (Gladkov and
Nikol’skii, 1935).
In Turkmenia, the badger lives mainly in montane regions,
occupying in particular the whole of the Kopet-Dag and Bol’shoi
Balkhan and, apparently, the Gyaz’-Gyadyk mountains (left bank
of upper Tedzhen), Kushka valley and mountains to the east of
Kushka (Chengurek mountains), and over the expanse lying along the
mountains (V.G. Heptner). In the Karakum, the badger is rare, but is,
apparently, met with almost everywhere (Yu.F. Sapozhenkov).
There are some unclear points and contradictions in the char-
acteristics of the northern border of the range sketched above, and
compared to several other works, may be explained not only by
incomplete information and imprecision concerning transgressions
of animals, and by natural fluctuations of the border during severe
1248
conditions in the north of the continent.° Apparently the badger,
like some other species, has expanded its range northward in the
last decade. Thus, according to relatively recent information
(Parovshchikov, 1959), the northern border in the European part of
the country shifted from Arkhangel’sk to Karpogory on the Pinega,
thence to the upper Mezen’ (Koslan) and across the middle course
of the Izhma to the right tributaries of middle Pechora—the
Shchuger* and Podcher’e. The border therefore passed consider-
ably to the south of 65° N. lat.
Geographic Range outside the Soviet Union
The range occupies Europe, to the west including England and
Ireland, and to the south to the coast of the Mediterranean Sea. It
is on the Balearic Islands, Corsica, Sardinia and Sicily, as well as
on the islands of the Aegean Sea except Rhodes, and, apparently,
on Cyprus, the badger is absent; it exists in Crete. The northern
border of the range in Europe begins on the Atlantic coast of
Norway at Trondheim Fjord (about 63° 30° М. lat.; Ekman, 1922)
and, describing an arc, curves northward (to approximately 64° 30°
N. lat.) reaching thé Baltic coast about the latitude 63° N. lat. In
Finland, the border begins on the Gulf of Bothnia approximately
at Raahe (about 64° 40’), and beyond forms a bend to the south,
but in the east of the country again rises somewhat northward and
nearly reaches 66° N. lat. (perhaps even as far as Kusamo; see
above).
In Asia, the range occupies all of Asia Minor, Syria and Pal-
estine, Iraq (apparently, only northern), northern, northwestern and
western Iran (except the extreme south and the belt adjacent to
southern Iraq) and northern Afghanistan. It may penetrate, per-
haps, the territories connected with the Hindu Kush, to the more
southern parts of the country contiguous with Baluchistan. In the
east, the range extends to Kashgariya and Dzhungariya (details
unknown—it is apparently absent in deserts); the entire northern
half of the Mongolian Republic, southward including the Mongo-
lian Altai, Khangai, Kentei, and sections lying to the south of these
montane parts of the country, the extreme east of it. In China, the
range occupies the entire western part of the country—from former
‘In Arkhangel’sk district, after cold winters with much snow and late springs, the
number of badger falls sharply (V.Ya. Parovshchikov).
*Misspelled “Shchugor” in Russian original—Sci. Ed.
830
1249
Manchuria to the extreme south. The range apparently extends to
the extreme north of Vietnam—north of Hanoi. In the east, the
range in China everywhere reaches the Pacific Ocean or to the
borders of our country; its distribution to the west is not clear. It
occupies Inner Mongolia, Gansu, Shanxi, Sichuan and, apparently,
Yunnan or а part of this province (the opinion that Uadger are
absent from this region has little basis), and possibly, the extreme
north of Burma (information of its occurrence in Arakan between
latitudes 20 and 15° N. is erroneous).
According to some data, the range also includes Tibet. In ac-
tuality, it does not, apparently, occupy all of this territory, which
is less suitable for the badger’s life, and occupies only a part of
it—perhaps the southern and southeastern margins. Thus, the Chi-
nese race was noted in Lhasa and Dzhangdze (Gyantse), southwest
of Lhasa, and the plains north of Sikkim (Pocock, 1941). Perhaps
the “Tibet” range extends through southwest China and Upper
Burma as a projection eastward along the Brahmaputra valley and
regions connected with it possessing relatively more favorable
natural conditions. The range does not penetrate adjacent India (or
Nepal) but the possibility of penetration of badger from the north-
west southward to Chitral, and from the north to Baluchistan is not
excluded. The range area also includes Japanese Islands® and Korean
Peninsula (V.G.).
Geographic Variation
Racial variation in the badger is considerable. It manifests itself
both in amplitude of change in several characteristics and in the
number of existing forms. About 40 names have been given to the
badger and 24 subspecies are usually accepted, of which 11 are
within our country (Ognev, 1931; Ellermann and Morrison-Scott,
1951; Stroganov, 1962). This number is evidently too great. To-
gether with that, those attempts to revise the racial composition of
the species that have been done do not reflect the actual situation
and themselves represent an extreme in the opposite sense—4 forms
for our territory (Novikov, 1956) and even 4 races for the whole
species (V. Petrov, 1953).
It is sometimes claimed that it occurs on Hainan (Pocock, 1941; Tate, 1947).
This error is apparently based on a misunderstanding. Pocock (1941) offered data of
Delacour about its occurrence “in the foothills north of Hainan”. The last word is a
misprint for “Hanoi”. This text was so used above.
1250
я ny
Ач
аа
а.
SS у
PLN
: <
830 Fig. 300. Head pattern of various race groups of badgers—above, European badger,
group meles; middle—‘sand” ог Siberian—Chinese, group arenarius—leptorhynchus;
below, Far Eastern, group amurensis—anakuma. Sketch by N.N. Kondakov.
831
1251
Geographic variation of badgers reveals itself in general di-
mensions, skull measurements (for the most part insignificant),
form of the infraorbital foramen (a character not very well defined
and permanent), form of the upper molar—its relative width (as
well), structure of roots of the second lower premolar, relative
frequency of loss of first premolars, general color tone and in form
of head pattern. Color characteristics are most stable, typical and
distinct. There are some geographic peculiarities in individual details
of skull structure.
According to color type, all races of badger are clearly divis-
ible into three groups. The remaining characteristics more or less
agree with coloration. Races directly “intermediate” are absent;
however, several other characters create “transitional” features in
separate populations.
1. Group meles, European badgers. General color quite light
and pure gray, “silvery”; sides of body a little lighter than middle
of back. A wide black, rarely black-brown, stripe passes through
eyes and envelops ear above and below (entire base of ear pinna).
A pure white stripe passes from nose between [black] bands across
forehead, behind ears and along neck. Cheeks and sides of head
white, the white stripe passing along sides of neck’. Upper molar
relatively wide—ratio of length to width less than 1:5; second lower
premolar usually with two well-developed roots, first upper and lower
premolars for the most part retained. Infraorbital foramen usually more
elongated in vertical direction. Dimensions large to moderate.
In Europe eastward to the Volga, Caucasus, Near East, south
and southeast Middle Asia.
2. Group arenarius—leptorhynchus, Siberian or “sand” badg-
ers®. Color lighter, but in some forms, color corresponds to that of
preceding group, and sometimes even darker, smudged with
ocherous and brownish highlights. Sides a little lighter than middle
of back. Dark facial stripe usually not black, but brownish in tone;
it narrows behind eye and extends above ear. It does not go far
posteriorly, sometimes hardly reaching ear. White color on head
"This type of coloration was described above in more detail in the section
“Description”.
*Animals of this type were recorded in our country for the first time from the
Ryn sands between the lower Volga and Ural [rivers]. The form described from there
by Satunin was called arenarius; i.e., sand. This Russian name was introduced in our
literature for the entire group of races, although the majority of them have no relation-
ship to sands (montane, forest).
1252
usually dirtyish. Light stripe passing along head dorsally between
dark stripes relatively narrow and short; it does not extend behind
ear or only extends behind it for a small distance and already
merges with color of dorsal body on occiput.
Upper molar relatively elongated—ratio of length to width 1.5
or nearly that; second lower premolar usually with one root; first
lower premolars for the most part absent. Infraorbital foramen more
extended in horizontal direction. Dimensions moderate.
In Trans-Volga; Siberia except Amur region and Ussuri basin;
Middle Asia except extreme south and southeast; and all remain-
ing areas of Asia except Near East and notheastern China (former
Manchuria) and Japanese islands.
From a zoogeographic and systematic point of view, it is note-
worthy that in Europe, the Volga, and in part, the Kama (lower
course) rivers separate two sharply differentiated race groups—
European and sand badgers. They evidently play the role of an
insuperable mechanical [physical]* barrier for badgers—animals
which in these latitudes are not active in winter. This fact has
multiple significance both in several general respects (problem of
barriers, problem of clines, and others). If in some forms (see
below), individual “transitional” characters are noticed, they do
not apply to facial pattern. At the same time, the limits of distri-
bution of both groups of badgers in the northeastern European part
of the country, and in the northern Ural region are not clear. In the
Pechora-Ilych preserve (upper Pechora), the Siberian badger
exists. In Tien-Shan (see below), the races of both groups also
approach each other closely, but are sharply separated by mountain
ranges.
3. Group amurensis—anakuma. Far Eastern badgers. General
color very dark, with brown tones strongly developed. Head pat-
tern corresponds to that of the Siberian badgers; however, head
dark, and dark stripes weakly defined.
Upper molar is elongated, second lower premolar with one
root, first premolars always absent. Infraorbital foramen wider in
transverse direction. Dimensions small.
In Amur region, Ussuri basin, northeastern China (former Man-
churia), and Japan.
Badgers of groups 1 and 2 were, up to the 30’s, considered
different species—M. meles and М. leptorhynchus (Ognev, 1931).
*In Russian original, the word mekhanicheskoi is used—Sci. Ed.
1253
The Amur-Ussuri badgers were affiliated with the latter. The
recent attempt to restore this completely neglected point of view
(V. Petrov, 1953), was naturally unsuccessful.
The scheme of races given below is preliminary.
Group of European badgers, meles.
1. Middle Russian badger, M. m. meles Linnaeus, 1758 (syn.
tauricus, caucasicus).
Dimensions large.
Color of back relatively pure silvery-gray tone. Main tone of
head pure white, dark stripes wide, black in color, white fields
extend far backward along upper and lateral parts of neck.’ Pelage
relatively soft with relatively dense underfur.
Skull with strongly developed crests, second lower premolar
with two separate or merely adjacent roots; first premolars, often
all, usually present; upper molar relatively wide—ratio of length to
width about 1.25, and usually not more than 1.35. Infraorbital
foramen extended in vertical direction, and its greatest diameter,
being directing upwards and somewhat internally, greater than trans-
verse diameter.
Dimensions maximmal for its group; apparently, for entire spe-
cies. Middle Russian badgers are largest—they are, evidently, larger
than Middle and Western European.
Condylobasal length of male skull is 125—140 mm, of females,
123.2—133.6 mm; zygomatic width of males is 75.2—89.5 mm, of
females, 68.0-85.2 mm; mastoid width of males 1$ 62.1—-71.9 mm,
of females, 58.5—67.0 mm; length of upper molar of males is 14.7-
17.1 mm, of females, 14.5-17.1 mm; greatest width of posterior
molar of males is 11.2—13.2 mm, of females, 11.0-13.5 mm (Ognev,
1931; some deviation in both directions possible). In animals of
this race, different numbers of first premolars retained in different
parts of range—in Middle and Northern Russia, only 77%, in the
south—60%, in Cis-Caucasus—66%, and in Crimea—about 72%
(V.G. Heptner).
Weight up to 20-24 kg (autumn), in exceptional cases, perhaps
a little more (see above).
In European part of Union eastward to Volga, in the Crimea,
Cis-Caucasus and northern Caucasus. Distribution to east in north
°The above-given description (page 1234) applies to this form.
1254
of European part of country not known (see below, separation
from Siberian badgers).
Outside the USSR, in remaining parts of Europe except Rhodes,
Crete, Spain (existence of a separate race in Denmark is unlikely).
Badgers from the northern slopes of the Caucasus, its foothills
and adjacent plains may be, on average (skull dimensions) slightly
smaller than Middle Russian. In this one might see some tendency
toward transition to the Trans-Caucasian form. This difference is,
however, completely insignificant and separation of a different form,
caucasicus, is not justified.
2. Kizlyar badger, M. m. heptneri Ognev, 1931.
Dimensions large (as in the preceding form).
Color very pale, dull, dirty-grayish-ocherous. Black stripes on
head narrow, but typical of European group.
Skull as in preceding form.
In Caspian steppes area (steppes of northeastern Cis-Cauca-
sus), right bank of lower Volga (Kalmytsk steppes), and Volga
delta.
Outside the USSR—absent.
A very poorly known form; exhibiting several characters simi-
lar to the Siberian badger group, but on the whole typical of Eu-
ropean [group]. Range is very small, especially as compared to
other subspecies, but on the whole, characteristic of a series of
mammalian races, mainly of eastern origin, which are also restricted
in their distribution to the above-mentioned section of steppes,
deserts and semideserts.
This form requires further study and here is provisionally rec-
ognized.
As yet there is no basis for placing it in synonymy with the
nominal form.
3. Trans-Caucasian badger, M. m. canescens Blanford, 1875
(syn. minor).
Dimensions smaller than preceding forms.
General color of dorsal side dirtyish-gray with highlights of
brown tones. Head color as in nominal form. Skull and tooth struc-
ture as in nominal form; i.e., infraorbital foramen high, but upper
molar somewhat elongated, though not so strongly as in Siberian
badgers, and crests somewhat more weakly developed.
Condylobasal length of male skull is 116.0-123.3 mm, of
females, 111.8—122.0 mm; zygomatic width of males is 68.0-81.5
1255
mm, of females, 66.0-72.0 mm; mastoid width of males is 56.0-
63.2 mm, of females, 56.1-58.0 mm; length of upper molar in
males is 15.9—17.3, its width 10.0-11.1 (Ognev, 1921). In badgers
of this form, about 57% of possible number of first premolars are
retained (V.G. Heptner).
In Trans-Caucasus, Kopet-Dag and apparently, Gyaz’-Gyadyk
mountains (right bank of upper Tedzhen); probably elevated re-
gions of Kushka (Chengurek mountains) in Turkmenia. Limits of
distribution in northern Turkmenia not known.
Outside the USSR—in Iran, Afghanistan, and possibly Asia
Minor.
4. Fergana badger, M. m. severzovi Heptner, 1940.
Dimensions quite small, apparently close to preceding form.
Color of dorsum relatively pure silvery-gray without yellow
sheen or with only insignificant development of it. Black stripes
on head wide occupying whole ear.
Upper molar short and broad; i.e., typical of European badger
group, preorbital foramen extended in transverse direction; i.e., а
character of Siberian badgers.
Condylobasal length of skull (10) is 109.4-М 118.4-127.0
mm; zygomatic width is 64.5-М73.2-81.6 mm. In animals of this
race, the least number of the first premolars—about 26% of the
number possible, among all groups of races (V.G. Heptner).
In region of right tributaries of Pyandzh and upper Amu-Dar’ya,
Pamiro-Alaisk system (apparently, Eastern Pamir only; the Alaisk
valley), Fergana valley and mountains bordering it on south and
north (Chatkal and Fergana ranges).
Outside the USSR—absent.
The Fergana badger stands closest to M. m. canescens.* How-
ever, it is substantially different from it, several skull characteris-
tics “transitional” to the Siberian badger group in particular being
most noticeable. It is probable that its range is separated from that
of M. m. canescens by the Pyandzh [river]. There is a remarkably
sharp boundary between its range in the north and badgers of the
leptorhynchus group (sand) which occupy the northern ranges of
the Tien-Shan system. Already along the southern slopes of the
Talas Alatau south of Issyk-Kul, and along the upper Naryn are
distributed typical representatives of the Siberian badger group.
The limits between both forms in the plains are not clear. In all
*See above, Trans-Caucasian badger—Sci. Ed.
834
1256
events, М. т. severzovi was recorded in Kugitangtau (Ishunin,
1961), and M. m. leptorhynchus in Nuratau (Kuznetsov, 1948).
Other data on badgers of Middle Asia are poorly defined, and
partially contradictory (Kuznetsov, 1948; Ishunin, 1961).
Group of Siberian, or “sand” badgers
arenarius—leptorhynchus
5. Siberian badger, M. m. sibiricus Kastschenko, 1900 (syn.
altaicus, raddei, aberrans).
Dimensions moderate—somewhat smaller than in nominal form,
but larger than in Trans-Caucasian.
General color tone of back light—gray, usually with yellowish
or straw-colored and sandy-straw highlights. Dark stripes on head
typical for group—narrow and extend along dorsum of head, not
including ear. Their color varies from brownish-black to tawny-
brown. Pelage long and relatively soft with dense undercoat.
Upper molar relatively elongated, ratio of its length to width about
1.5. Infraorbital foramen extended in transverse direction, second
lower premolar with one root, and first premolars very rarely occur.
Body length of males (6) is 675-М720-750 mm, of females
(4), 620-M668-692 mm; tail length of males is 185-М214-235
mm, of females, 174-М196-222 mm; length of hind foot of males
is 100-М116-127 mm; of females, 82.0, М84.2-86.0 mm; ear
length in males is 38-M39-41 mm, of females, 37-M38—40 mm.
Condylobasal length of male skull (7) is 123.0-М125.8—128.7
mm, of females, (5) 116.0-M117.4-118.7 mm; zygomatic width of
males is 75.6-М78.0-80.5 mm, of females, 68.4-М69.2—70.2 mm;
interorbital width of males is 27.3-М27.8-28.3 mm, of
females, 26.2, M26.6—27.0 mm; postorbital width of males is 22.0—
M23.8-—24.6 mm, of females, 20.0-M23.2—24.0 mm; mastoid width
of males is 64.0-М68.7-75.4 mm, of females, 58.6-М60.2—62.0 mm.
Percentage of retained first premolars differs in different
populations of this race—in Bashkiria and southern Urals—about
13[%], in Siberia and Altai—3[%] and in Trans-Baikaliya—O[%]
(V.G. Heptner).
Weight of adult males reaches 10-13.6 kg, and evidently even
more (all data on dimensions after Stroganov, 1962).
In Siberia including Trans-Baikaliya and Altai, northern parts
of Kazakhstan and probably northern Trans- Volga.
835
1257
Outside the USSR—unknown.
The independence of the Siberian form of badger was some-
times, and is now, doubted. At the same time, its differences from
the form arenarius, and from the Mongolian form leptorhynchus,
is sufficiently real. Separation of this race is arguable, but the
racial identity of, for example, badgers from the middle course of
the Ob’ and from the deserts and semideserts of southern
Kazakhstan and Uzbekistan one may hardly admit even theoreti-
cally. Neither is the view by which all badgers from the
Trans-Caucasus, Siberia and Turkmenia to Central Asia, China and
Tibet belong to one form (Petrov, 1953; Novikov, 1956) acceptable.
The separation of the form aberrans Stroganov, 1962, demands
confirmation. This form is, to a high degree, doubtful, both as
regards its characters and its range (“northern regions of eastern
Kazakhstan”—Tselinograd, Kokchetav, Pavlodar, Zaisan Lake and
middle Tarbagatai).
Concerning Trans-Volga badgers, see the following form.
6. Kazakhstan or sand badger, M. m. arenarius Satunin, 1895.
Dimensions moderate—somewhat less than in nominal form,
but greater than in Trans-Caucasian form.
Color is lighter and paler than in northern forms, conditioned
by decrease in black bands of guard hairs and their frequent full
reduction on sides of body, where the color acquires a pale-straw
tone. A clayey tone usually develops over the entire skin, which
impinges on the silvery-gray color characteristic of northern races.
Pelage is coarse, bristly and underfur is scarce.
Structure of skull and dentition as in Siberian badger, M. m.
sibiricus.
Body length of males (4) is 700-780 mm, of females (4), 610—
700 mm; tail length of males is 200-220 mm, of females, 185-250
mm; length of hind foot in males is 115-126 mm, of females, 95—
110 mm; ear length in males is 58—70 mm, of females, 40-80 mm
(Sludskii, 1953).
Condylobasal length of male skull is 115.0-131.0 mm, of
females, 109.0-118.0 mm; zygomatic width of males is 68.6-77.7,
of females, 62.2-73.8 mm; mastoid width of males is 55.9-65.0
mm, of females, 52.3-58.8 mm; width of upper molar in males is
10.1-12.1 mm, of females, 10.1-12.3 mm; length of upper molar
in males is 14.1-16.6 mm, of females, 14.0-16.0 mm. Percentage
of retained first premolars—about two (V.G. Heptner).
1258
Weight of males in March—May is 7.8—8.3 kg, in March—June
is 5.6-7 kg. Weight of young animals in November is 10.5—12.5
kg, of an adult female, 14.5 kg (Sludskii, 1953).
In southern Trans- Volga (Volga-Ural steppe), Kazakhstan ex-
cept the northern and montane parts, and plains of Middle Asia
except the parts occupied by the forms canescens and severzovi
(extreme southern Turkmenia, Pamir-Alaisk mountain system,
Fergana valley and the mountain ridges bordering it on the north).
Outside the USSR—absent.
This described form of badger is morphologically and
geographically well-characterized. Its distribution in northern Trans-
Volga is not clear. It was recorded in the southern Urals (Kirikov,
1952), at the latitudes of Samarsk Luka and Bashkiria. However,
these data indicate only the presence of badgers belonging to the
“sand” group in these places, but they do not confirm its identity
with the form arenarius from the Volga-Ural steppe and Middle
Asia. This is all the more so, since badgers of northern Kazakhstan
belong to the Siberian form sibiricus. It is possible that these Trans-
Volga badgers (except southern) also belong there. In general, as
was shown, limits of distribution of the sand badger group to the
north in the Cis-Urals are not known to be contiguous with the
meles group.
Information on the occurrence of this form in the steppes of
the Cis-Caucasus (“Caucasian steppe”, Ellerman and Morrison-
Scott, 1951) was mistaken. As was shown, all of the forms of the
leptorhynchus group are absent west of the Volga and Caspian
Sea.
7. Tien-Shan badger, M. m. tianshanensis Huene, 1910 (syn.
talassicus).
Dimensions moderate.
General color same as type, and as in Kazakhstan badger, M.
m. arenarius, but somewhat darker, the yellow sheen is weaker or
not developed, the fur longer, denser and fluffier. Percentage of
retained first premolars is about 8.3% (including material from
around Dzhungariya; V.G. Heptner).
In the northern ranges of the Tien-Shan including Dzhungarsk,
Zailiisk, Kirghizsk and Talassk [Alatau]. In the eastern parts of the
[mountain] system within the USSR, southwards as far as the upper
Naryn [river].
Outside the USSR—probably in the Tien-Shan system in
Dzhungariya and Kashgariya [China].
836
1259
It is a completely unclear and entirely doubtful form which
requires additional study and confirmation. It is to the highest
degree close to the Kazakhstan badger, M. m. arenarius (many
individuals are not distinguishable at all) and apparently, differ
only in somewhat denser underfur, as is frequently the case with
montane populations.
The southern limits of this form and the whole “sand” badger
group requires more precision. The Fergana badger, M. m. severzovi
(see above), exists along the southern slopes of Chatkal and, prob-
ably, Fergana ranges and Tien-Shan form along the upper Naryn.
Note. The problem of variation and nomenclature of badgers
of the northern Tien-Shan is not clear. Hoiningen-Huene (1910)
named the Tien-Shan form from living animals received from
Hagenbeck* and caught in “Tien-Shan” but not noted whether
within the borders of the [present] USSR or in China. From pho-
tographs and the description it is clear only that this is a typical
“sand” badger. Ognev (1931), without attempting to precisely limit
the type locality (t. t.** restricta) of the form tianschanensis, allo-
cated to it individuals from the extreme eastern—Chinese—parts
of the Tien-Shan (Kul’dzha, Kunges river, Kaitsalai, Yuldus, Boro-
Khoro range). At the same time, he described the distribution of
this form, according to data of N.A. Severtsov, in a series of places
in the eastern parts of the Tien-Shan within the borders of the
USSR westward to the Chu river as well as in the western Tien-
Shan, “for the entire range in general” and even for the Karatau
(page 478). Factually, therefore, it is to take the habitation of the
form tianschanensis throughout the entire Tien-Shan, within the
borders of our country and beyond them.
On a par with this, in the same work [Severtsov, 1873] (pages
478—479), М. т. talassicus was described from Talassk Alatau
(southern slope) from two specimens; the occurrence of which is
assumed to be also on the Chu river (Frunze-“Pishpek’”***). Dif-
ferences from the form tianschanensis given are very insignificant
(somewhat darker color). The place of origin for M. m. tianschanen-
sis has not yet been accurately established—and besides it is en-
tirely doubtful—that it is possible that there are different badgers
of the type arenarius—leptorhynchus from the different parts of
Tien-Shan, there is no basis for admitting the independence of the
form talassicus.
*German animal dealers—Sci. Ed.
**For “terra typica”; i.e., type locality—Sci. Ed.
***Presently Bishkek, Kyrgyzstan—Sci. Ed.
1260
Group of Far Eastern badgers, amurensis-anakuma.
8. Amur badger, M. m. amurensis Schrenk, 1858 (syn.
schrenkii).
Dimensions very small, smaller than in the Trans-Caucasian
badger M. m. canescens. It is the smallest race of the species.
General color very dark, darkest among all races. Dark stripes
on head extend above ear and disappear in dark field of occiput;
they are black or dark blackish-brown or sometimes coal-black in
color. Entire area between stripes and cheek region, i.e., places
which in other races are white or very light, are dirty grayish-
brown in color. It may be so dark that stripes are weakly
distinguished or almost undetectable. Color of back very dark due
to strong development of dark bands of the hairs, grayish-brown
with silvery highlights, depending on white tips of hairs. Sides of
body only slightly darker than back. Pelage with little wool, but soft.
Skull small, with smooth outlines, relatively light. Protu-
berances, etc. on skull weakly defined, and crests very weakly
developed even in oldest individuals (wear on crown of carnassial
teeth). Entire skull has infantile features. Infraorbital foramen ex-
tended in transverse direction, upper molar is elongated. First
premolars always absent (extreme degree of reduction within
limits of species).
Body length 60-70 cm.
Condylobasal length of male skull (6) 1$ 111.0-117.9 mm, of
females (6), 104.8—112.2 mm; zygomatic width of males 1$ 67.8—
75.4 mm, of females, 62.1-73.2 mm; mastoid width of males is
57.3-—63.5 mm; of females, 53.0-58.2 mm; interorbital width of
males is 24.0-29.2 mm, of females, 22.1-24.6 mm; postorbital
width of males is 20.8-22.4 mm, of females, 18.2—21.9 mm
(Stroganov, 1962 and materials of Z[oological] M[useum of]
M[oscow] U[niversity]). Length of upper molar in males is 16.2—
17.0 mm, of females, 14.0-15.3 mm; width of upper molar in
males is 11.2—11.5 mm, of females, 9.6-10.8 mm (Ognev, 1931).
Badgers of this form are always characterized by complete
absence of first premolars and corresponding change in dental for-
mula (V.G. Heptner).
In Ussuri Territory and Priamur’e westward approximately to
123° E. long. (Albazin).
837
1261
Outside the USSR—in contiguous parts of northeastern China
(former Manchuria), and probably westward to include the Bol’-
shoi [Great] Khingan [range] and Korean peninsula.
The Amur badger is a sharply distinguished form. In its head
pattern, it resembles races of the “sand” badger group, but is sharply
separated from them and all other forms of the species by its satu-
rated darker color, by size and complete reduction of all first
premolars; i.e., by dental formula. This form, undoubtedly, pos-
sesses features similar to the Japanese badger, M. m. anakuma;
however, the union of both races is unfounded.
The borders of the range of the Amur subspecies are still not
fully clear. There was some information that badgers of Trans-
Baikaliya possess characters transitional to the Amur form, but
this is not so. In the Mongolian Republic, near Beijing, and
possibly in the central Manchurian plains lives a badger of the
typical “sand” type, extremely light.
The Amur badger is a clear example of one of the character-
istic features of geographic variation of some widely distributed
species of Palearctic mammals—darkening or more saturation of
color, and partly decrease in size in the south of the Far East
(Cervus elaphus, Capreolus capreolus,* Mustela sibirica and
several others).
Beyond the borders of our country, a quite large number of
farms have been described. The independence of many, if not the
majority, of them needs confirmation. Usually accepted are: 1) M.
m. anakuma Temminck, 1844—Japanese Islands (Hondo [Honshu],
Shikoku, Kyushu, ? Hokkaido); 2) M. m. melanogenys J. Allen,
1913—Korean Peninsula (evidently a synonym of M. m. amurensis
V.H.); 3) M. m. leucurus Hodgson, 1847—southeastern Tibet
(Lhasa); 4) M. m. leptorhynchus Milne-Edwards, 1867—China,
Mongolian Republic’®; 5) М. т. blanfordi Matschie, 1907—
*Now considered a distant species, C. pygargus—Sci. Ed.
Osgood (1932) considers the forms [еисигиз and leptorhynchus identical. This
point of view was, apparently, also supported by G. Allen (1938). In this case, the
Chinese-Mongolian badger must be called leucurus (not leptorhynchus as accepted by
Allen). In general, the identity of badgers from Lhasa and from around Beijing (the
type localities of both forms) is very doubtful, and this question needs further study.
Allocation to leucurus of the badgers of Siberia, Middle Asia and even Trans-
Volga (Bashkiria; У. Petrov, 1953) belong to Jeucurus is, of course, not accepted.
838
1262
Kashgariya; 6) М. т. marianensis Graells, 1897—Pyrenese [Ibe-
rian] Peninsula; 7) M. m. arcalus Miller, 1907—Crete; 8) M. m.
rhodius Festa, 1814—Island of Rhodes, Aegean Sea; 9) M. m.
ponticus Blackler, 1916—Asia Minor (very probably, a synonym
of M. m. canescens; V.H.); 10) M. m. danicus Holton, [1935]—
Denmark (evidently a synonym of the nominal form) (V.H.).
Biology
Population. Badger populations within the limits of its range are
variable. It is determined chiefly by the abundance of food, breed-
ing burrow conditions and sources of drinking water. In the breed-
ing period, in the large spruce swamps, and on the water divides
of the Volga and Zapadnaya [western] Dvina, its density was not
great, reaching 0.13-0.16 per 10 km’. In the forest-steppe, in is-
land of broad-leaved oakgroves (Tula abatis; Likhachev, 1956),
average density over 13 years was from 1.6 to 2.1 рег 10 km’. In
one of the great islands (28 km?) of the delta of Ili river in
Kazakhstan (Sludskii, 1953), density reached a maximum of 21.0—
26.0 рег 10 km? (15-16 litters). Data on the number of inhabited
burrows are less reliable: some burrows are only periodically in-
habited and in others, up to three litters live (badger “сиу”).
In the European part of the USSR, the badger is most numer-
ous in the middle zone—there where islands of forests prevail, and
the locality itself is hilly and cut with ravines. Thus, in Tatariya,
3—9 animals are found in 10 km? of wooded lands suitable for the
habitation of badger (Gorshkov, 1964).
The greatest number of badger skins are obtained from
Kazakhstan, although the animal itself is far from numerous through-
out the region. The least number of badgers is in the northern
Kazakhstan steppes. It is most numerous in Alma-Altinsk, Dzhambul
and Chimkent districts (Sludskii, 1953). In the taiga, both lowland
and montane, the badger is rare. The same must be said for the
open steppes.
Habitat. The badger reveals great flexibility with respect
to habitats. Just as in steppe and forest foxes, wolves, tundra
and taiga reindeer, etc. badgers of various races differ in their
ecology. The badger in the forest zone is a forest animal, mainly
restricting itself to forest edges, forest islands, coppices and wooded
ravines. It is most numerous in the zone of islands of mixed forests
1263
alternating with fields, meadows and settlements. It avoids large
forest massifs, rarely settling those, and if so mostly along edges;
such forests have little food for them. On the contrary, in a small
island forest of 15—20 h, there are sometimes up to three litters of
badger. In some localities (Byelorussia, Kirov district), the popu-
lation of badgers corresponds to the percentage of woodlands, which
is explained by the excessive pursuit of the badger by man, and not
by environmental conditions. In Moldavia (Korchmar’, 1963), the
badger inhabits the forest tracts of Kodra, in insular forests of the
northern regions and in southern forests. It is most numerous in
small islands of forest. For the badger, the forest is merely a good
refuge sheltering it during foraging, and not a necessary sur-
rounding for its life.
In the habitat, besides abundance of food, favorable burrowing
conditions are the most important requirement. This is natural for
an animal pursuing a semisubterranean way of life, in which the
greater part of the [24 hour] day and several winter months are
spent in the burrow. The badger prefers dry, sandy-loam soils with
a deep ground water level for easily-yielding excavation. In ex-
treme cases, it settles in more or less moist soils, choosing the
most elevated, well-drained places. Forest attracts the badger not
only as a shelter, but also thanks to the presence of the Ortstein
horizon which is impermeable to water (Kolosov, 1935), under
which it digs its burrows. For the same reason, it prefers sand
lying beneath a layer of clay or loam. The water-impermeable upper
layer of soil then serves it as the vault of its underground construc-
tions (Lukhachev, 1956). The badger sometimes digs burrows in
sands of pine forests, and also among rocks.
The second condition for the badger is the proximity of water,
especially when the approach to it from the burrow is hidden. In
open dry steppes of southeastern Trans-Baikaliya, habitats of the
badger are always connected with water resources. In Kazakhstan,
the badger sometimes makes its burrow not more than 5—10 metres
from water (Sludskii, 1955). The badger does not avoid human
settlements as long as the burrow itself is located in a little-popu-
lated, hidden place, most often on the slopes of a deep ravine
overgrown with trees, bushes and tall grasses.
In the middle zone, the badger prefers the edges of mixed
forests but it does not avoid coniferous forests. And here, it prefers
sections dissected by ravines, but sometimes settles on flat plains.
1264
839 Fig. 301. Montane oak and mixed coniferous-broadleaved forests. Typical habitats of
Amur badger in southern Ритог’е. “Kedrovaya Pad’ ” preserve. Here also exists a
fauna complex of Ussurian Territory—kharza, racoon dog, white-chested bear,
leopard, and others. July 1964. Photograph by A.G. Pankrat’ev.
In the forest-steppe zone, the badger more often burrows along the
slopes of ravines and gorges which are very often covered, if not
by forest then by steppe shrubs and rank growth of grasses.
In Kazakhstan, the badger is an animal of steppes and
semideserts, ecologically different from the badger of the forest
zone. In northern Kazakhstan, it settles along the slopes of ravines
839
840
1265
(saev), the high ridges near lakes in islands of pine woods, оп high
terraces, and along the river valleys. In central Kazakhastan, it
settles in the bank precipices and the slopes of hills and ravines,
especially the shores of lakes and rivers. In the desert zone, it lives
in consolidated sand hills alternating with solonchaks, streams and
lakes, in flood lands and deltas of rivers. Many badgers are also
found on “terraces”* of foothills, especially in apple forests on the
slopes of mountains. In mountains here and in the Caucasus it
ascends to a height of 2300-2500 т. In the montane Altai, it
restricts itself to the lower belt of the mountains—not higher than
1000—1200 т.
In Ussuri Territory (Yu.A. Salmin and V.D. Shamykin), the
badger is associated with more gentle relief. It is encountered in
valleys of large mountain rivers—in well-drained dry valleys at the
foot of slopes.
Food. Of the whole order of carnivores the badger, like the
brown bear, is the least carnivorous. In the badger as well, the
development of powerful teeth and masticatory musculature is con-
nected not with carnivory, but with omnivory. While in the pine mar-
ten with mixed feeding the intestine length surpasses body length by
4.5—5 times, in the badger it is 8 times (in the bear, by more than 10
times).
The badger’s food consists of rodents (mainly voles), rarely
small birds, amphibians (mainly frogs) and reptiles (mainly liz-
ards), insects and their larvae, molluscs, earthworms, various fruits,
nuts, berries and other plant food.
The species composition of food, and within it the ratio of
each food type, is subject to considerable variation, both seasonal
and geographic. The effect of geographic variation upon food
composition can be judged by Table 70, which presents the data
on the most important foods of 7 separate parts of the range.
Actually, the diversity of the food components is still more signifi-
cant. Apparently, in the northern parts of the badger’s range there
is more carnivory, less in the south where sources of its food are
more abundant and varied (Crimea, Kazakhstan).
Among mammals, common and red-backed voles are the most
important. The role of birds is not great. Usually, they are small
birds nesting on the ground, mainly during the nesting period, or
some kinds of wounded birds. The shores of large bodies of water
*In Russian original “prilavok,” or counter, shelf—Sci. Ed.
1266
840 Fig. 302. Broad-leaved forest in the valley of Kedrovaya river—typical habitat of
Amur badger “Kedrovaya Pad’ ” preserve, southern Primor’e. October 1963.
Photograph by A.G. Pankrat’ev.
constitute an exception (Kaletskaya, 1957). In destruction of eggs
and young of game birds, badgers, for the most part, are accused
without cause. In the forest zone, frogs are, in places, of great
importance. In the dry forests of Zhigula, the importance of
slow-worms [anguine lizards]* increases, while in the Buzulak pine
forest—it is lacertid.
Among invertebrates in forests, of greatest а аге
dung beetles, various ground beetles, stag-beetles, longhorn bee-
tles, forest may beetles and june beetles and their larvae, sawflies,
wasps and bumblebees and their larvae. The badger eats many
mollusks (mainly slugs) and earthworms (in large amounts). Among
plant foods, of greatest importance are acorns, hazelnuts, strawber-
ries, wild apples, pears and others. In Kazakhstan, the main foods
of the badger consist of insects, mainly mole crickets, and in some
years—Asiatic locust on which it is quickly fattened (Sludskii,
*The Russian word “veretenitsa” is applied to both Anguis fragilis and Coronella
laevis—Sci. Ed.
841
1267
Table 70. Geographic variation in badger foods (occurrence, %)
Food type Tula Abatis Tataria Zhiguli Buzuluk Montane Delta of Ryabinsk
(Likhachev, and (РВ: pine Crimea Ш river water
1956) adjacent Yurgenson) forest (Savina, (Sludskii, reservoir
regions (Shilova- 1940) 1953) (Kaletsk-
(Zharkov Krasova, aya, 1957)
and 1951)
Teplov,
1939)
Mammals 70.6 54.2 57.5 17.1 34.1 23.6 No data
Rodents 69.0 53.6 56.6 ТТ 29.8 20.8 14.5
Birds 27.0 23 2.6 4.1 3:5 8.3 17.6
Bird eggs 3:2 2.1 — — 2.2 12.5 15.3
Reptiles 4.0 0.6 8.8 22.5 — 31.9 24.6
Amphibians 26.3 8.6 — — — — 88.2
Insects 96.0 76.8 86.7 95.4 98.0 94.4 97.0
Beetle larvae 17.7 18.4 ПТ 36.9 9.3 0.6 —
Molluscs — 0.8 15.0 0.1 14.6 -—- 12.3
Plant material 23.8 28.3 23.9 — 50.7 15.0 77.6
Sample size 126 474 113 221 224 144 134
1953). In the Crimea, among insects in the badger’s food, ground
beetles and stag-beetles predominate; a large role is played by
fruits of dewberry, wild pear, apple, acorn, and beechnut. The
badger also raids vineyards (Savina, 1940).
Seasonality of food types is conditioned by the degree of their
abundance and availability, which is not the same throughout the
year.
The seasonal changes in food composition of the badger in
Moldavia are as follows (Korchmar’, 1962; % occurrence):
First half of summer Second half of summer—autumn
(572 excrement samples) (226 excrement samples)
Mouse-like rodents 18.3 Mouse-like rodents 67.5
Subterranean voles 10.4 Small birds 5
Stag-beetles 45.2 Ground beetle 10.0
Ground beetle 45.2 Rose chafer 10.0
Rose chafer 31.3 Beetle larvae 7.0
Beetle larvae 33.3 Plum 65.0
Sweet cherry [P. avium] 60.3 Dogwood 44.0
Strawberry 12:9 Apple 10.0
Pear 5.0 Grape 7.0
1268
In Zhiguli (Р.В. Yurgenson), dung beetles are most frequently
found in the badger’s food in May, but in June—twice less; later
they rarely occur. Land molluscs are most often found in July—
August, as is hazelnut. Red-backed voles [Clethrionomys rutilus]
and wild apples are utilized most of all in August. Mountain
leafhopper larvae are more encountered in the badger’s food in
June, beetle larvae in June—August, and wasps (3 species) in July—
August. All of this is associated in a very close way with the
natural abundance of these foods.
It is characteristic for the badger that during one hunt it catches
one food type predominantly, but in considerable amounts: thus, in
the stomachs of three badgers caught on 3 November, were 57, 63
and 75 frogs (А.М. Skorodumov); in one caught on 31 May—97
maybeetle larvae (Likhachev, 1956), on 27 May—65 maybeetle
larvae and 50 dung beetles, on 26 September—285 gm of wasps
and their larvae (P.B. Yurgenson), on 26 June—146 barbel beetle
larvae, on 5 July—362 larvae of bumblebees, and on 22 August—
152 earthworms (Likhachev, 1956). In Moscow district near Klin,
during autumn, the stomachs of several adult badgers killed in the
morning when they returned to the burrow after a night’s foraging,
were full of earthworms only, or with small terrestrial frogs only.
2 The latter were, evidently, taken from hibernating aggregations
(V.G. Heptner). In Central Europe (Rorig)*, 9 old and 74 young
voles and 7 frogs were found in one.stomach, etc. In general, the
badger most often eats not more than 0.5 kg food per day, and
young (up to one year old), more than adults. Therefore, an adult
badger with an average weight of about 15 kg eats daily a quantity
of food equal in all to 3-4% of its live weight.
In the badger are clearly revealed differences in foods in dif-
ferent biotopes. Thus, in Tataria and adjacent regions, amphibians,
gray voles, fish, briar, blackberry and orthopterous insects рге-
dominate in foods of animals living in water-meadows; in pine
woods—maybeetle, click beetles, red-backed voles and blueberry;
in broad-leafed forests—dung beetles, mouse-like rodents, earth-
worms and acorns (V.A. Popov). The significance of the badger in
destruction of the harmful rodents and insects is quite noteable. In
Tataria, in young pine woods, up to 60 thousand places dug by the
badger were located, which here destroys up to 75% of all maybeetle
larvae (Gorshkov, 1964).
*Not in Lit. Cit.—Sci. Ed.
842
843
1269
Fig. 303. Complex old burrow of European badger in mixed fir-larch forest. South of
Vyazniki, Vladimir district. October 1964. Photograph by V.G. Heptner.
Home range. Almost no data. The badger, apparently, moves
away from its burrow a considerable distance, but is most active
within a radius of about 400-500 т from it; i.e., within an area
of about 100 h. Isolated ranges, it seems, occur only in those
places where badgers are few and the area yields little food. With
abundant food, badgers live near each other, sometimes 2-3 fami-
lies in one burrow of complicated construction. A series of cases
exist in which the inhabitants of one burrow visit a neighboring
one and peacefully live together (Neal, 1948). In Moldavia, in
early spring and late autumn, the radius of badger activity equals
2—3 km; in summer and the beginning of autumn, this range shrinks
to 0.5-1.5 km (Korchmar’, 1962). In Tataria, in early spring, the
badger feeds near the burrow, in summer, it moves away from it
for 2-3 km, maximally for 5 (Gorshkov, 1964).
Burrows and shelters. The badger is a typical burrowing ani-
mal. Only in very extreme cases, with insufficient places suitable
for burrowing, does it creep in winter into a haystack or into a hay-
loft in a forest glade. In England, there was a case where a badger
constructed a surface nest-lair in bushes, where it gave birth to 5
young.
1270
843 Fig. 304. Entrance into permanent burrow of Amur badger, made among stone blocks
at the trunk of a Korean pine. “Kedrovaya Pad’” preserve, southern Primor’e.
June 1962. Photograph by A.G. Pankrat’ev.
The number of exits in one burrow fluctuates from 1-2 to 44—
50 (Neal, 1948). In the vast old underground constructions of this
animal—badger “cities’—several badger families sometimes set-
tle. In this event, most frequently, several systems of passages
and nesting chambers occur, most often isolated from one another.
Usually, some exits are continuously used, the remaining are only
used in case of danger, or serve for the play of young badgers. In
Tula abatis (Likhechev, 1956), 68.8% of the breeding burrows had
from 2 to 5 exits. Their width at the base is from 22 to 63 cm,
height—14—32 cm; they are more often semicircular (arched) form.
The nesting chamber is positioned 5-10 meters from the exit open-
ing, at a depth of not less than 1 т from the soil surface, more
1271
often at a depth of about 1.4 т, but in individual cases, it is even
deeper—up to 2-3 meters. General length of the passages varies
from 35 to 81 m, with cubic capacity of the entire construction
ranging from 8.3 to 19.2 плз. In one case, the volume of the freshly
thrown-up ground near the burrow was 38 т? (Moscow dist.).
Average dimensions of the nesting chamber are 74 x 76 cm, with
an average height of 38 cm (Likhachev, 1956). Bedding consists of
dry leaves and grasses, which are periodically renewed. Leaves are
always collected on the dry days. There occur 2-3 nesting cham-
bers. Some of them are open from both ends, and others are blind.
In some complicated burrows the system of passages and chambers
form certain layers (“floors”). Special “ventilation” passages do
not occur in badger burrows. In winter, the openings are closed by
ground and leaves. By winter, up to 5 kg leaves and grasses for the
bedding are collected and the temperature in this bedding is kept
constant. The soil at the depth of the nesting chamber most often
does not freeze (Likhachev, 1956).
BS Ae neg SRR Say eee Tae! Bi ERS
844 Fig. 305. Occasional day lair of a badger, secluding itself after a night hunt, under a
small twisted aspen. Boggy dark aspen wood. In the background of the photo is a small
lake. South of Vyaznikov, Vladimir district. October 1964. Photograph by V.G. Heptner.
845
1272
Spring cleaning of the burrow is connected with birth of the
young (Neal, 1948). Each year, apparently, a new nest chamber is
dug for the birth. During the course of the summer it is changed,
sometimes several times, apparently in order to rid it of
ectoparasites. For the same reason, periodic cleaning of the burrow
occurs. The main work of building a burrow dates to August—
September, when a newly formed pair of young dig a new burrow,
and old burrows are prepared for hibernation.
Daily activity and behavior. As a rule, the badger is active at
twilight and at night. On bright moonlit nights, activity drops
sharply. Time of emergence from the burrow clearly coincides
throughout the entire year with the beginning of the twilight (a
little after sunset), and changes proportionately with day length
(Neal, 1948). Only in extremely remote places does the badger
emerge from the burrow during the day, but does not leave it—it
usually lies in the entrance and basks in the sun.
Among the sense organs, the best developed of all is smell—
it is the main organ of orientation. The badger’s vision is mono-
chromatic. The badger does not react to a lantern with red light.
Only moving objects attract the badger’s attention. Its hearing is
no sharper than that of the human. Since it makes significant noise
while moving itself, the badger cannot catch the simultaneous move-
ment of a human. In capturing food, it is mainly oriented by smell.
Torpor, winter sleep. Winter sleep in the badger is the adap-
tation of an omnivorous predator to existence under severe
climatic conditions with cold, snowy winter, when its main foods
disappear or become unavailable. Winter sleep in the badger, as
also in the bear, is not accompanied by a lowering of body
temperature and slowing down of all vital functions.
Already in the second half of summer, the badger begins to
accumulate fat reserves, which reaches its limit in October. By this
time, the burrow is cleaned and the nesting chambers are filled
with bedding. In the middle zone of the USSR, the badger retires
for winter sleep in the period from the second half of October to
the middle of November. It usually stops leaving the burrow after
the snow falls. In very warm winters, badger tracks are observed
up to the end of January (Tula abatis; Likhachev, 1956). Where
winter is not severe and almost without snow, for example in the
Trans-Caucasus and England, badgers do not retire at all or its
sleep is repeatedly interrupted, especially by thaws.
1273
845 Fig. 306. Hollow next to root of Mongolian oak—temporary shelter of Amur badger
during the period of fattening on acorns. “Kedrovaya Pad’” preserve, southern
Primor’e. 20 October, 1964. Photograph by A.G. Pankrat’ev.
Upon retiring, the badger blocks the exits of the burrow with
dry leaves and earth.
In spring, the badger emerges from the burrow in March-first
half of April. In Tula abatis, the average date is 10-15 March, with
fluctuation during the multiyear period from 3 to 21-23 March
(Likhachev, 1956); in the neighborhood of Gor’ki (A.N.
Formozov)—the middle of April. The earliest emergence near Kazan
was recorded on 2 April (1955)—the average date—mid-April
846
1274
(Gorshkov, 1965). It has been noted that the time of the badger’s
emergence from the burrow coincides with the spring arrival of the
white wagtail and the beginning of the nuptial flight of woodcock;
however, this coincidence is not always observed. In Middle
Russia, the badger usually emerges after this, when the average
temperature remains for several days above 0°С. The short warm
spells in the beginning of March do not induce emergence of the
badger (Likhachev, 1956).
Seasonal migrations and transgressions. Annually, in autumn
young badgers disperse. Their dispersal is closely connected with
breaking up into pairs and the construction of new burrows. The
animals sometimes occupy burrows the inhabitants of which have
died. The distance to which young badgers dispense depends upon
the food resources of the land. In one case, a pair settled 1.5 km
from the parental burrows (Neal, 1948).
Beside such local mixing, cases occur when individual animals
appear at a considerable distance from the primary habitat. Thus,
in 1951, a badger was found in the upper Pechora, where earlier
it was unknown (Teplov, 1960). Such cases are not frequent, but
are observed everywhere. How badgers are able to penetrate for
such distances during their dispersion and what stimulates them to
do this are not known.
Reproduction. Reproduction of the badger is, at the present
time, much better studied than in the majority of animals having a
latent period of pregnancy. Nevertheless, all data are still contra-
dictory.
In the Moscow zoo, young males copulated at the age of one
year, but it is not known whether these matings are productive
(Osmolovskaya, 1948). In England, badgers have already united in
pairs at the age of one year, but offspring do not occur in the first
year, and the first productive mating is observed at the age of 17
months (Neal, 1948). In the G[erman] D[emocratic] R[epublic]
and Sweden, adult males are able to fertilize females from January
to October, but yearling males, not earlier than in March—April or
May (Shtive, 1952*). Testes of these males are found in active
condition from mid-March to August (other data from Sweden are
contradictory).
Females are capable of reproduction at the age of two years
(Osmolovskaya, 1948). On attaining sexual maturity in the second
*Not in Lit. Cit.—Sci. Ed.
1275
846 Fig. 307. Temporary burrow of Amur badger in oak wood on south slope. “Kedrovaya
Pad’preserve, southern Primor’e. 20 October, 1964. Photograph by A.G. Pankrat’ev.
year, young females first ovulate either early in spring or in sum-
mer and autumn (Neal, 1948). In young strong females, follicles
develop during the period of winter repose (in weaker ones—from
January to May and later; Unger, 1955). In the middle part of the
G[erman] D[emocratic] R[epublic], some of the females born in a
given year are fertilized in September-November, and in the
particularly warm winters, also in December—January. In Sweden,
1276
847 the females of that age are not fertilized (Shtive, 1952). Maturation
of follicles in the ovaries of English badgers was observed
in February—March, as well as mature, recently ovulated follic-
les. Large, maturing follicles were also noticed in June and
September—October (Harrison and Neal, 1956).
Ovulation and formation of new corpora lutea also occur in
pregnant females during the whole latent period, but esterus,
copulation and fertilization do not take place. Therefore, the number
of corpora lutea may be considerably larger than of blastocysts in
the horns of the uterus. During the latent period, not less than
three ovulations take place. Ovulation proceeds within 72 hours
after being provoked by copulation (Notini, 1948). For implanta-
tion of blastocyst, there is no need for the stimulation of copula-
tion. The female allows the male access to her only if it has ready
mature follicles. In middle Sweden, as shown by materials from
several hundred badgers, females are fertilized from May to July.
Copulation in July was also observed in Central Europe, and in
general here, cases of summer mating are known to be quite nu-
merous (Unger, 1955). In England (Neal, 1948), copulation was
observed in March—May, аз well as in June—September, and at-
tempts of it are in February, March and the beginning of
October. The greatest sexual excitement occurs in February—March.
Rut and copulation proceed for short periods, between which the
mutual interest in sex disappears. The least sexual activity occurs
in May and up to mid-July, when all badger activity is concen-
trated on food.
In relation to the ability of the lactating females to be ferti-
lized, data are somewhat ambivalent. According to some data
(Shtive, 1952)*, they cannot be fertilized during this time, and
fertilization occurs in autumn or in the following year. The influ-
ence of lactation on the ovulatory cycle has not been shown
(Harrison and Neal, 1956). In the Moscow zoo, estrus and copu-
lation in Amur and “sand” (steppe) badgers were observed pre-
dominantly in the end of March—April, 3-6 days after parturition,
and in individual cases—in July (Osmolovskaya, 1948). Material
of 120 badgers from France showed that, in the south of
the country, in 80% of the cases copulation takes place during
February, just after parturition (Canivenc, 1957).
*Not in Lit. Cit.—Sci. Ed.
847
848
1277
Fig. 308. Track of a badger, emerging early from hibernation, on deep snow. April.
“Stol’by” preserve near Krasnoyarsk. Photograph G.D. Dul’keit.
In badgers, there are two periods of pregnancy: a prolonged
period of the latent condition of the fertilized egg, and a much
shorter period of active development of the embryos after implan-
tation of the ovum. In the latent period, the fertilized eggs reach
the blastocyst stage and are found free in the horns of the uterus.
Their nutrition at this time is probably fulfilled by the secretory
activity of uterine epithelial cells. In the period March—July, the
blastocyst has a diameter from 1.5 to 2.0 mm—this is the
primitive trophoblast. From October to December, the blastocyst
already has a diameter of 3.5—4.0 mm. Their placentation proceeds
in December, when they are evenly distributed along the uterine
walls (Canivenc, 1957). In England (Harrison and Neal, 1956),
blastocysts were found from 30 April; from placentation to partu-
rition takes 60 days.
In general, the duration of pregnancy the case of summer mat-
ing is 271-284 days, and for early spring—up to one year. In
1278
females fertilized in January and later, the latent period extends to
the end of December, and the duration of pregnancy consists of
14—15 months (Shtive, 1952)*.
With castration and ovary ligation, it was shown that up to
implantation, hormonal secretions of the ovary, including the
corpus luteum hormones, have no effect at all on nutrition and
development of the blastocysts (Unger, 1957). Introduction of
proestrone and progesterone with estradiole did not stimulate
placentation. It is possible that the latent stage is the result of the
absence of the element necessary for this process, perhaps ovarian
hormone. The quick development of the embryos after placentation
is coincident with the lengthening of day light; i.e. with the effect
of cortical hormones (Unger, 1955).
The average number of corpora lutea in material from England
(Harrison and Neal, 1956) during the entire period of pregnancy
was 6.5 (from 2 to 11), the average number of embryos in Janu-
ary—February—3.1; in Sweden, it is equal to 2.5 (Notini, 1948); in
Germany—2.7 (Fisher, 1931).* In England, the average number of
young in a litter is 2.25 (1-5); in 22 cases, it was 2; in 8 cases, it
was 1 and 3, and in 4 cases—S (Neal, 1948). In Moscow zoo, the
number of newborns ranged from 2 to 5 (average—4). In
Kazakhstan, in the Dzhungarsk Alatau, litters were 2—4, in the
lower Ili—3-4, and in Tselinograd district—S—6 (Sludskii, 1953).
On the whole, the size of litters varies from 1 to 6 and the
number of embryos reaches 7, but most often in the litter 2 badger
whelps occur.
In the G[erman] D[emocratic] R[epublic], young are born in
March-April; in Sweden—in March; in Western Europe, from
19 December to 14 April. There was one record in July (Neal,
1948). In southern England, [birth] occurs more often in February—
March and in the north of the country—more often in March. In
the Moscow zoo, in 7 cases parturition occurred in the period
26 March—21 April. Therefore, the reproductive process in badgers
is quite complicated. The major periods depend on geographic
variations in living conditions. These also have, undoubtedly,
significance also for age, and attributes of early or late litter, etc.
In nature, badgers are monogamous, and most often the female
copulates with one male; however, rare cases of mating with two
*Not in Lit. Cit.—Sci. Ed.
849
1279
males have also been observed. The male remains with the female
for several years, possibly for its entire life (Neal, 1948).
Growth, development and molt. In England, the newborn badger
pups weigh about 85 gm, with body length of 12 cm (Neal, 1948).
Weight of newborns in the Moscow zoo was 63-84 gm; on aver-
age 75 gm. They are born completely helpless, blind, and with the
ears covered by skin. The newborn cubs are covered with sparse,
white wool and the dark stripes on the head are well differentiated
At the age of 3-5 days, the claws become pigmented and indi-
vidual dark hairs appear. Strong darkening of color is observed at
the age of 10-14 days. The juvenile pelage is replaced at the age
of 41-68 days. Ear pinnae are formed at the age of 3 weeks, and
eyes open on the 35th—42nd day. Milk teeth erupt at the age of 1
month, but are not fully developed, and at the age of 2.5 months,
their replacement by the permanent ones begins. Reduction of the
milk dentition may be the result of prolonged feeding exclusively
on maternal milk. The young shift to independent feeding at the
age of 3 months (Osmolovskaya, 1948). At the age of 4—6 months,
a reduction in intensive growth is observed.
At the age of 6-9 weeks, badger pups begin to emerge from
the burrow, but they do not decide to abandon it. The first emer-
gence under the mother’s protection was noted in England at the
end of April. At the age of 11 months, they come out near the
burrow to play, but only in the presence of parents. Dispersal of
young badgers proceeds in October, and at that time, they break
up into pairs.
Enemies, diseases, parasites, mortality, competitors and popu-
lation dynamics. The badger has almost no enemies. Wolves, lynx,
and dogs are dangerous to it, but cases of death from them are very
rare. In practice, only human activity has significance for the
badger—direct destruction, and also forcing it from occupied areas
by destruction of burrows. This latter is particularly severe where
possibilities of burrowing are very restricted because of unfavorable
ground and hydrological conditions.
In nature, cases are known of a die-off among badgers, after
which the animals disappeared from the given locality for a series
of years. This points to the effect of an epizootic, but the cause of
the die-off were not established. Only an invasion of helminths
spreading in badgers is known. The most dangerous of these are
pulmonary invasions, in particular eluerostrongylosis and
1280
ef
w
VW
И
ii
849 Fig. 309. Track of right paw of badger on moist sand and the scheme of tracks during
slow movement of the animal. Naurzum preserve, Kazakhstan. 7 April 1957. Sketch
by A.N. Formozov, about 2/3 natural size.
krenozomatosis (Rukhlyadev, 1940). In England, mortality due to
acute tonsillitis was recorded (Neal, 1948). The extent of mortality
among badgers is unknown. The age limit is supposed to be deter-
mined as 12-15 years.
With respect to burrows the badger’s competitors are shown to
be the red fox and raccoon dog. However, the badger is by no
means appears here as the weakest. The old, widespread represen-
tation that the fox itself, being not purely carnivorous, drives out
badgers from burrows, is untrue. Often, fox and badger live side
by side in isolated sections of a wide burrow. However, a series of
cases are known where a badger drove away a fox and destroyed
its litter, discarding the fox carcasses on the surface (Likhachev,
1956).
In England, badgers often die under the wheels of cars and
trains at night, but in the majority of cases, they apparently die in
burrows. This is supported by the frequent discovery of skulls and
bones in burrow scrapings during cleaning. Apparently, badgers
seal the chamber in which the badger died and dig a new one in
another part of the burrow. In England, they observed, as a pair of
1281
badgers pulled a badger corpse from а burrow and buried it in a
rabbit burrow widened for this (Neal, 1948).
There are fluctuations in badger populations, but have not yet
been studied. The greatest influences on them are rendered by
humans and diseases. In England, for example, a considerable
increase in their number was recorded in the war years in connec-
tion with the cessation of its pursuit by humans (Neal, 1948).
Field characteristics. In twilight and night time, the badger
can be detected in the forest by its noise made while moving and
feeding. Under the cover of darkness in distinction from the ma-
850 jority of wild animals, it does not conceal itself, and is heard from
a far. Characteristic signs indicating the presence of the badger are
the beaten tracks going in various directions from its burrow to a
distance of up to 0.5 km, numerous, very typical holes made by it
while catching mouse-like rodents, beetles and their larvae, de-
struction of bumble bees and wasps nests, and also by overturned
forest litter, and feces left here and there.
Tracks of badger on dirt, moist soil or on sand are difficult to
confuse with anything else. They resemble most of all tracks of
bear in miniature. Prints of the plantigrade feet and long, massive
claws are particularly characteristic of them. Most often, prints of
the tracks occur in pairs—prints of hind and fore paws overlap
each other. Length of the forefoot is about 8 cm and of hind—
about 9 cm; length of claws of fore paw in prints is about 2.5 cm,
length of doubled prints of the track, about 10-14 cm (Formozov,
1957).
Practical Significance
In the hunting economy, the badger has little, mostly local, signifi-
cance. The price of fur product from it is not high. The skin is
used for shagreen [leather] and the guard hairs for making shaving
brushes of high quality. Badger meat is eaten in several districts;
however, in the majority of cases, it is thrown away. Badger fat is
highly valued, in Middle Russian districts (Moscow, Vladimir) for
example, and the badger is mainly hunted for its fat rather than its
less valuable skin. The fat is used for greasing leather, but it is
chiefly used in folk medicine in case of injuries, rheumatic dis-
eases and even tuberculosis; i.e., analogous to bear fat. On the
market, it is very scarce (V.G. Heptner).
1282
Not less than 50% of the yearly catch of skins is obtained
from Kazakhstan and other Middle Asian republics of the Union.
Here the badger constitutes about 5% of the commercial fur trade
and in other places considerably less.
At the same time, the badger is useful in destroying insects
and mouse-like rodents harmful to forestry and agriculture. Among
insects and their larvae that are destroyed in great numbers are—
larvae and adults of maybeetles, larvae of mountain leafhoppers,
mole crickets, locusts, larvae of longhorn beetles and sawflies.
Among mouse-like rodents, the common vole has greatest signifi-
cance in its food, and pests of forestry and agriculture. The badger
feeds also on injured animals and birds, and on their carcasses.
Harm caused by the badger to the hunting economy is very insig-
nificant. The accusation of badger attacks on small domestic live-
stock is not confirmed. In England such cases in relation to domestic
fowl were observed, but very rarely.
Methods of hunting the badger vary, except that the badger
does not serve as the object of special hunting. Badgers are caught
in jaw traps, or opportunistically by night hunting with dogs, am-
bushed at burrows with guns, and also caught with special dogs
(fox-terriers and dachshunds) used to excavate the burrow. Finally,
the badgers are sometimes smoked out of their burrows, but more
often, they die in the burrow.
Calculated against the useful activities of the badger, its taking
in the zone of broad-leaf forests and forest-steppe and in the re-
gions of field-protecting forest belts should be entirely prohibited
throughout the whole year, as also should, any kind of excavation
and damaging its burrows, especially where the number of badgers
is determined by the conditions for burrowing. Spring and summer
hunting should be prohibited everywhere as well, except in the
autumn months: from 15 August or 1 September to 1 November (P.Yu.).
851
1283
Subfamily of Otters
Subfamilia Lutrinae Baird, 1857
Genus of Otters
Genus Lutra Brisson, 1762
1762. Lutra. Brisson. Regn. Anim., 13. Mustela lutra Linnaeus.
Dimensions large.
General construction light, body strongly elongated and thin,
extremely flexible; head small, small ear pinnae; tail long, ex-
tremities strongly shortened, plantigrade, claws small, between fin-
gers webbing strongly developed. Pelage very close-fitting, with
coarse guard hairs and very dense, soft underfur. Color almost
constant over whole body, more or less lightening on lower sur-
face, of tawny tone. Glandular area at base of tail absent. Teats 2
or 3 pairs.
Skull relatively light, broad and strongly flattened, with strongly
diverging zygomatic arches and very sharply narrowed postorbital
region. Braincase volume enlarged, rostral region short and broad—
its width greater than length. Posterior part of hard palate lying
posterior to toothrow narrow and relatively short. Mastoid proc-
esses flattened, approaching anteroventral part of occipital crest,
paroccipital processes low but separated from auditory bulla.
Mastoid width of skull a little less than zygomatic. Auditory bullae
flattened, with small bony auditory meatus.
Dental formula 15 Co PIM =36 In its main features, the
dentition is like that of the badger subfamily, Melinae. It also
represents the same combination of cutting and tubercular (grind-
ing, crushing) type, with however, great development of incisor
characters. Compared to the general dimensions of the rostral part
of the skull and of the palate, the dentition is large. Upper carnas-
sial tooth large and triangular in form. Upper molar rhomboidal in
form, its area approximately equal to carnassial or very slightly
less. Lower carnassial tooth long, anterior part has three well-
developed apices; posterior part a little longer than anterior. Loss
of anterior premolars usually does not occur.
853
851
1284
Specialized freshwater forms, excellent swimmers and divers,
always connected biotypically with water bodies, both in plains
and in part, mountains. Swims both by paddling movement of the
limbs and by sinuous bending of extremely flexible trunk and tail.
Feeds mainly on fish, in part on aquatic invertebrate (crayfish) and
amphibians. Builds bank burrows. Hibernation is absent, appar-
ently, as is a latent stage in development of the fertilized egg.
The range of the genus is extremely extensive—one of the
very largest among mammals. It is exceeded only by the ranges of
several bats. This is particularly remarkable, given the great spe-
cialization of species in the genus and the relatively small number
of them (see below). The genus is distributed throughout Europe,
occupies the mainland of Asia (except the extreme north and
Arabian Peninsula), Ceylon, Sakhalin, the Japanese islands, Tai-
wan, Hainan, Java, Sumatra, Calimantan (Borneo) and Banka. Rep-
resentatives of the genus are absent on the remaining islands of the
Pacific Осеап'.
In Africa, the range includes the extreme northwest of the
continent (Atlas countries) and the entire continent south of the
Sahara, except its extreme southern and southwestern parts. In
North America, the range occupies the whole continent, except
several areas in the extreme north and desert regions in the south,
the islands adjacent to the continent along its western shore (Kodiak,
Alexander archipelago; absent on islands of the Bering sea), and
islands off the northeastern shore (Newfoundland and some smaller
ones). In the range is included Central America and South America,
and the whole of Tierra del Fuego. In the range is included the
island of Trinidad, but species of this genus are absent on all
islands of the West Indies. The range of the genus, therefore,
occupies almost the entire range of the subfamily—its species are
absent only on Palawan, Commander, Aleutian and Pribiloff is-
lands, in middle India and in South Africa, where species of other
genera are encountered.
The genus Lutra is the very largest in the subfamily. In it are
11 species of the 16 assigned to Lutrinae. There were attempts
to divide it into several independent genera (Lutrogale for
‘Information on the Kuril islands is incorrect (see below, in description of
otter). Pocock’s reference to its occurrence in the Philippine Islands is, apparently,
wrong—it was not confirmed by anyone. In the Philippine sensu stricto, true
otters are absent, but the clawless otter, Aonyx cinerea, exists on Palawan.
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L. perspicillata, or Hydrictis for L. maculicollis); however, the
majority of authors do not accept this division, and it is, appar-
ently, more correct to recognize one genus with two subgenera—
Lutra and Lutrogale. On the other hand, some authors (Chasen,
1940; Carter, Hill and Tate, 1946) believe that the separation of
the genera (subgenera) of the so-called clawless otters Aonyx
(capensis) and Amblonyx (= Microaonyx: cinerea) has no basis.
They combine the species mentioned with Lutra, which is an hy-
pothesis that demands a wider foundation.
Besides the most widely distributed Palearctic and Indo-
Malayan species, the common otter, L. lutra, the genus also in-
cludes the following species: L.* canadensis (North America, south-
ward to Мех!со)?; L. sumatrana (Indo-China and Malayan
archipelago; evidently, the least specialized species); L. maculicollis
(Africa south of the Sahara); L.* annectens (extreme south of North
[America], Central and South America); L.* enudris (South
America); L.* felina (southern South America; a marine form,
evidently the most specialized form in the genus); L.* incarum
(South America); L.* platensis and L.* provocax (the same;
subgenus Lutra) and L. perspicillata (South and Southeast Asia;
subgenus Lutrogale). Therefore, the greatest richness and diversity
of species are obviously, characteristic of South America (the center
of origin)—there are 6 species out of 11 here. North America is
characterized by 1 species, there are 2 species in South Asia (one
represents a separate subgenus), 1 species in Africa and 1 species
has the above-mentioned wide distribution in Eurasia and north-
western Africa. It is noteworthy that with the great diversity of
otters in South America (besides those mentioned, there is also the
giant river otter, Pteronura brasiliensis). North America possesses
only one endemic species. One is characteristic also for northern
Eurasia—apparently the diversity of forms in the genus is in the
south. In this sense, geographic variation of L. lutra is also indica-
tive (see below).
As already mentioned (see characters of the family), in this
complicated mix of forms representing the marten family,
*These New World species, here placed in Lutra, are now considered a distinct
genus, Lontra—Sci. Ed.
This form, with its great general similarity to the Eurasian otter, itself represents
a separate species, well characterized craniologically. The separate species,
L. mitra (Prince Wales Island in Alexander archipelago), accepted by American
authors, represents only a subspecies of L. canadensis.
854
1287
Mustelidae (perhaps itself a polyphyletic group), the subfamily of
the otters—Lutrinae—is one of the most accurately circumscribed.
Perhaps it may even be considered the most natural and sharply
distinguished one.
Besides the genus Latra, the subfamily includes 4 other gen-
era, the so-called giant river otters: Pteronura, in South America
(1 species—P. brasiliensis), the clawless otters, Aonyx—a genus
comprising two species belonging to two subgenera—A. (Aonyx)
capensis in Africa and A. (Amblonyx) cinerea in Southeast Asia,
Paraonyx—with 1 species in West and Central Africa (P. congica:
“species” microdon and philippsi are not taken into consideration
here) and sea otter* (kalan)—Enhydra, with 1 species (Е. lutris)
from the shores of the northern part of the Pacific Ocean (see
below).
The identification of the position of Lutra within this circle of
genera is very difficult. The thing is that within the limits of the
subfamily, the genus of sea otters (“kalan”)—Enhydra—is totally
sharply set apart from all the genera of the subfamily by the ex-
tremely unique structure of its limbs, skull, completely distinctive
teeth and other characters (see description of this form below). All
the remaining genera (and species), together and separately, are
very far different from Enhydra and are, to a greater or lesser
degree, close to each other. Differences between them are com-
pletely unequal to differences between each of them and Enhydra.
By this criterion, they might perhaps be considered as one genus;
so to speak, river otters, Lutra; corresponding to the genus of sea
otters, Enhydra.
Characters of external structure and craniology, by which the
otters are usually grouped, create patchwork combinations. Thus,
the group Aonyx has no claws (A. capensis) or they are very small
(A. cinerea). This [latter] is usually considered a specialized char-
acter (“progressive”). At the same time, finger webbing is reduced
or actually almost absent as in the posterior limbs of A. capensis.
Their skulls are relatively short with a voluminous braincase. The
dentition is modified only weakly and unsubstantially. As regards
the structure of its skull, extremely developed, large finger web-
bing and certain other characteristics, evidently the entire genus
Lutra must be considered a highly specialized (“progressive”) group
*In Russian original, “Kamchatka beaver”’—Sci. Ed.
1288
among the genera of freshwater otters. Most closely related to it,
apparently, is the genus of giant Brazilian otters Pteronura. The
southern Asiatic and African Aonyx and Paraonyx are, apparently,
more primitive, although Aonyx is also very close to Lutra.
In its origin, the subfamily is apparently connected to certain
extent with the subfamily of badgers, Melinae. However, the ear-
liest representatives of the subfamily as highly specialized forms
are already known from the Upper Oligocene of Europe
(Potamotherium). More than 10 genera of fossil otters have been
described, mainly from the Upper Tertiary period (Miocene,
Pliocene—Paralutra, Enhydriodon, Vishnuonyx, Sivaonyx and
others) and partly from the Pleistocene (Cyrnaonyx). The genus
Lutra is known from the Lower Pliocene of Europe and Asia and
from the Upper Pliocene of America.
All the species of the family, even in the tropics, represent
valuable fur animals.
In the USSR, there is one species: Lutra (Lutra) lutra Linnaeus,
1758, which represents about 0.3% of the species of the fauna and
about 10% of the species in the genus.
The range occupies almost the whole territory of the country.
The species is everywhere closely associated with water.
A valuable fur-bearing animal (V.H.).
Subgenus of the Common Otters
Subgenus Lutra Brisson, 1762
[EURASIAN] OTTER, PORESHNYA
Lutra (Lutra) Linnaeus, 1758
1758. Mustela lutra. Linnaeus. Systema Naturae, ed. X, 1, p. 45.
Uppsala, Sweden.
1777. Lutra vulgaris. Erxleben. Regn. Anim., 1, p. 448. Renaming
of lutra.
1834. Lutra nudipes. Melchior. Den Danske Stats og Norges
Pattedyr, p. 50. Coast of northern Norway.
855 1839. Lutra aurobrunnea. Hodgson. J. As. Soc. Bengal, 8, p. 320.
Nepal.
1289
1839. Гита monticola. Hodgson. Journ. As. Soc. Bengal, 8, р.
320. Nepal.
1844. Lutra kutab. Schinz. Syn. Mamm., p. 354. Kashmir.
1867. Lutronectes whiteleyi. Gray. Proc. Zool. Soc. London, p.
181. Japan.
1912. Lutra lutra seistanica Birula. Ezhegodn. Zool. mazeya Ak.
nauk, 17, p. 274. Gil’mend river in Seistan, eastern Iran.
1915. Lutra lutra oxiana. Birula. Ezhegodn. Zool. muzeya Ak.
nauk, 19, p. XXI. Lyangar, Pyandzh basin. Western Pamir.
1922. Lutra vulgaris var. baicalensis. Dubowski. Arch. Tow. Nauk.
Lwow, 1, p. 349. Nom. nud. Near Baikal.
1922. Lutra vulgaris var. amurensis Dybowski. Ibidem, p. 349.
Nom. nud. Amur and Ussuri Territories.
1922. Lutra vulgaris var. kamtschatica. Dybowski. Ibidem, p. 349.
Nom. nud. Kamchatka.
1931. Lutra lutra meridionalis. Ognev. Zveri Vost. Evropy i Sev.
Azii, 2, p. 527. Vic. of Teheran, Iran.
1936. Lutra stejnegeri. Goldman. Journ. Mammal., 17, p. 164. Vic.
of Petropavlovsk, Kamchatka (У.Н.).
Diagnosis
The only species of the genus in the USSR.
Description
Body strongly elongated and relatively thin, more or less equally
thick along its whole length—posterior part only slightly more
massive than shoulder region. Limbs short, from whence its stat-
ure. Tail long and comprises about half the length of head and
body, very thick at base, strongly narrowing to tip and flattened in
dorso-ventral direction. Such tail structure is determined by strong
development of ligaments but most of all by the powerfully mus-
cular tail tracts running along the vertebrae. Moreover, masses of
collagenous tissue are strongly developed along the lateral sides of
the tail, somewhat less on the lower side, and still less on the
upper side, forming loci for fat deposition. Tail thickness, there-
fore, depends to a certain degree on nutrition of the animal, and
characterizes it. The tail, contrary to all remaining species of the
856
1290
family (except sea otter, does not bear long hairs. It is covered by
short (as on the trunk or shorter) closely investing fur.
Head relatively small, elongated oval, narrow, no wider or
hardly wider than the neck, strongly flattened. Neck relatively short.
Ears short, with rounded apexes, weakly protruding from fur, widely
separated. External ear openings covered by special valves. Eyes
quite large; bare area on tip of nose large, transversely elongated—
its width greater than its height or approximately equals to it.
On the whole, the appearance of the otter’s head is somewhat
snake-like.
Digits fused together by strongly developed webs, reaching
middle or nearly to end of distal phalanx. Digits, including first,
somewhat elongated, which increases area of hand and foot.
Second and third digits on both fore and hind limbs are longest,
and equal in length. Lower hand and foot naked, edged with elastic
hairs somewhat increasing propulsion surface. Claws small, but
relatively little reduced.
The otter body is extremely flexible and the animal is very
active. On land, the otter moves in a walk or trot, with lowered
head, bent back, and outstretched tail usually dragging its end on
the ground; more rarely leaps, strongly hunching [its back]. On the
whole, its movement on land is not very agile. Movements of otter
in water are particularly confident and agile. Swimming slowly, it
usually paddles with its paws but when moving quickly, it presses
limbs against body and moves with energetic snake-like move-
ments of the whole trunk and muscular tail. The moveability and
flexibility of the vertebrae and entire body of the animal are
exceptional. Overall, in general appearance and habits, the otter is
entirely unique and is strongly differentiated from all remaining
species of the families of our fauna.
Pelage in winter is short, very even over the whole body (dif-
ference between length of hair on back and belly is small), close-
fitting, shiny, with often coarse, elastic guard and very dense and
close-fitting soft, silky underfur. The fur on the belly is denser
than on the back. In northern otters, the average length of contour
hairs on the back averages 24.2 mm, on belly 21.0 mm; guard
hairs, correspondingly, are 18.4 and 17.2 mm, and underfur 14.6
and 11.2 mm. The number of hair on 1 сп? of dorsum reaches
35,000 (34972), on venter about 50,000 (50668). On the back, for
every outer hair, an average of 155 underhairs occur and on the
856
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АНЯ
МЕРЕ
4.
I и, Й,
ng Fg:
Bae
cated,
Fig. 311. Otter, Lutra lutra L. Sketch by A.N. Komarov.
belly—120. Outer hairs have very wide and flattened distal thirds—
the ratio of their width to length is 1:3-3.5 (in terrestrial forms,
not more than 1:1.5—2.5). Moreover, the outer hairs are strongly
inclined with respect to the skin surface (30-35°) and are strongly
bent near the base (“in the neck”) where they are relatively thin.
On the whole, the outer hair has the form of a small plate and
together they tightly cover over the underhairs. The bases of these
latter, as well as bases of the smallest top hairs, are strongly crimped
and form a wavy fur coat. All these characters, especially the struc-
ture of the outer hairs, themselves represent typical adaptations to
an aquatic mode of life, and they prevent wetting of the fur, even
during extended immersion in water. The cortical layer of the hair
857
1292
is greatly increased, thus its durability greatly augmented
(Tserevitimov, 1958)’.
Full growth of the otter is attained only in the third year of
life; at that time, its fur is also completely developed. Skins of
animals born in the current year (“КозШаК”) have a long and soft
pelage composed of sparse underfur and thin, soft guard hairs. In
second-summer animals, the fur is longer and softer than in adults,
but coarser than those born in the current year, with sparse underfur.
Glandular areas at the tail base are absent; anal glands are
present, but not strongly developed (?). Teats 2-3 pairs’.
Color of the upper body in winter fur is bright dark-brown or
dark nut-brown. Upper head somewhat darker, tail of the same
color as back. Lower surface of body, including ventral neck and
lower and lateral head are slightly lighter, with tinge of silvery
yellowish tones, sometimes whitish. On the sides, the color forms
a gradual transition from the tone of the back to the color of the
belly. Legs are dark brown, and claws light horn, darker at the base.
In summer coat, the fur is somewhat more sparse, but of the
same color as in winter. There are no differences among sexes in
color. Individual variability of fur is insignificant and lies only in
more or less intensity, and in greater or lesser lightening of venter
relative to dorsum. In some localities, two color types are differ-
entiated—lighter and darker (G.F. Bromlei; southern Ussuri Terri-
tory); however, color dimorphism does not оссиг.5 This is normal,
simple individual deviation, apparently, even mostly age variation.
The soft “fluffy” fur of the young otter is dark-chocolate
color and lacks luster. At first winter, the young animals don a
coat similar in color to that of the adult, but their venter is darker
than in adults. Upon attaining the adult condition, color continues
to change somewhat, and later color changes in the direction of
3Otter fur is considered very durable and “long-wearing” and in determining the
relative quality of other furs, its “long-wearing” is accepted as 100. In mink, it equals
70, in sable 50 and in squirrel 25 etc.
“For otters of England, 3 pairs recorded (inguinal; Miller, 1912), for middle
European 2-3 pairs (Gaffrey, 1961), for our otters, 2 pairs (Ognev, 1931), for the
Indian form of the species, 2 pairs (Pocock, 1941).
‘It is considered that the dark otters, called “river”, have a uniform dark-brown
color and small dimensions, and live in rivers. The other form—‘sea” otter—is larger,
its color is lighter—with a dirty-ocherous belly and legs, lives and feeds in the sea
(С.Е. Bromlei). Similarly, two types of color were also noted for otters in the upper
Amu-Darya basin (Chernyshev, 1958).
859
1293
lightening of the general fur tone, while the venter becomes lighter
relative to the dorsum.
Geographic variations of color within the USSR are insignifi-
cant and are only expressed in lightening of the general color tone,
relatively greater lightening of venter, and decrease in the fur density
in some southern regions.
The skull is very broad, both in rostral, and particularly in the
zygomatic and cranial regions, and is strongly flattened. Height of
the braincase in its middle section constitutes only slightly more
than half the mastoid width. Height in the region of the auditory
bullae is less than mastoid width. The upper line of the skull pro-
file is almost straight—it is only insignificantly depressed in front
of the middle of the interorbital area; and in the young—also in the
occipital region. On the whole, this line is almost parallel to the
line of the lower surface of the skull. Such a flattened skull is
absent in any other carnivore of our country.
The braincase is very voluminous and broad in its middle and
posterior parts. The width of this region is increased still more by
strong development of the occipital crest which extends also onto
the lateral surface of the skull, and the mastoid processes, which
are strongly flattened and fused with them in its anterior part. The
anterior region of the braincase (in the interzygomatic area) is
sharply narrowed. The postorbital area is sharply narrower and
forms a strong intersection, particularly striking to the eye thanks
to the wide separation of the zygomatic arches. Width of the
postorbital constriction is considerably less than the width of the
interorbital area and is approximately 4 times less than mastoid
width. Postorbital processes are well developed, but short and blunt,
and border only a very small part of the orbit.
Orbit quite large—its greatest diameter almost equal to inter-
orbital width. Zygomatic arches are strongly separated laterally
throughout their whole extent (as well as in anterior part) and
generally form a rounded figure. They are strong and massive,
noticeably curved upwards, and in the anterior part bear a well
defined tubercle on their upper edge, bordering the orbit from
behind and below. Infraorbital foramena, perforating base of
zygomatic arch anteriorly, very large. Its diameter is 1.5—2 times
more than diameter of the cusp of the upper canine.
Rostral part of skull very short (about 2 times shorter than
cranial), high and broad. Distance from anterior point of zygomatic
1294
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858 Fig. 312. Skull of otter, Lutra lutra L.
arch to anterior edge of alveoli of incisors approximately equal to
width of muzzle below canines, and approximately 4.5 times less
than condylobasal length of the skull. Sagittal crest is almost un-
defined or only weakly so. Palate between toothrows relatively
quite narrow and elongated—distance between inner parts of
859
860
1295
ata XE
Fig. 313. Skull of a very old otter, Lutra lutra L. Posterior cheek teeth only remain,
worn to gums. Sudzukhinsk preserve, southern Primorye. Sketch by G.F. Bromlei.
molars fits into length of palate from anterior edges of alveoli of
incisors to posterior edge of molars about three times. Portion of
palate lying behind the toothrow is wide anteriorly, quickly nar-
rows posteriorly, and is short. Its length is approximately equal to
distance between canines. Interpterygoid notch narrower anteriorly
than posteriorly.
Auditory bullae small, flattened and only insignificantly swol-
len along inner part. Paroccipital processes have form of short
rounded outgrowths and do not contact auditory bullae. The exter-
nal auditory meatus is short, opens below mastoid (anterior) part
of occipital crest, and are directed forward and upward. The lower
jaw massive compared to general lightness of skull. Coronoid part
short, but quite wide, its apex not pointed.
Compared to general dimensions and width of palate and
rostrum, teeth are large—maximum transverse diameter of upper
carnassial tooth constitutes half of distance between carnassial teeth.
Canines are thin, long and very sharp. All small premolars are well
developed and of tearing form, and by their location in relation to
teeth of opposite jaw (upper—lower), form a strong tearing appara-
tus. First upper premolar well-developed and not reduced. It is
displaced deep into the toothrow and does not lie directly behind
the canine, but on its intero-posterior side.
Upper carnassial tooth large, quite regularly triangular in form,
one angle directed inward. It bears a high cutting apex along outer
edge, behind which lies a second apex, shorter and connected with
first by a crest. In front of the main apex lie small teeth. All of
them are arranged regularly next to one another. The inner blade,
1296
the area of which constitutes about half the area of the tooth, is
flat and does not bear apexes.
Upper molar large, rhomboidal in form. Its area is approxi-
mately equal to area of carnassial tooth or slight smaller, and its
greatest diameter is directed almost at a right angle towards the
axis of the toothrow. The flat surface of the tooth bears four small
tubercles, of which two are along the outer edge. Lower carnassial
tooth elongated in form, with three well-developed sharp apices,
almost identical in dimensions on the anterior part of the tooth.
Posterior half of tooth somewhat larger than anterior one and does
not bear sharp apices. In area, second lower molar significantly
smaller—approximately three times—than carnassial.
Sexual dimorphism of the skull is well manifested. Beside some-
what smaller general size, the female skull is distinguished by
almost complete absence of a sagittal crest, even in old individu-
als, somewhat smaller infraorbital foramen, narrower rostral part
of skull (breadth above canines), somewhat smaller nasal opening,
and, apparently, somewhat narrower and shorter interpterygoid
notch. Skulls of young animals have less-developed protuberances,
crests, etc., and are less angular; nasal region is relatively low,
lower than occipital region (with age, massiveness and height of
anterior part of skull increases), postorbital region is broader and the
supraorbital processes are small (Ognev, 1931). Sagittal crest does not
develop and occipital is very small. Geographic variation of the skull
within the USSR was not noted, and if expressed, then very weakly.
Posterior section of os penis is thickened and somewhat el-
evated. On anterior end has two rounded lobes, tips directed down-
ward separated by groove. Left lobe 1.5 times thicker than right
and ventrally bears incision (Ognev, 1931).
Ratio of body length to length of intestine (two males) 1:6.36
and 1:6.10; cardiac index (male, January) 13.8 (from data of
Chernyshev, 1958; Pyandzh basin).
Body length is 550-950 mm, length of tail 260-550 mm, hind
foot length 110-200 mm, ear length 17-25 mm.
Condylobasal length of male skull is 107.0-126.0 mm, of fe-
males 96.5—121.0 mm; zygomatic width of males 1$ 62.0-83.0 mm,
of females 60.5-72.0 mm; interorbital width of males is 17.5-22.5
mm, of females 16.2—21.3 mm; postorbital width of males is 12.3—
18.3 mm, of females 11.8—18.3 mm; mastoid width of males is
54.5-71.3 mm, of females 55.9-67 mm.
861
1297
Weight of adults 5.7-10 kg.°
Females, as is evident from the craniological indices presented,
are noticeably smaller than males. Differences in body dimensions
and weight of individuals from one restricted territory (Sudzukhinsk
preserve in southern Ussuri Territory) are as follows (15 speci-
mens): body length of males; 550-810 mm, of females 550-680
mm; tail length of males; 280-455 mm, of females 280-380 mm;
hind foot length of males, 110-120 mm, of females 82-93 mm; ear
length of males, 19-25 mm, of females 17-22 mm. Maximum
weight of males is 9470 gm, of females, 6543 gm. Animal in the
first year of life, in November—February, have a body length of
about 350 mm and weight from 1100-1500 gm (G.F. Bromlei).
The latter figures are relatively small—young otters from Pechora
(preserve; Z[oological] M[useum of] M[oscow] U[niversity]) in
November-—February weighed 2250, 2400 and 4430 gm (males)
and 2250 and 3500 gm (females).
Very old individuals approaching the age-limit experience
poorer nutrition than those of intermediate age and, despite some-
times large dimensions, weigh less (G.F. Bromlei). Nutrition and
weight change seasonally.
Length of os penis is 58-65 mm.
Geographic variation of dimensions within the boundaries of
the USSR was not noted, but some, apparently, exists, and south-
ern otters are somewhat smaller than northern. Several forms from
the extreme south of Asia (nair, barang) are quite strongly differ-
entiated by their dimensions (V.H.).
Geographic Distribution
In Europe, Asia except the extreme north and Arabian Peninsula,
and in northwestern Africa.
‘Data on body dimensions and weight from Ognev, 1931; Novikov, 1956;
Stroganov, 1962; G.F. Bromlei and from materials of the Z[oological] M[useum of]
M[oscow] U[niversity]. Skull dimensions from Ognev, 1931; Stroganov, 1962 and
collections of Z[oological] M[useum of] M[oscow] U[niversity]. Altogether approxi-
mately 150 skulls. Reference to body length of 100 cm and weight of 15 (16) kg
(Central Europe; Gaffrey, 1961 and several other sources) is apparently an exaggera-
tion. In any event, such measurements and weight were not recorded in our otters,
even the very largest populations. However, one case is known of an otter caught in
Turkhan Territory with tail length (measured from skin) of 175 cm (Zalesskii, 1930).
It is possible that this is a case of gigantism, analogous to kolonok and polecat (see
[above]).
1298
Geographic Range т the Soviet Union
Very vast and constitutes the greater part of the range of the spe-
cies. It occupies almost the entire territory of the state except the
extreme north and arid and desert regions.
In the European part of the USSR, the northern border of the
range on the west is formed by the northern coast of the Kola
Peninsula (also exists on Kil’din [Island]) and eastward along the
White Sea coast as far as the mouths of the Kuloi and Mezen’
[rivers]. On the Kanin, the otter as a permanent inhabitant is
absent, but it is sometimes transient there. Eastwards to the Pechora
delta, the border passes along the seashore, but beyond the Pechora,
somewhat lowers and turns to the south, and reaches the
Urals somewhat farther from the sea. However, here also there are
places of transgression into the tundra (Malozemel’sk and
Bol’shezemel’sk; V.Ya. Parovshchikov). In the Urals, the border
falls approximately along the Arctic circle (Shvarts, Pavlinin and
Danilov, 1951).
In the northwest of western Siberia, the border passes some-
what to the north of Salekhard and along the extreme southern
edge of Yamal (upper Yana river). Going to the east, it crosses
Ob’ gulf at its southernmost part and passes to the eastern shore
of the Ob’ gulf at the Arctic circle (Laptev, 1958). Farther, at the
same latitude, it crosses the lower Pur and passes beyond the Taz
approximately 100 km north of Sidorovsk on the Arctic circle and
at the same level or a bit south of the upper Turkhan. Thence, the
border suddenly rises to the north, crosses the Yenisei, and east of
the Yenisei, Lomo Lake (at Norilsk) and on to the southern Taimyr,
reaching there the upper Avam (right tributary of upper Pyasina)
at a point a little north of 70° N. lat. It also includes the Kheta
(V.N. Skalon), apparently, in its upper and middle course. The
Avam is the most northernmost point of the range in the USSR.
The otter is absent along the lower Khatanga (Middendorf, 1867).
There is a reference to otter hunting in West Siberia, on the
Gydansk Peninsula in the upper Yuribei about 71° N. lat., in the
northern part of the tundra zone (Laptev, 1958; Stroganov, 1962).
This information is very doubtful, and if it is true, speaks only to
occasional long transgressions.
The eastern border is very poorly known; however, it can be
considered established that in Yakutiya (especially in the west and
863
1299
middle) bends considerably towards the south and then again rises
strongly to the north in northeastern Siberia. This is, apparently,
connected with particularly low winter temperatures in this part of
Siberia, leading to formation of very robust ice cover, to freezing
of many water bodies to the bottom and even to freezing of small
montane creeks.
The course of the border in the eastern half of Siberia is drawn
in the following form: from southern Taimyr, the border in a way
not yet completely established, goes to the southeast to the upper
УПуш in the region of Chona mouth (about 62°45” [N. lat.]. Then,
directing itself towards the east, it crosses the Ygyatta (Igetta)
river and several other left tributaries of the Vilyui (Tyung, Tyukan)
and passes to the lower Vilyui below Vilyuisk. Thence, the border
passes generally in the same direction, crosses the Lena somewhat
below the mouth of the Vilyui, extends through the middle courses
of the right tributaries of the Aldan—the Tumara and Tompo and
passes to the sources of the Indigirka and upper Oimyakon. Then,
it goes along the left tributary of the Kolyma—the Yasachnaya
river, approximately at Sredne-Kolymsk and to the right tributaries
of the Kolyma—the Berezovka and Omolon (border in Yakut ac-
cording to data of Maak, 1859; Tugarinov, Smirnov and Ivanov,
1934; Belyk, 1953 and Ognev, 1926, and mainly from material
made available by О. Ivanov, Yakutsk).
Along the described northern border (not considering the
Omolon), the otter is very rare and only individual cases where the
animal or its tracks are encountered are known. The border of
normal, regular occurrence lies to the south. Nevertheless, there
are some less definitive data (Maak, 1859) on occurrence (very
rare) of otter along the left tributary of the Lena—the Muna—
which flows into the Lena from the left at 68° N. lat. and, a little
to the south—on the “Serka” river (apparently the Syarga) in the
upper Linda (approximately on the Arctic circle west of the Lena)
and at Zhigansk on the Lena—a bit north of the Arctic circle.
These data are in poor agreement with contemporary materials
and invite doubt. It is possible that they refer to occasional indi-
viduals transgressing far along the Lena. Mentions of “Alazeya in
Middle Koymsk region” (Belyk, 1953) obviously belongs in this
category. The same may be said of the upper Alazeya (Stroganov,
1962, plotted this occurrence on the map very far to the north).
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In the lower Kolyma and along the Bol’shoi and Malyi Anyui
it is, apparently, absent. Farther to the east, the border includes on
the north the Anadyr basin, and the range also includes its left
tributaries—Belaya, Tanyurer and Konchalan (Belopol’skii, 1937;
Portenko, 1941). At the mouth of the Anadyr’, the border passes
to the Pacific Ocean.
The range includes Sakhalin and Shantar Islands (at least
Bol’shoi Shantar; Ognev, 1929). There are no data concerning
Karagin Island. Of the Kurils, the otter was recorded for
Kharimkotan (Kharumukotan) in the north and Iturup (Etorofu)
and Kunashir’ (Kunasir’) in the south (Sergeev, 1947).
On the whole, data on occurrence of otter in the Kuril chain
are confused and contradictory. Several authors, both geographers
(Solov’ev, 1946) and zoologists (Kuznetsov, 1949) do not refer to
this species at all in the composition of the fauna, and according
to new data (V.G. Уогопоу) it 1s actually absent there. The reason
for mention of the otter was probably its confusion with the
Kamchatka beaver (sea otter—Enhydra lutris) and old information
of Snou (1902) who recorded for Iturup and Kunashir the “абип-
dance” there of “river beaver, Lutronectes whiteleyi’*.
The southern border of the otter’s range in the European part
of the country, starting from the mouth of the Danube, generally
passes along the Black Sea coast to the mouth of the Dnepr, and
farther, around the extreme south of the steppe to the mouth of the
Don. In the Crimea, the otter is absent. In the past the otter lived,
apparently, in small creeks in the southern steppes. Farther, the
border of the range extends along the Don and somewhere in the
region where the Don and Volga come close, passes to the Volga.
Along it, the otter spreads to the very delta.
To the south of the Don and Volga, in the Trans-Don, Kalmytsk
and a considerable part of the Cis-Caucasian steppes the otter is
absent. It is encountered along the Kuban’, Kuma and Terek (no
data concerning Manych), and the line extending along the Kuban’
and Kuma represents the northern border of the Caucasian part of
the range. To the south of it, the otter exists throughout the entire
Caucasus. The Caucasian part of the range, therefore, is isolated
from the northern. The possibility, and even the probability, exists
that, in the past, the otter lived along the rivers flowing into the
*This “species” is a synonym of L. lutra—Sci. Ed.
864
1302
Azov Sea between the Don and Kuban’, and thus isolation by the
steppes was not so complete. It may also be assumed that, under
certain conditions, contact between populations might have
occurred between the Kuma and Volga.
The otter is absent in the steppes between the lower Volga and
Ural [rivers] and in Saratov Trans-Volga. From the Volga to the
Ural river, the southern border passes, apparently, somewhere along
the latitude of Irgizi. The otter extends along the Ural [river] to the
[Caspian] Sea.
From the mouth of the Ural, the border rises northward along
the river and then turns to the east, including also the lower Ilek.
The farther course of the border is entirely unclear and may be
designated only tentatively. It bends around the Urals [mountains]
along the Ural river and, after rising somewhat to the north, it
passes somewhere along the border between the forest and forest-
steppe zone, or along the forest steppe eastward to the Irtysh, and
along it upstream to its sources (recorded at Ust’-Kamenogorsk
and Chernyi Irtysh; Sludskii, 1953). It cannot be excluded that in
the cut-off section mentioned, before the Irtysh, the line of the
border passes somewhat more to the south—there is information
on the occurrence of otter to the southwest of Petropavlovsk and
to the west of Kokchetav (Sludskii, 1953), and, perhaps, even in
the upper Ubogan river at Kushmurun Lake to the southeast of
Kustanai (V.G. Heptner).
Beyond the Irtysh, the otter is absent in the Barabinsk,
Kulundinsk and Cis-Altai steppes. In this zone, the border of the
range, proceeding somewhat to the north of Lake Chana, reaches
Ob’ south of Novosibirsk, then goes directly south and, passing
around the Altai from north [to east], reaches the Irtysh.
From the Zaisan depression and southern Altai to the Pacific
Ocean, the range passes to the south outside the boundary of the
USSR.
The range of the otter in Middle Asia is very narrow and
complex. Moreover, its distribution is sporadic and it is every-
where rare. Locally, the otter is encountered only as a transient
and has been observed or caught once in several years. In some
regions where it occurred in the past, this animal is absent at the
present time. Finally, data for Middle Asia are in general very
scarce, fragmentary and in part indefinite. Therefore, it is almost
865
1303
impossible to outline a more or less accurate range in this region
at the present time.
The Middle Asian range of the otter must be considered as the
northern edge of the Near Eastern part of the general range of the
species, and in part, perhaps, the Central Asian (Pamir-Altai and
Tien Shan, and Semirech’e). There is no direct connection with the
main European-Siberian region of occurrence of the species in the
USSR (between the Dzhungarsk Alatau and Chernyi Irtysh otter
are absent), or-it is weak and may exist in some form in the east.
This is demonstrated by the systematic peculiarities of the northern
and southern otters (see below).
In the west, a small section of otter occurrence is occupied by
the Kopet-Dag [mountains] and Atrek [river] to its mouth. This is
the northern edge of the Iranian part of the range, included within
the limits of the USSR from the south. Another analogous section,
isolated in our country from the first, is situated along the Tedzhen
river, occupying its upper and middle courses and not reaching
Tedzhen city. A third section is occupied by the Kushku river,
where, as possibly in some sections of the Tedzhen, the otter ap-
parently occurs only as a transient from the south, and extremely
rarely (V.G. Heptner). Concerning the occurrence of otter on the
Murgab, at the present time or in the past, positive data are absent,
though it is not excluded, at least for its upper course.
Farther to the east, the section inhabited by the otter is, obvi-
ously, connected with Afghanistan, occupies the western Pamir;
i.e., the region of rivers flowing from the east to the upper Pyandzh
(Rozanov, 1935), Pyandzh [itself] and the uppermost Amu-Dar’ ya
and the basins of the rivers flowing into it from the right, in par-
ticular the Surkhandar’ya, Kafirnigan, Vakhsh-Surkhab, Kizyl-su
and smaller rivers—Varzob, Khanak’, Ilyak (Flerov, 1935;
Chernyshev, 1958; V.G. Heptner). In the basin of Surkhandar’ ya,
the range includes the regions of the Tupalang and Sangardak
rivers (Ishunin, 1962; V.G. Heptner); i.e. the southwestern part of
the Gissar range. Along the Amur-Dar’ya, the range does not reach
beyond Termez. Concerning the occurrence of the otter in the
remaining part of the Amu-Dar’ya, data are absent, and apparently
it does not exist along this river. The extreme turbidity of water
probably prevents its occurrence.
The otter is absent in the eastern Pamir. Concerning all re-
maining parts of the Pamir-Alaisk montane system with its water
1304
bodies, we have no definite data on the occurrence of otter, ex-
cept one reference to its occurrence on the Zeravshan below and
somewhat above Samarkand (Fedchenko, 1950; Ishunin, 1961).
In Semirech’e, in the Tien Shan system and in the Syr-Dar’ya
basin, the range of the otter occupies Dzhungarsk Altau and the
region to the north of it to the Alakol’ and Sasykkol’ (basin of the
Tentek river), the Karatal basin, the Ш basin and, apparently, other
rivers flowing into [Lake] Balkhash; Kungei-Alatau (at least the
northern slopes of its eastern half), Zailiisk-Alatau (in particular,
Chilik basin), and the Chu river to its lower reaches (Sludskii,
1938, 1939, 1953). The otter was recorded in the Syr-Dar’ ya in its
lower course and for Chatkal [range] (Zarudnyi, 1915; Kuznetsov,
1948; Sludskii, 1963). Some of the data mentioned refer only to
the past, and at the present time in the otter is, apparently, absent
from a series of places.
For the entire remaining parts of the Tien Shan system, Fergana
and adjacent places, southward as far as Surkhandar’ya and the
Pamir, and westward to Samarkand there are no definite positive
data on the otter’s presence. There is only vague and indefinite old
information about occurrence of the otter in the lower reaches of
rivers flowing into Issyk-Kul’ (N. Severtsov, 1873) and for
Przheval’sk [city] (Kuznetsov, 1948). There are even data, that in
the past the otter was distributed throughout Kirghizia (i.e., through
the entire Tien Shan system, in the Alaisk range and Alaisk val-
ley), but by the beginning of the current century, it was “almost
destroyed”, although, “it is possible that it was somewhere still
preserved” (D. Dement’ev, 1938). Equally, there are direct refer-
ences about the otter’s absence in Kirghizia (Kuznetsov, 1948). At
the same time, the ecological conditions permit at least sporadic
occurrence of the otter almost everywhere in the mentioned moun-
tain system. One may think that, in the past, the otter’s natural
range occupied the whole Tien Shan and Pamiro-Alaisk districts,
except for such places as the western Pamir where conditions of
life for this species are unfavorable.
The otter’s distribution is more or less sporadic almost every-
where. In forest-steppe and steppe districts of the above outlined
northern range, there are, moreover, areas where the otter is com-
pletely absent (see for instance, Grigor’ev and Egorov, 1964). The
otter nowhere occurs in great numbers and moreover, at the
periphery of the range, but aiso in some regions far from the
866
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periphery, the animal sometimes disappears for a considerable time
and then appears anew. Local and long-distance transgressions may
occur, as mentioned previously.
Geographic Range Outside the Soviet Union
This occupies all of Europe, to the west including Ireland, on the
north the northern part of the Scandinavian Peninsula. On the
Balearic Islands, Corsica, Sardinia, Crete and Cyprus, the otter is
absent. The range farther includes northwestern Africa (Algeria
and Morocco), Asia Minor, Syria, Palestine, more northern parts
of Iraq except central and southeastern desert areas, Afghanistan,
Kashgaria, parts of Tibet (details unknown; apparently those con-
nected with Himalayas), Kashmir, Himalayas and northern
India to the north of the Ganges. Southern India and Ceylon form
a separate region of otter occurrence. In the central parts of India,
L. lutra does not occur, and therefore, the range of this species
here, like the ranges of the kharza [Charronia] and thar,
Hemitragus) are disjunct. The northern border of the range of the
southern Indian race (L. I. nair) is not accurately established. In
Central India, Lutra (Lutrogale) perspicillata lives.
Farther, the range includes Assam, Burma, the Indochinese
Peninsula, Malacca, Sumatra and Java. In Asia, the range includes,
moreover, the Mongolian Republic (mainly the northern and
montane parts of the country), northeastern China (former Man-
churia), the greater part of the rest of China (except the steppe and
desert regions), Japan, Taiwan and Hainan.
Within the limits of the outlined region of otter occurrence
Outside our country, there are considerable expanses where this
species is absent. Thus, it is absent in the greater part of the deserts
of Central Asia; apparently, in the greater part of Tibet, etc. De-
tails are, however, unknown. In some parts of the range the otter
has been exterminated (V.H.).
Geographic Variation
Although the otter’s range is vast, it is remarkable that its geo-
graphic variation is very small. It is greater in South Asia, al-
though one may think that presentations of it (for example, Pocock,
1942; Ellerman and Morrison-Scott, 1951) are nevertheless over-
1306
Е
are)
OM oS
NS
Fig. 315. Species range of the otter Lutra lutra L. V.G. Heptner.
estimated. Increase in variation along the southern edge of the
range in Asia is manifested both in the number of forms and also
in the degree of their separation. Thus, for example, the character-
istics of Java otters, L. (L.) 1. barang (material of Z[oological]
M[useum of] M[oscow] U[niversity]) in structure of not only the
skull but also dentition are so considerable that, from a purely
morphological point of view, they are analogous to specific char-
acteristics. It is characteristic that all the southern “border” forms
of Asiatic otters (nair, barang, chinensis) are small, while on the
whole, geographic variation of Eurasian otters reflects Bergman’s
Law well. A certain decrease in dimensions southward is also
noticed within the boundary of the USSR, but it is very weak.
867
1307
Himalayan otters (monticola, kutab) do not differ in dimensions,
or differ very slightly only, from the northern otters. Moreover, in
the south, there has been observed in places lightening of general
color and of the relative color of the venter especially of neck and
throat; i.e., increased contrast between the upper and lower body.
Geographic variation of otters in the Old World is still com-
pletely insufficiently studied. This also applies to the territory of
our country. Descriptions of separate races have been done on very
little material and have not subjected to serious revision. For the
USSR, the following forms may be preliminarily accepted:
1. Northern otter, L. (L.) [. lutra Linnaeus, 1758 (syn. vulgaris,
baicalensis, amurensis, kamtschatica, steinegeri).
Dimensions and mass relatively larger.
Color of back and sides dark-tawny, color of ventral surface
relatively dark. Underfur silvery, with dark chestnut-colored hair
tips. Pelage close-fitting and soft.
Whole range within boundary of the USSR except the northern
Caucasus, Trans-Caucasus and Middle Asia, and southern and
southeastern Kazakhstan.
Outside the USSR—in Europe, Mongolian Republic, north-
eastern China (former Manchuria), Japan.
The above given species description applies to this form [p.
1289]. It retains its principal characteristics in the whole huge
expanse mentioned and under all the diversity of conditions in
which it exists. Thus, even the separation of the Kamchatka otter
as a separate subspecies did not receive approval. Furriers within
the outlined limits also do not distinguish separate fur types, not-
ing only that skins obtained from Yakutiya and from the Far East
are of particularly high quality—of darker color and have very
dense underfur. The entirely weak variation in otter of northern
Eurasia is, apparently, associated not only with the fact that it is
an aquatic form but also with its ability to settle in new and quite
distant places.
2. Caucasian otter, L. (L.) 1. meridionalis Ognev, 1931.
Dimensions and mass relatively less large.
Color of back and sides lighter than in northern form—light
tawny. Ventral surface also somewhat lighter. Tail monotone both
above and below. Underfur silvery with yellowish tint and light
tawny hair tips. Pelage less close-fitting, somewhat coarse.
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Body length (14) 530-710 mm; tail length 340—470 mm; length
of hind foot 115-135 mm; ear length 20 mm. Weight 3.7-8.5 kg
(adult individuals from Okum’ river basin in western Trans-Cau-
casus and Kura; M.P. Pavlov, 1964).
In northern Caucasus and Trans-Caucasus.
Outside the USSR—in Iran, perhaps Iraq, Syria, Palestine and
parts of Asia Minor.
This is a well-distinguished form, but still poorly studied. In-
dividuals from Trans-Caucasus are quite typical, and northern
Caucasian population comprises some numbers of more darkly
colored individuals; i.e. it bears characters transitional to the north-
ern form. Distribution outside the borders of our country are en-
tirely unclear. Systematic and geographical relationships of this
form to the following one are unclear (see below); nevertheless,
affiliation of the Trans-Caucasian otter with the southern group of
forms is evident. Its range, moreover, if not completely, then per-
haps to some degree may be isolated, from that of the northern
form by the steppes of the Cis-Caucasus. One may, apparently,
consider that this isolation is greater than that from the southern
parts of the species range.
3. Middle Asiatic otter, L. (L.) 1. seistanica Виа, 1912 (syn.
oxiana).
Dimensions similar to preceding form. General color is lighter—
back is grayish-light tawny, usually with gray hairs; sides lighter
than back, the belly very light whitish-yellow or whitish-gray.
Sometimes has yellowish tone lightening throat forming a distin-
guishable patch variable in size and color. The tail is bicolored—
upper like back and lower a light dirty-yellow-white. Pelage is
coarse and not very close fitting.
In Middle Asia, on northeast including Dzhungarsk Alatau;
possibly also region of occurrence in Turkmenia.
Outside the USSR—in eastern Iran and Afghanistan.
This form is well differentiated from the northern, although
among the described type in the population are dark individuals
with fur color and characteristic very similar to that of the northern
form, especially in the mountains. It is very similar—if not
identical—to the Trans-Caucasian form to it. Together with it, they
themselves represent members of the group (its size not clear)
apparently of the Near Eastern type distributed along the southern
edge of the western half of the country’s Asian part. Ranges of the
1309
Trans-Caucasian and Middle Asian form (meridionalis, seistanica)
in their northern section are, to a considerable degree, quite fully
isolated from southern regions inhabited by the nominal form. This
applies to the Cis-Caucasian steppes, steppes of Kazakhstan, the
expanse between the Altai where the nominal form lives and
Dzhungarsk Alatau occupied by the Middle Asiatic (Sludskii, 1953).
If the otter’s range in the Near East were not so limited, one could
assume that forms of this type have settled (penetrated) with us
from the south. The group of otters mentioned is analogous to the
southern group of races of brown bear—isabellinus-syriacus.
Systematic and geographic relationships between the mentioned
forms are not clear. They are described from quite nearby places
(lower Gilmend-Tegeran) from very few materials, having similar
characteristics. It cannot be excluded that, in the future, with ac-
cumulation of new material, these forms may be united. Also not
clear is the relationshin, at least of the Middle Asiatic form, to the
other adjacent races described from the Himalayas and northern
India. The forms monticola, aurobrunnea and kutab were described
from Nepal (first two forms) and from Kashmir respectively, and
were also recorded in Kashmir and Punjab; i.e., in those places
nearest to our southern Middle Asiatic regions in part directly
neighboring (Kashmir).
Leaving aside the problem of relationships among the listed
Indian forms (they are hardly entirely independent), one may be
assured with great likelihood that our Middle Asian (at least Pamiro-
Alaisk and Tien Shan) otters are related to them. Apparently, they
must bear the name of the Kashmir form kutab (if it is separate
from monticola, which is quite doubtful).
One must also keep in view that the form seistanica was
described from the delta channels of a river which disappears in
the hot lowlands on the border of Iran and southern Afghanistan,
while all the other named races were described from montane coun-
tries. The name seistanica was left here, therefore, conditionally,
as a traditional one in our literature until direct comparison of our
material with Indian, and revision of all Himalayan and northern
Indian forms. Published data (Pocock, 1941) give a completely
confused picture both of systematic and geographic relationships.
It seems highly probable that otters of the Himalayas and the Middle
Asian mountains, at least to the east of Amu-Dar’ya, all belong to
one form, which should be designated by the name monticola. It is
869
1310
possible that the name seistanica may be related to the group
kutab, perhaps in the form of a synonym.
Outside the boundaries of USSR, the following forms are ac-
cepted: 1) L. (L.) I. angustifrons Lataste, 1885—Algeria, Morocco
(possibly a synonym of the nominal form); 2) Г. (L.) 1. chinensis
Gray, 1837—continental China, except Tibet and northeastern China
(former Manchuria), Taiwan, Hainan; 3) L. (L.) 1. monticola
Hodgson, 1839—northern India (Punjab, Kumaon, Nepal, Sikkim,
Assam); 4) Г. (L.) l. aurobrunnea* Hodgson, 1839—Nepal (the
most elevated parts), Garhwal (apparently, a synonym of the pre-
ceding form); 5) L. (L.) I. kutab Schinz, 1844—Kashmir, and to
Tibet (probable synonym of the form monticola); 6) L. ¢L.) 1. nair
Е. Cuvier, 1823—southern India and Ceylon (a well-distinguished
form); 7) L. (L.) l. barang F. Cuvier, 1823—Sumatra, Java,
Malacca, Indochina (a sharply-expressed race) (V.H.).
Biology
Population. The otter belongs among a number of low-density
animals. In the past, when exploitation did not yet considerably
affect the status of its stocks, the annual yield within the boundary
of the USSR was 5000 otters, over the course of a series of years.
Annual growth of this animal ranges from 20 to 30%. Taking into
consideration the absence of overhunting at that time, it may be
assumed that the number of otters in the USSR did not exceed
20,000 individuals, falling to about 10,000 in years of more inten-
sive hunting. The smallest number of otters is in Siberia, while in
West European countries, in spite of the predominance of culti-
vated lands and intensive settlement of the territory, it is more
frequent than in the USSR. The yield from the USSR constitutes
10% share of the global one, that of West Europe—a 25% share
of the skins; the cause of this is the fact that otter populations are
most of all limited by the character of the ice regime of water
bodies.
The otter is distributed throughout almost the whole USSR,
but very unevenly. Its range is not continuous, but forms a sort of
net, which is generally characteristic of all animals closely associ-
ated with fresh water. Density of river distribution determines,
first of all, density of the otter occurrence. However, there are
*Misspelled aureobrannea in Russian original—Sci. Ed.
869
870
1311
Fig. 316. Habitat of otter in Altaisk preserve. Lower Chul’chya river. Photograph by
G.D. Dul’keit.
other very essential causes limiting the number of otters and the
density of settlement in fresh waters of the USSR.
Habitat. The otter is a true freshwater animal. It usually
captures food in water, and here also takes its shelter in case of
danger. It usually only comes ashore to rest. The otter inhabits
freshwater bodies in all landscape zones. It also lives on the
seashore where it even becomes frequent locally, but usually only
in the mouths of rivers, or even creeks. Height above the sea level,
characteristics of relief and geographical latitude, as well as the
direct influence of climate are of no importance to otter in the
presence of other favorable life conditions. Where the otter is not
intensively hunted, it does not avoid the densely populated places
or even the neighborhoods of large cities. In Middle Europe, cases
are known of otters even living in city. On the whole, the number
of otters is inversely proportional to the number of hunters. For
the same reason, numbers of otters usually increase parallel to
increasing: afforestation of a place.
1312
ты.
870 Fig. 317. Habitat of otters in “Kedrovaya Pad’” preserve, southern Primor’e.
May 1963 (Photograph by А.С. Pankrat’ev).
The characteristics of river banks and their relative inacessibility
to humans are very important for occurrence of the otter. Almost
impassable wooded banks, reed thickets and turgai, and general
inaccessibility of the shore zone from land, are favorable for it.
Because of this, the otter evidently often prefers forest rivers, with
greatly cluttered channels choked with obstructions and broken
twigs and branches which make these rivers impassable not only
for boats, but also for a walking person. In spite of the fact that
the nature of these rivers hinders the otter itself in catching food,
it settles willingly in such rivers.
Rivers with transparent waters, rapid flow and rocky beds are
preferred by otters to water bodies with slow flow and polluted
water, muddy bottoms, matted water plants and opaque water al-
though it also settles in such bodies. All of this is related to con-
ditions for catching food. Abundance of its food is also very
essential: of deciding significance is the abundance of fish in the
water body. One of the main conditions in the otter’s ecology also
871 proves to be the character the water and snow regimes of the water
1313
871 Fig. 318. Winter aspect of otter habitat on Teletsk lake in Altai preserve. Tracks of
otter (here also tracks of kolonok and American mink). Photograph by G.D. Dul’keit.
body in the winter period. The presence of polynias [open water],
thawed patches, ice holes, and “spring places’”—opportunities for
Otters to penetrate into the water and catch food there—are
necessary for the otter.
The feeding possibilities in unfrozen sections of water bodies
are usually limited, and therefore open areas under ice, which are
formed during winter reduction of water [level] are of no less
importance to otter. This gives it the possibility of finding food
872
1314
under the ice without coming out onto the surface and in this way,
the significant part of the water body being utilized is extended.
With the freezing over or complete freezing of the water body, it
is not suitable for the otter in winter. The impregnable ice cover
and the rarity of unfrozen sections practically deny the otter the
possibility of exploiting even those rivers richest in food. Pro-
longed ice cover is also unfavorable for these animals.
Food. Vertebrate animals, and above all fishes, everywhere
predominate in the food of the otter. Frogs, birds and mammals
sometimes play an essential, but strictly seasonal, role. The signifi-
cance of invertebrates is small—these are mainly large water
beetles. Locally, crayfish and freshwater mollusks have some
significance. Consumption of plant food has been noted only for
otters held in captivity.
In the fish-rich channels of the Volga delta, the frequency of
fish in the otter’s diet constitutes 88% on average for the year. In
the warm period of the year, it remains at a level of 91.6-97.6%,
but in winter falls to 55.6%, while during the period when water
body are frozen, and because of great difficulty of their capture at
this time—to 27%. Among all fish eaten at that time, cyprinids
constitute 60.2%, carnivorous fish—23.6%, and sculpin—10.7%.
Fry, eaten avidly, constituting about 1/3 of all fish. The otter pre-
fers small fish to large. In first place in frequency stands the carp
(33.8%), constituting 25% of all fish eaten; in second place—pike
(24.3%); in third—roach (18%); followed next by sculpin (13.2%)
and shiner (11.6%). Many other fish are encountered, but in small
amounts (403 samples; A.G. Romashova).
In the upper Pechora, the proportion of fish in the diet consti-
tutes about 65% annually, fluctuating seasonally from 58% (sum-
mer) to 74% (spring). The significance of fish here is less than in
the Volga delta. Grayling (24.2%), rock sculpin (26%) and loach*
(13.4%), i.e. rheophilic forms, predominate, constituting on the
whole 79% of fish eaten. Of these 43% are pelagic species, and
57%—bottom [fish]. Limnophilous forms constitute only 21%, in-
cluding only 4% bottom [forms]. This is explained in that the otter
prefers water bodies with fast transparent water where it is easier
to catch fish. Inhabitants of deep water—whitefish, ide, are rare
here in the otter’s diet, and, as in the Volga delta, small fishes
*The Russsian word “golets” can mean either loach or brook trout—Sci. Ed.
1315
872 Fig. 319. Pra river in Oksk preserve (Ryazan district)—habitat of otter. Complete ice
cover. Path of otter (trench) along deep trail at -42°C. January 1950. Photograph by
V.G. Heptner.
(weight to 200 gm) and fry predominate. Large fish weighing
more than 1—2 kg are rarely met with; in individual cases, capture
of burbots weighing up to 4 kg were recorded. Pikes are not often
caught, especially small ones not more than 10 cm long. Commer-
cial species constitute 45.5% (1530 specimens; Teplov, 1953).
In the Lapland preserve (Kola Peninsula, Kislaya river basin;
414 samples—261 excrement in summer, 38 in autumn, 115 in
winter; Vladimirskaya, Lebedev and Nasimovich, 1953) fish also
873
874
1316
occupies first place in the otter’s food. In the yearly cycle, brown
trout predominates (35.5%), followed by pike (15.2%), burbot
(15.1%), whitefish (10.8%), grayling (8.1%), minnow [Phoxinus]
(7.8%), and perch (5.8%). Riverine species predominate over lake
forms and constitute 78%. Small fishes (up to 20 cm long) consti-
tute 70.2%, and large—29.8%. Only 4.4% were more than 50 cm
long. The otter begins to eat large fish in the trunk region. In
summer, fish constitute 23.6% of the otter’s diet, in autumn—
5.3%, and in winter—29.6%.
The otter feeds on grass frogs mainly in winter (29.6% occur-
rence) when it does not have adequate food. It obtains them from
the bottom of small unfrozen rivulets. In summer, it catches them
along the river banks and in swamps (23.6%). It occasionally feeds
on ducks (teal, golden-eye) and mammals (voles, water shrew,
squirrel) in isolated cases. Among insects, in winter it feeds mainly
on caddis flies (12.2%) and stoneflies (8.7%), and in summer—
ground beetles (2.6%). Among mollusks, it eats primarily Anadonta
(1.5%) in small numbers.
In the Altai, fish constitutes 95.9% occurrence in the otter’s
food, and the larvae of caddis flies—83.6% (Ternovskii, 1956).
In general, the species composition of fish in the otter’s food
depends on the composition of ichthiofauna of the given water
body and the vulnerability of each species. In Caucasus, brown
trout, barbel, Chondrostoma and others predominate. In forest riv-
ers of the Volga basin, the otter feeds in winter mainly on burbot,
and in summer on other bottom fishes. On the Murmansk coast, it
feeds on cod, salmon trout and others. In the otter’s diet, frogs are
a typical winter food, exhibiting sharp seasonal fluctuations. In the
upper Pechora, the average annual frequency of occurrence is 18%,
fluctuating from 10% in spring to 26% in winter (Teplov, 1953).
In the Volga delta this fluctuation is sharper—from 2.4-3.5% in
the warm period of the year to 46.1% in winter and 69.2% in
periods of complete freezing of the water body. It is a very impor-
tant food for otters in winter, but by no means a preferred food. In
spring, frogs are numerous in the Volga delta, but the otter hardly
catches them at all. It only hunts them under ice in their hibernat-
ing places where up to 1156 frogs may congregate per 1 hectare
(A.T. Romashova). On the Pechora, in the warm period of the
year, the otter feeds more intensively on frogs (10-18%). Here, the
13
873 Fig. 320. Polynias [open water] in Kedrovaya river—the otter’s place of permanent
residence in winter. ‘“Kedrovaya Pad’” preserve, southern Primor’e. December
1963. Photograph by A.G. Pankrat’ev.
water bodies are less rich in fish and conditions for capturing
them are, apparently, also less favorable.
The significance of birds in the otter’s food varies greatly. In
the upper Pechora their average annual frequency is 13%. In the
Volga delta, in spite of the abundance of birds, is two times less—
6.1%. In the upper Pechora in summer, birds comprise 26%; among
1318
them, ducks—7.9% and snipe—6%. Ducks of open waters are
encountered somewhat more often than those species which
confine themselves to dense growths of water plants. In the Volga
delta in summer, frequency of birds reaches only 1.8%, but in this
case, not ducks, but to coots and their nestlings. Cases of otters
hunting moulted ducks are very rare—the birds restrict themselves
to places unfavorable for hunting by the otter. Apparently the
increased frequency of frogs and birds in the food of the Pechora
otter is associated with small numbers of fishes.
The mammals represented in the otter’s food are mainly water
voles [Arvicola], less frequently water shrew [Neomys] and still
more rarely, other species which can appear on the shore of the
water body or may swim across it. Due to local conditions, the
significance of water voles varies. For example, in the upper
Pechora, depending on the season, the percentage of occurrence
ranges from 2 to 5%, and in the Volga delta, reaches 12% both in
summer and in winter. During the period of complete freezing, it
then increases to become one of the major sources of food—19.2%.
Complaints have recently appeared that the otter in the Ural [river],
noticeably affects the number of muskrats (Bakeev and Koryakov,
1960).
The daily ration of the otter consists of about 1 kg of fish.
When constantly supplied with food, the otter eats during a night
0.4—0.9 kg of fish, in the breeding зеазоп—0.2-0.4 kg, and after
prolonged starvation—up to 1.5 kg. (A.T. Romashova). Feeding
on small fishes, the otter becomes satiated overnight with 2-3
helpings, and with a large catch—after 1 helping. It is curious that
one water vole of about 145 gm can fully satiate it (A.T.
Romashova). This is probably conditioned by the higher nutri-
tional value of warm-blooded animals in comparison to fishes.
Home range. The home range of the otter is limited in width
to the narrow [river] bank zone—not wider than 100 m on each
bank. Depending on the food capacity of the water body, the home
range may be unitary or consisting of a series of separate ranges,
separated from each other by non-feeding areas. The length of the
section and its area depends also on the abundance of food and
conditions for its capture. The separate feeding sections may form
a chain extending for 15-20 km along the course of one river, or
the otter may periodically visit a series of adjacent lateral tributar-
ies, reaching them by way of their mouths or crossing the
875
1319
watersheds on land. In water bodies with abundant food, the otter
leads а settled life in a section extending from 2 to 3—5 km. In the
great taiga rivers of the Pechora basin, the average length of the
home range for a series of years was 6 km (Teplov, 1953). There-
fore, the area of the range inhabited by the otter fluctuates from 4—
12 to 300 hectares. On the Kola Peninsula, the otter covers 4-6 km
in 24 h period and sometimes more (Vladimirskaya, Lebedev and
Nasimovich, 1952).
In winter, when roaming around the watersheds, and also with
strong freezing of water bodies, the otter moves along snow and
river ice 15—20 km рег 24 h period. On the Kola Peninsula, leading
almost a migratory mode of life in winter, the otter commits a
transgression into a neighboring section 20-30 km in length and
sometimes including neighboring rivers and creeks. On ice, it moves
up to 15 km, and on snow, up to 8 km per 24 h period.
Burrows and shelters. Within the boundary of the home range,
the otter usually has a permanent burrow, and also a series of
temporary shelters and hiding places. The latter are entirely vari-
able depending on local conditions. These may be eroded areas of
steep banks protected by tree-roots, heaps of driftwood and fallen
branches, and caves in the rocky shore, etc.
The permanent burrow is often constructed in a steep, although
not high, bank. Its entrance opens under water, at a depth of 0.50-
0.75 т. The sloping passage, 1.5-2 m long and about 20 cm in
diameter, leads to a nest chamber 0.4—0.5 m in diameter, often
lined with dry grass or moss. In one case, it was found at a depth
of 20 cm from the soil surface (Teplov, 1953). In the family bur-
row, the nest chamber is always located so that it may be sub-
merged only during spring flooding. During that period the otter,
like the beaver and desman, is deprived of permanent shelters. In
low-lying places where high level of groundwater does not permit
construction of burrows, the otter builds a shelter for the litter of
young in high piles of reeds or fallen branches, in the heaps of
driftwood half-covered by sand and silt, flotsam,* and other cover.
In well-protected places, the otter sometimes gives birth to its young,
even on the surface, but does so in a carefully concealed nest—a lair.
*In the Russian original this word is misspelled “plavnik” (fin, flipper) instead
of “splavnik” (flotsam)—Sci. Ed.
1320
In winter, in Ападуг Territory, the otter sometimes settles т
a shelter under a steep overhanging bank which has an exit to the
sub-ice space. Its activity is thus hidden to a considerable degree
from the eyes of observers. On Kola Peninsula, otter forms occur
under snow, rarely on the shore or on snow-covered rocks. “The
dining room” is located on rocks which have on one side an edge
sloping to the water. “Latrines” occur under spruces with the crown
hanging near ground, usually directly on ice (Vladimirskaya,
Lebedev and Nasimovich, 1953).
x —
875 Fig. 321. Shore rocks with niches and clefts—shelters of otter in the course of
the entire year “Kedrovaya Pad’” preserve, southern Primor’e. May 1963.
Photograph by A.G. Pankrat’ev.
1321
Daily activity and behavior. The otter leads mainly а сгериз-
cular-nocturnal mode of life. It is especially active on moonlit
nights. In places where it is not disturbed, it sometimes also ap-
pears during the daytime; this more often occurs in winter than in
summer. Increase in diurnal activity in winter sometimes manifests
dependence on diminished conditions of illumination under the
876 layer of snow and ice, which hinders the otter in catching fish at
Fig. 322. Winter shelter of the otter on an eroded bank. A bunch of dry grass at the
edge of an ice-free area, which the otter was carrying to the shelter, and dropped upon
the approach of a human. “Kedrovaya Pad’” preserve, southern Primor’e.
January 1961. Photograph by A.G. Pankrat’ev.
877
1322
night under the ice (Teplov, 1953). On the Kola Peninsula, it is
active round the clock (Vladimirskaya, Lebedev and Nasimovich,
1953).
The otter leads a secretive mode of life. It hunts, by waiting
for its prey on shore, or by pursuing it. Obstructions in the water
body, a silty bottom, unclear water and growth of water vegetation
hinder capture of food by the otter, but at the same time increase
the protected nature and safety of the habitat.
On dry land, the otter is clumsy and less mobile, but has en-
durance, and even in deep snow, progresses for ten kilometers or
more. No dog at all can overtake the otter, even following fresh
tracks. In water, the otter is very quick, dexterous and mobile. It
catches large fish, by diving under them, seizing the abdomen, or
rarely taking the back. It may happen that a very large fish pulls
away from its teeth and escapes with wounds (A.T. Romashova).
Diving under the water, the otter is at first not evident, but then its
path may be followed by the bubbles exhaled when it breathes out.
For breathing, when there is danger, the otter sometimes thrusts
only the tip of its nose to the surface. Sallying out of the otter onto
shore most often occurs in definite, well-concealed places, and
usually they have the character of beaten tracks, similar to those of
the beaver. In some places, so-called “otter slides”* occur. These
are smoothly polished, well-beaten paths on steep slopes of the
bank, often damp and slippery. There are observations that otters
“amuse themselves” by sliding many successive times along these
steep tracks into the water. Moving along the surface of the ice,
the otter quite often, making a running jump, slides along on its
belly, leaving a long trough-like track. Such behavior in the otter,
as in mink, is explained by the need to dry the wet fur (Ternovskii,
1956).
The otter mainly hunts “stationary” fishes or those slowly swim-
ming. It also hunts at “fishing holes” and hunts fry on small bars.
When hunting, the otter swims with its head and part of its back
on the surface. After seeing its prey, it dives.
Otters usually live singly. However, females and their litter
stay together for the whole autumn and winter, and usually remain
with it until mating again. The female defends and protects the
litter, and the male, apparently, sometimes participates in its up-
bringing.
*Literally, “sliding hills” in Russian original—Sci. Ed.
1323
Seasonal migrations and transgressions. The otter is an ex-
tremely mobile animal. However, its frequent wanderings, from
one water body to another, giving the impression that it is migra-
tory, nomadic animal, actually take place within its home range.
These wanderings are explained by water bodies becoming food-
deficient especially in winter, with an unfavorable ice regime, the
home range occupies a considerable length of shoreline. Neither
can the otter’s crossing of watersheds be called migration. These
also bear the character of movements within a vast home range.
Reproduction. Reproduction in otters is still very insufficiently
studied. A series of contradictory points of view exist. These, as
well as the existing facts, may be harmonized according to the
existing level of our knowledge in the following way. 1. As in
many species of the mustelid family, there is a latent period in
embryonic development in the river otter. The duration of this period
is completely unstable and is determined by environmental factors.
In several sections of the range, the latent period is liable to indi-
vidual fluctuations and in others—it is more or less stable and is,
at the same time, different in different sections. After the end of
this period, pregnancy apparently proceeds for about 60 days with-
out deviation. 2. Corresponding to variations in duration of the
latent period, the times of estrus and mating are subject to some
deviations. Most frequently, apparently, estrus and mating take place
during March and last about one month. This applies to the Volga
delta region (A.T. Romashova), Mari Republic (V.A. Popov), up-
per Pechora (Teplov, 1953) and southwestern Kalinin district (P.B.
Yurgenson). The date of beginning of this period is changeable
(February—April).
On the other hand, there are reports of otter cubs of one and
the same age (one month) found in June and October-December.
Quite well-developed embryos were found in otters caught in Janu-
ary. In February, an otter cub the size of a domestic cat was taken.
Such dimensions of young were attained both in January and July.
An otter cub which could hardly walk was once found on the snow
(A.T. Romashov). The data obtained from the tributaries of Amur
also confirm that otters reproduce at any time of the year—“like
dogs’. There is a report that in the Caucasus, estrus in otters ос-
curs at the beginning of winter. For the Volga delta, the possibility
of mating in times other than March is not excluded, but such
exceptions are rare. Late litters usually die in winter. Parturition
878
1324
occurs here in June—beginning of July. In the upper Pechora, it
occurs at the end of April—beginning of May, and in June active
young otters are observed. On 18 June, two young males were
caught, with the eyes opened, but the teeth had barely erupted
(weight 1 kg, body length 33.5 cm; V.P. Teplov). Most often,
births occur in May. Previously mentioned differences in dates
serve as adaptations to the local ecological conditions. The number
of young in the litter varies from two to four. Individual cases are
known when there were 5 otter cubs in a litter (Khorol river,
Ukrainian SSR). In the upper Pechora, no more than two otter cubs
were observed in litters. The number of young is closely related to
the degree to which environmental conditions are favorable.
In England, the newborn cubs are encountered at any time of the
year, not excluding winter (Stevens, 1954). In one region the majority
of births took place late in autumn and in winter (October—February),
and in others—in the period from January—February to April inclu-
sive. In 134 cases (for several years), the occurrence of new-borns
was distributed by months of the year as follows: January—14;
April—10; July—11; October—12; February—8; May—11;
August—13; November—11; March—10; June—11; September—
11; December—12.
In the British Isles, winter is mild, and the otter is not subject
there to sharp seasonal fluctuations in conditions of existence. In
Captivity, estrus in the female is repeated each month until mating
is productive. Mating is accompanied by a sharp whistle produced
by males. At this time, a fight may occur between them. Pregnancy
lasts 61-63 days. Duration of the latent period is variable and may
be more than 9 months.
Growth, development, and molt. Data are few and fragmen-
tary. Young animals are born blind and without teeth, with short
dark underfur. They begin to see at the age of about 35 days
(Stephens, 1954). In the female, there are three pairs of nipples;
however, usually only a part of them functions. Young females
gain in dimensions and weight more slowly than males. Young
otters gain about 400 gm in weight each month, and at 10 months
weigh about 4 kg. At the age of one year, they attain the size (but
not the weight) of adult individuals. Sexual maturity in individual
cases (in Captivity) is attained in the second year, and probably
more often, in the third.
1325
879
Fig. 323. Otter on the ice of a creek near an otter slide. “Kedrovaya Pad’” preserve,
southern Primor’e. January 1961. Photograph by A.G. Pankrat’ev.
Molt in the otter proceeds very gradually, and is little noted.
Enemies, diseases, parasites, mortality, competitors, and popu-
lation dynamics. Other than humans, the otter has no dangerous
enemies. Neither has it any serious competitors. It usually forces
out mink from the borders of its home range, often destroying
them. On the Kola Peninsula, the wolverine was referred to as an
enemy of the otter, and the white-tailed eagle as a competitor
(Vladimirskaya, Lebedev and Nasimovich, 1953). Beaver activity
is usually useful for the otter—in winter the latter uses beaver
outlets from under the ice. Diseases of epizootic character are
unknown. In captivity, coccidiosis infection has been recorded. In
a severely emaciated otter killed while attacking a domestic goose,
pulmonary tuberculosis and endocarditis were diagnosed (Stephens,
1954). Helminth infections are not major.
The main cause of mortality, especially up to one year of age,
is unfavorable weather conditions in winter, in particular, the ice
regime. Late litters usually die in winter. In the upper Pechora, a
connection exists between the survival of young, magnitude of
autumn flood favoring the formation of sub-ice spaces and the
879 Fig. 324. Track and prints of night life of an otter family—a female with young.
18 December 1964. Photograph by A.G. Pankrat’ev.
duration of the ice-cover period. The otter population there is
determined by intensity of reproduction and survival of young.
The average percentage of young in otter populations was 19.0,
with variation from 12 to 26%, i.e. more than 2 times (Teplov,
1953, 1954).
The otter responds very quickly and positively to prohibition
of hunting (launching)* with real observance of it. The cultivated
landscape does not directly prevent the growth of the otter
population if they are protected from direct pursuit and destruc-
tion. Only measures giving rise to impoverishment of the
ichthyofauna of water bodies have proven to be harmful: floating
forest litter, sewage drainage from industrial enterprises, etc.
Field characteristics. Sightings of river otter are rarely suc-
cessful. It is difficult to confuse with any other animal. Tracks of
otter, and their excrement are often encountered. The large tracks
of this animal most often bear well-marked outlines of the swimming
*Meaning of parenthetical word “zapusk”, or “launching” not clear—Sci. Ed.
880
881
2327
membrane between the digits. Its track is clearly separable from
that of the beaver in that it usually extends along the bank at the
edge of the water, while the path of the beaver is always perpen-
dicular to the line of the bank—these are its sallies onto the bank.
Excrement of the river otter usually possesses a greenish tint,
and is often watery. The remains of fish bones and scales of fish
are typical of them. From mink excrement it differs in large size.
The otter usually leaves them [feces] on rocks in and around the
water body or on the bank, as well as in holes on sandbars. It is
often buried. Its urine typically has a strong odor peculiar only to it.
On snow, the otter leaves not only its footprints, but also the
track of its dragged tail. In deep snow it makes a whole furrow
with its body. On river ice, a long ribbon of a trail (sliding after
running) is typical, as are tracks emerging from water.
The otter sinks into the snow for 10-15 cm; the length of its
leaps equal 50-110 cm more often 60-80 cm; the length of the
trail made sliding on snow is 2—3 meters (Vladimirskaya, Lebedeve
and Nasimovich, 1953). The otter sometimes proceeds for up to 6
km along ski-tracks. The weight loading on 1 cm? of the foot
surface is equal to 53.2 gm for the male and 37.4 gm for the
female (Teplov, 1953). In the period of reproduction, the otter’s
presence may be revealed by whistle of males at night (P.Yu.).
Practical Significance
The river otter provides very valuable fur—beautiful, warm
and durable. From it are manufactured fur articles in natural shape
but with plucked guard hairs, or even dyed a black color as
“sealskin”. Such articles are usually sold as a “true seal”.
The otter is not numerous and is captured in limited numbers,
more or less occasionally and accidentally. At the present time,
capture of this species is carried out by licensed trappers. In RSFSR,
6100-6700 otters were trapped by license in 1956 and 1957. The
annual catch for the USSR was about 7.5 thousand for the same
years. The world catch of otter before the Fatherland War [World
War II] was 80-100 thousands.
Due to the small number of otters in the USSR, the harm from
it to the fishing economy is negligible.
The otter is usually hunted with guns and dogs, more rarely by
still-hunting on moonlit nights. Harvest by jaw-traps is of a certain
880 Fig. 325. Tracks of ап old otter male on light, wet new-fallen snow on the ice.
“Kedrovaya Pad’” preserve, southern Primor’e. March 1962. Photograph by
A.G. Ponkrat’ ev.
significance, but good traps for catching this animal are not avail-
able. All other methods are either not productive or are not al-
lowed as inhumane.
882 The otter is easily tamed and capable of domestication, and
after a more detailed study of its reproductive processes might be
bred in captivity. This is, however, possible only where there
exists abundant and cheap food (fish) (P.Yu.).
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881 Fig. 326. Prints of hind and fore feet of otter on a sand bank and the scheme of
arrangement of the tracks during different gaits. Neya river, Kostromsk district.
24. X-1950. Sketch by A.N. Formozov, about 2/3 natural size.
3
1330
Genus of Kamchatka Beavers ог Kalans*
Genus Enhydra Flemming, 1822
1816. Pusa. Oken. Lehrb. 4. Naturgesch., 3, р. 985. Nom. ргаеосс.
Pusa Scopoli, 1777, Phocidae.
1822. Enhydra. Flemming. Philos. of Zool. 2, p.187. Mustela lutris
Linnaeus, 1758.
1827. Latax. Gloger. Nova acta phys.-med. Acad. caes. Car.-
Leopold, 13, p. 511. Renaming of Enhydra Flemming, 1822.
1829. Enhydris, Fischer. Synops. Mammal., p. 228. Substitute for
Enhydra Flemming. Nom. praeocc. Enhydris Merrem, 1820,
Reptilia (V.H.).
Measurements very large—largest in the subfamily and family.
General body construction heavy, trunk very long, cylindrical
in form. Limbs short, hind [limbs] displaced backward. Foot flip-
per-like, paw of fore limb short, digits unsegmented externally.
Tail muscular, its length constitutes one-quarter to one-third that
of body.
Fur long and very dense. Color tawny and brown in tone. Ex-
ternally, glandular areas at base of tail absent. Two pairs of teats,
but only one functions in female.
Skull massive and heavy. It is high and broad, with facial
portion strongly shortened and elevated anteriorly, and terminated
nearly vertically. Postorbital constriction only slightly narrower
than interorbital. Brain case short, but very wide. Mastoid width
only slightly less than zygomatic. Protuberances and crests of skull
sharply defined; not only occipital but also sagittal crests well
developed.
api aah feat
Dental formula Dae Мат” i.e. Somewhat reduced.
Incisor formula 3/2—a rare exception among carnivores (one
bear has 2/3). As a form of anomalous deviation, an additional
underdeveloped lower incisor may be manifested. It is probable
nie vik wb by) sbiainig
that in milk dentition is ie
*“Kamchatka beaver” was the common name originally bestowed by Russian fur
traders. At present, the accepted common name in English is “sea otter,” and it will
be used in this account, together with the native name “kalan”—Sci. Ed.
883
1381
Anterior premolars with obtuse apices, posterior premolars
and molars are almost flat, with low, blunt, rounded apices. “Car-
nassial” structure of upper third premolar not expressed. It has a
rounded triangular form, with one angle directed inwards. Upper
molar considerably larger than last premolar, of rounded triangular
form, one side directed forwards toward premolar and other side—
inwards. First premolar only slightly shifted deeper into toothrow.
Lower carnassial tooth not developed either. This is an extreme
deviation in the family in the sense of loss of “carnassial” structure
of the dentition formula, is also observed in badgers and otters
(Lutra). Here, the cutting form of the tooth is entirely lost. Such
a structure itself represents an adaptation to feeding on sea-urchins
and mollusks.
Vertebrae 50-51 (С 7, D 14, L 6, $ 3, Ca 20-21), thoracic
region elongated and composed of 14 pairs of ribs, of which 10
articulate with sternum. Scapula broad with poorly developed
acromian process; pelvis narrow and long, situated to a consider-
able degree parallel to vertebral column, humerus very massive,
noticeably spirally twisted. Femur shortened.
Stomach relatively large, intestine very long (exceeding body
length by approximately 10 times), gall bladder present. Ligamen-
tum teres absent. Lungs relatively large, diaphragm lies obliquely:
fixed more posteriorly on dorsal side than on ventral (characteristic
feature of truly aquatic animals, associated with increased dimen-
sions of lungs).
Testis located under skin (scrotum is not formed).
On the whole, in all of its morphological features, the genus
Enhydra represents an extreme degree of specialization in the sense
of adaptation to water, not only in the subfamily and family, but
also in the order as a whole. From a purely biological and
bio-morphological viewpoint, sea otters occupy an intermediate po-
sition between those types of aquatic mammals which are repre-
sented by pinnipeds and terrestrial mammals, standing in the series
relatively closer to the former. Of course, the described genus must
not be considered as genetically linked between these groups; how-
ever, it indicates one of the possible biological stages in the path
of transformation of terrestrial mammals to aquatic.
A purely marine littoral form of temperate and cold temperate,
and in part warm temperate waters, specialized for feeding on fish
and marine invertebrates (mainly sea-urchins).
1332
Distribution restricted to shores of the northern part of the
Pacific Ocean, north from 23° N. lat. in the east and 40° N. lat. in
the west.
The genus Enhydra is restricted very sharply, and stands apart
from other genera of the subfamily, especially Lutra and Pteronura.
In fact, differences from the genera Aonyx and Paraonyx are also
just as sharp, and attempts to relate them are deprived of any basis.
On the whole, the genus Enhydra contrasts with the remaining
genera of the subfamily, each of which is nearer to one another
than to Enhydra. It represents an extreme type, and a terminal link
in the development and specialization of the otter group. Attempts
to separate it even into a special subfamily have been made more
than once.
The origin of the genus is usually associated with the extinct
genus Potamotherium comprising several species (at least two)
from the upper Oligocene and the lower Miocene of Europe. Con-
necting links between Potamotherium and Enhydra are not clear;
in the mind of some authors (Pohle, 1919), for this may serve one
of the fossil European species of the present African genus Aonyx
(A. hessica); however, this is doubtful. The genus Enhydra is
known from the upper Pliocene, when it was represented by the
extinct species, E. reevei, in the North Atlantic (England). This
species is considered the direct ancestor of the present kalan [sea
otter]. It is supposed that in the Pleistocene, from the Atlantic,
Enhydra colonized the Pacific Ocean (Pohle, 1919). It is entirely
possible that the path of colonization was the sea washing north-
ern Eurasia. Generally, the geological history of the genus is un-
clear, and there is a basis for assuming its earlier occurrence in
the Pacific Ocean.
This is one of the characteristic species of the northern Pacific
(“Beringian”) center of development of aquatic and inshore fauna,
in which are included such remarkable forms as Steller’s sea-cow,
sivach [Steller’s sea-lion], and a series of bird species (black guil-
lemots, cormorants).
The genus includes only one species: Enhydra lutris Linnaeus, 1758.
One of the most valuable fur-bearing species. Its falls under
full protection throughout its entire range (V.H.).
884
1333
KAMCHATKA OR SEA BEAVER [OTTER], OR КАГАМ!
Enhydra lutris, Linnaeus, 1758
1758. Mustela lutris. Linnaeus. Syst. Nat., Ed. X, 1, p. 45. Com-
mander islands?.
1777. Lutra marina Erxleben. Syst. regni. anim., p. 445. North
Asia and America.
1799. (1800) Lutra gracillis. Bechstein. Thomas Pennants allg.
Uebers 4. vierfiiss Thiere, 2, р. 177. “Statenland”—one of
the southern islands of Kuril chain, apparently Kunashir.
1816. Pusa orientalis. Oken. Lehrb. d. Naturgesch., 3, p. 986.
1827. Lutra stelleri. Lesson. Man. de Mammal., p. 156. Kamchatka.
1922. Enhydra lutris kamtschatica. Dybowski. Arch. Tow. Nauk.
Lwow, 1, р. 350. Nom. nudum (У.Н.).
Diagnosis
Only species of the genus.
'The true Russian name of this species must be considered “Kamchatka bobr”
(more correctly bobior), or “sea bobr’. The old popular name of the Bering
Sea—“Bobr Sea”—was derived from this word. The name of the animal came into the
language of literature through Pushkin (“Its bobr collar silvers as the frosty dust”).
The word “kalan” somewhat modified from the word “kalakha” was used by Russian
people only in the 18th century and then very rarely (Pallas, 1811). At the present
time, because of the danger of mixing it up with the river bobr [beaver] (!), some
zoologists use it in our literature. Following this “principle”, evidently such names as
“sea slon” (northern elephant seal) and “sea lev” (sea lion), “sea ezh” (sea urchin),
“sea liliya” (sea lily) etc. must also be changed. The name “kalan” came into
daily use, but must now be considered as artificial and bookish. It is not known
there. Such is also the name “sea vydra” [sea otter] a translation from German.
Fur manufacturers name the young animal suckling still feeding on milk
“medvedok” [young cub], the semiadult (1 year)—“koshlak”, the female—“matka”
[dam], and the adult male—‘bobior, bobr” (=otter).
*The assignment of Linnaeus “habitat in Asia et America septentrionali” was
fixed in that sense by Barabasch-Nikiforov (1947). Some authors consider
Kamchatka as the type locality of the species (Ognev, 1931; Ellermann and
Morrison-Scott, 1951, and others), which has little basis. To avoid undesirable
nomenclature changes (renaming), the submitted interpretation of the restricted
type locality must be sustained.
Steller’s description of “Lutra marina” was given before 1758 (in 1751) and has
no nomenclatral significance. “Viverra aterrina” of Pallas is sometimes submitted in
lists of synonyms (Yakobi, 1938) but has no relation to the described species (Heptner,
1934). It is a kharza (see [Vol. II, pt. 1b]). In Erxleben, the name Г. marina is
associated with the given species only partially, because the author also had in mind
the Brazilian otter.
885
1334
Description’
Externally the sea otter is to the highest degree unique. Trunk
elongated, of cylindrical form, and quite massive. Neck thick—
thicker than head, and short. Limbs very short, hind ones displaced
posterior. Animal appears clumsy and heavy appearance. This
impression is partially due to not only very luxuriant, although not
very long fur, but also to skin loosely enveloping the trunk—as if
in a sack. On land, movement of the animal is hesitating and
relatively slow; while lying or moving, it usually flexes its back
strongly. In water, it is very agile, mobile and flexible.
Head rounded, with dense long vibrissae and small eyes. Ear
pinna small (resembles external ear of seals), auditory openings
slit-like, may close (during submergence into water), nostrils are
slit-like, closeable. The iris is blackish-tawny.
Foot flipper-like, all digits invested with common covering
and fused to last phalanx; digit V (outer) very long, remaining
gradually decrease in length to very shortest first; foot above and
below covered with wool. Fore limb strongly shortened. Paw short,
digits not segmented externally and only weakly defined. On lower
side is found somewhat bulbous, round bare area, in anterior part
of which four small parts marked by slight grooves corresponding
to digits, and in posterior—one common narrow part extending
transversely, [Fig. 328]. A second external part of fore limb in-
cludes two very long digits (on upper surface, there are corre-
spondingly two claws), remaining one of three include one digit
each. Claws of both fore and hind limbs very small, dark horn in
color. Tail covered with a dense short fur, relatively short (about
25-30% of the body length), muscular and slightly flattened
dorsoventrally.
Fur of sea otter not especially long, but exceptionally dense,
soft and silky. It is one of the most precious furs, in fact is not
comparable to any other fur as regards its beauty (especially “gray
otter”) and durability.
Guard hair and underfur differ little from each other in length,
and are, moreover of more or less equal length throughout the
whole body. The longest fur is in the middle of the back. In
Commander otters, average length of guard hairs here is 27.7 mm,
’Mainly according to Barabash-Nikiforov (1947) and materials of
Z[oological] M[useum of] M[oscow] U[niversity].
1335
Fig. 327. Sea otter, or kalan, Enhydra lutris L. Sketch by A.N. Komarov.
Fig. 328. Lower surface of fore foot of kalan, Enhydra lutris L. Sketch by
N.N. Kondakov.
886
1336
of underfur—22.5 mm. For withers, 24.3 and 19.5 mm, гезрес-
tively; for sacrum—24.5 and 21.7 mm. On scapula, hair length is
24.5 and 19.0 mm; on middle of side, 25.9 and 20.3 mm; on hip
20.3 and 16.4 mm. On breast, guard hairs have length of 25.3 mm;
underfur 15.0 mm; on belly, 23.4 and 17.0 mm; on groin, 21.1 and
15.0 mm. Maximum thickness of guard hairs is on breast (169.5
micron). On belly, it equals 147.6 [uw]; on middle of side, 136.6
[4]; on sacrum, 125.0 [4]; on middle of back, 123.9 [4]; and on
hip, 115.3 [4]. Tail is covered with shorter hair compared to trunk—
along its dorsum, length of guard hairs and underfur is 19.0 and
16.5 mm; along venter, 18.5 and 15.1 mm [respectively]. Guard
hairs on dorsum of tail even coarser than on whole trunk (except
breast): their thickness is 153.2 [в]. On lower surface of tail, their
thickness is 115.7 [и] (all figures are mean of 50 hairs). On 10
mm? of back, an average of 12.4 (in summer), and 14.6 (in winter)
guard hairs occur, as against 1910 and 1725 underfur hairs, respec-
tively; on belly in summer, there are 20.2 and 1674, and in winter,
17.2 and 2221, respectively (Barabash-Nikiforov, 1947).
General color of fur is dark brown. Facial portion of head
brownish-gray, light straw or almost white; vibrissae are white.
сосен ра soagoee оо и НН
886 Fig. 329. Kalan оп Mednyi Island, March 1964. Photograph by S.V. Marakov
2837
Rear portion of head and occiput dark brown, sometimes mixed
with separate white (“gray”) and light reddish hairs (guard hairs
with light tips). Fur on neck, withers and back dark brown, of an
especially dense, deep, almost black, color on back. On the neck
and withers, there is a significant mixture of yellowish-gray hairs,
white and reddish guard hairs, and also on base of tail. Lower
surface of neck and breast brownish-gray or even dirty-white. Ab-
domen dark brown, only a little lighter in tone than back. Transi-
tion in color from back to abdomen is gradual. Fore and hind legs
similar in color to sides, but slightly darker. Gray hairs are absent
on them. Upper side of tail same color as back and lower side—
as in abdomen (prevailing type of Commander kalan, chiefly after
Barabash-Nikiforov, 1947).
Individual variation in color is significant. General tone of fur,
from the [above-]described and most widely-distributed type, pro-
ceeds through gradual gradations to deep pitch-black, or through
Fig. 330. Old kalan. Mednyi Island, May 1960. Photograph by S.V. Marakov.
887
888
1338
tawny to light brownish. These differences are determined by
characteristics of pigmentation of the terminal part of the hair—
their bases have a very constant gray-straw color. Not rarely,
there are significant exceptions to the described lightening of the
anterior part of the trunk, and individuals are sometimes encoun-
tered with an almost white (cream-colored) head, neck (upper and
lower) and breast. Otters with white heads and dark-sandy trunks
belong to the category of extreme color deviants; completely white
otters are also met with (albinos; Fig. 331).
Admixture of gray hairs is entirely variable. Usually, they are
distributed on withers and anterior part of upper body, but some-
times occupy a greater expanse. Sometimes they are unevenly dis-
tributed, as in patches of different degrees of density. In a series
of cases, they may be completely absent. Beside individual varia-
tion, the path and condition of molt itself have influence on degree
of gray hair development (see below).
Seasonal changes in color and density of fur are insignificant.
As in several other typical aquatic animals, molt in sea otters is not
limited to a defined period, extending throughout the entire year,
proceeding slowly, and pelage is always actually complete, or al-
most complete. However, molt is more intensive in summer months,
which is responsible for noticeable differences between “summer”
and “winter” fur. Before shedding, there is, in places, lightening of
guard hairs, and therefore during the period of more intensive molt,
the quantity of light (gray) guard hairs on the skin is greater.
Therefore, intensity of molting, i.e. mainly summer animals, appear
lighter—“gray”. Except change in quantity of gray hair, which gives
general fur color its tint, the basic color of the animal also changes
somewhat. In summer, it acquires a lighter brown tone, apparently
because of fading of the fur under the influence of the sun’s rays
(Reshetkin and Shidlovskaya, 1947).
Sexual differences in color are absent, or is only revealed in
slightly lighter color of females. In females, the quantity of guard
hairs is, on the whole, somewhat less, in connection with which
their fur is generally somewhat softer.
Age variations in color are quite significant, and proceed slowly.
Recently-born animals (medvedoks) have light brownish-red or
reddish-tawny fur. Head and neck are lighter than remaining part
of trunk, but the contrast is not as sharp as in adults. Abdomen is
somewhat darker than back; fur is dense, but relatively short, with
1339
Fig. 331. А group of kalans in Zapalat Вау on Mednyi Island. In the fore-
ground—Albino kalan. On rocks, glaucous winged gulls (Larus glaucescens Naum.).
6 August 1962. Photograph by S.V. Marakov.
coarse guard hairs and is quite harsh to the touch. Such fur occurs
in animals 5—6 months old or less. In a yearling, but not yet adult,
otter (koshlaka), fur does not differ in color from that of adults,
but, apparently, still contains some quantity of coarse guard pelage
of the medvedoks. In both of these stages, gray hairs are absent.
Maturity is attained by sea otters, apparently, in the fourth year. In
young, the iris is nut-brown.
Geographic variations in color are poorly studied and, gener-
ally speaking, are insignificant.
Skull large and massive, high and broad, but short, with sharply
shortened, high facial portion, as if vertically cut anteriorly [Fig.
332]. Its length is approximately two times less than cranial part of
1340
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889 Fig. 332. Skull of sea otter, or kalan, Enhydra lutris L.
skull. Upper profile of skull almost horizontal, weakly rising
posteriorly. Cranial portion of skull relatively very large and wide—
its width only slightly less than width of zygomatic arches.
Zygomatic arches massive and strong widely separated laterally.
Auditory bullae large, located transversely, auditory openings very
small, directed obliquely upwards. Nasal bones short and broad.
Upper processes of premaxillae narrow, extend to nasal bones, but
889 contact only their anterolateral processes. Postorbital processes
890
1341
weakly developed. Postorbital constriction short and broad,
constituting about one-fourth condylobasal length, but less than
interorbital constriction.
Nasal openings terminate almost vertically, very wide—their
width greater than width of orbit (from postorbital process to
zygomatic arch). Nasal turbinates very strongly developed—intri-
cately convoluted, filling the whole nasal opening and extending
forwards nearly to its outer termination. Hard palate cleft behind
cheek teeth very short and almost unconstricted. Sphenopalatine
notch very short and wide. Diameter of infraorbital foramen is
nearly equal to diameter of alveolus of upper canine. Mastoid proc-
esses isolated, blunt and massive, directed obliquely downwards.
Opening of jugular foramen very large—its diameter approximately
equal to diameter of alveolus upper canine or slightly less. Paroc-
cipital processes very weakly developed. Coronoid process of lower
mandible very large (60-80% of mandible length from angular
process to alveoli of incisors), angular process small. Protuberances
and crests of skull on the whole, and sagittal occipital crests in
particular, sharply expressed in old [individuals].
Dental formula—see the characteristics of the genus.
Individual variations in skull lie in several changes in general
dimensions and proportional details. It is a fact, that not rarely
there is in places asymmetry of the skull—somewhat stronger
development of the left side of the braincase. This phenomenon is
observed not only in old animals, where it is manifested in particu-
lar in twisting of the sagittal crest, but also in much younger
animals.
Age changes in skull very great. In young, postorbital proc-
esses weakly developed, width of postorbital constriction less than
width of interorbital area; facial portion of skull relatively still
shorter; braincase, on the contrary, still more developed, more
rounded and swollen; crests (sagittal and occipital) absent; suture
between basioccipital and basisphenoid not fused*. In the upper
toothrow, last tooth is third permanent premolar.
Sexual differences in skull not sharp, but obvious. Skull of
female, beside smaller general dimensions, is distinguished by its
relatively weakly developed crests, less massive zygomatic arches,
narrower hard palate and some weaker teeth.
*Russian word is zarosshii; lit. “grown over’—Sci. Ed.
1342
Geographic variation of skull insignificant, manifested,
apparently, only in general dimensions.
Os penis very large, quite massive and, on the whole, bent
slightly upwards; anterior end forms slight bend towards ventral
side. Usually, also small bend in lateral direction. At base, bone
broadens and flattens, tapering anteriorly and becoming round in
cross section. At extreme end, there is a small thickening with a
deep depression on lower surface.
Body length of full adult (from four years old) and old indi-
viduals (Commander islands; 36), 100-136 cm (usually 120-130
cm); tail length, 30-36 cm; length of hind foot, 20-23 cm; length
of ear, 24-26 mm. One especially large old kalan had a body
length of 146 cm. Probably, in individual cases, body length may
reach 150 cm (Ognev, 1931; Barabash-Nikiforov, 1947, 1962;
Stroganov, 1962). Females somewhat smaller than males (accurate
data absent).
Condylobasal length of male skull (31) is 124.2-145.9 mm, of
females (9), 120.2—142.9 mm; zygomatic width of males 1$ 96.9—
116.2 mm, of females, 90.0-102.0 mm; interorbital width of males
is 42.0—46.5 mm, of females, 36.7-42.2 mm; postorbital width of
males is 28.0-36.0 mm, of females, 27.0-35.5 mm; length of upper
toothrow (cheek teeth and canine) of males is 44.4—51.4 mm, of
females, 44.3-49.2 mm (Ognev, 1931; Barabash-Nikiforov, 1947;
materials of Z[oological] M[useum of] M[oscow] U[niversity];
Commander islands, a few specimens from Kamchatka).
New-borns have body length of about 38 cm (Brandt, 1880)
and weight of about 1.5 kg; “not long after birth’—about 44 cm,
tail length about 15.5 cm and weight about 2 kg; at two years old
on average (36)—body length 80.6 cm, tail length 29.7 cm, and
weight 19 kg; at 3 years old, on average (75)—body length 90.5
cm, tail length 31 cm and weight 21.2 kg; at 4—5 years old on
average (17)—body length 101.5 cm, tail length 31.6 cm and weight
23.8 kg. Later, dimensions and weight gradually and slowly grow
and the fully adult animal often weighs from 23.0-36.4 (37) kg.
Males may attain weight of 40 kg, while weight of adult females
is 20-25 (30) kg. With age, relative tail length decreases, consti-
tuting 35% of body length in recently-born animals; in 4—5-уеаг-
old animals—about 31%; and in 9-year-old and older—about 25%
(Barabash-Nikiforov, 1947, 1962).
Os penis length 150 mm, width at base 15 mm (У.Н.).
1343
891 Systematic Position
Only species in the genus.
Geographic Distribution
Islands and in part shores of northern part of Pacific Ocean from
approximately 62—64° М. lat. in the north, to Tropic of Cancer (in
east [Pacific]) and 40° М. lat. on the south (in west [Pacific]).
891 Fig. 333. Range of the sea otter or kalan, Enhydra lutris L. in the USSR.
Solid line—present range; broken line—reconstructed. Cross designates local
settlement on Moneron Island. V.G. Heptner.
892
1344
Geographic Range т the Soviet Union (reconstructed mainly ac-
cording to materials of Snou, 1902; Tikhenko, 1914; Barabash-
Nikiforov, 1947; Uspenskii, 1955; Gribkov, 1963).
This occupied the southern coast of Sakhalin (apparently, Aniv
Bay and somewhat to the north of Cape Aniv along the Okhotsk
coast)* and the whole Kuril chain from the southernmost islands
(Shikotan and Kunashir) to Shumshu and Atlasov.
In Kamchatka, along its western (Okhotsk) coast, sea otters
even in the 18th century, apparently, were present only in the
extreme south and did not extend north beyond the mouth of the
Igdyg River (Ozernaya) and the Kambal’ and Kitov capes (Steller,
1753, mentioned that in the “Penzhin River” sea otters were ab-
sent). In any event, they did not live in Shelekhov Gulf, or perhaps
only very rarely intruded there (Steller’s data, given below, con-
cern only the eastern shore of the peninsula). Frozen sea and fast
shore ice excluded their existence.
Along the eastern shore of Kamchatka in the “Otter Sea”, in
the first half of the 18th c[entury], kalans were distributed be-
tween 50° and 56° М. lat. chiefly from Cape Lopatka to the Kronotsk
Peninsula (Steller, 1753), more precisely, to Cape Stolbov (a little
north of the latitude of Bering Island—about 56° N. lat.). Accord-
ing to less definite information, in the past (18th and first half of
19th c[entury]) they extended northward to 60° М. lat. i.e. approxi-
mately to Olyutor Bay and even to Cape Faddey and the mouth of
the Anadyr’ [river], in other words, approximately to 64° N. lat.
However, information concerning its occurrence at 60° N. lat.**
and more to the north, is doubtful, or related to occasional trans-
gressions of drifting.
A separate section of occurrence, not connected with the main-
land portion of the range is found in the Commander islands, situ-
ated at approximately 180 km in the open sea from the nearest part
of the Kamchatka coast. It is also isolated from the other part of
the range which occupies the Aleutian islands—about 350 km from
Attu Island.
During the 18th, and especially the first half of the 19th
c[entury], there was very intensive, entirely unregulated hunting of
the animal throughout the whole range within, as well as outside,
the boundaries of the USSR. The highly prized fur of the sea otter
*Not shown on map, Fig. 333—Sci. Ed.
**Contradictory—Sci. Ed.
1345
was, аз is known, one of the main stimuli for taking possession
of the northern coast of the Pacific Ocean, at first, by individual
Russian explorers and later by the “Russian-American Company”.
In the second half of the 19th c[entury], after the Company ceased
its activities, and in some places much earlier, the animal was
practically exterminated in the greatest part of its region of occur-
rence in our country. It completely disappeared from the continen-
tal coasts and Kamchatka, except Cape Lopatka. On the eastern
coast of Kamchatka, in Kronotsk Bay, the last time kalans were,
observed, apparently, was in 1852 (Ditmar, 1901).
At the beginning of our [20th] century (up to 1943), otters
temporarily disappeared completely, apparently, from Lopatka, or
their number decreased so greatly that information of their exist-
ence there was questionable. However, in 1943, there were about
300 individuals (Averin, 1948); they are there now at the present
time (beginning of the 60’s).
On the Commander islands, as a result of the senseless harvest
in the 18th c[entury], it is considered (Marakov, 1964) that kalans
were completely exterminated and were absent here for about 100
years—until the 70’s of the 19th c[entury], when they were ob-
served again. It is supposed that they resettled on the Commander
islands across the sea. This is, however, quite doubtful. At least
the present situation of populations of Mednyi and Bering islands
points to the settled mode of life of the Commander kalans and to
the low probability of their surmounting the large expanse of open
ocean. On the first island, the otters are numerous and on the
second they occur individually, although the islands are separated
by a strait whose width is all of 24 km. Therefore, it is more
probable that a very small number of animals was preserved from
extinction on the islands and they, under conditions of complete
prohibition of their pursuit, had the possibility of reproducing again.
In any event, at the beginning of our [20th] century, there were
already about 400 (Suvorov, 1912). Afterwards, they were mainly
preserved because in this place, long before the Revolution, the
state hunting of sea otters and fur seals was organized. On Mednyi
Island, animals were always more numerous than on Bering Island.
Otters had already disappeared from southern Sakhalin very
long ago, and at the present time are absent there. On the Kuril
islands, otters were from time immemorial, as is said, always present
everywhere. However, pursued both by Russian and also many
893
1346
foreign fur companies in the 19th c[entury], otters quickly
decreased in number and their distribution was reduced. Their num-
bers continued to decrease in the 20th c[entury]. On the brink for
two centuries, in part at the beginning of our century, they, al-
though rare, occur on a quite considerable number of islands—
Shumshu, Paramushir, Onekotan, Kharumkotan, Ekarma, Matsuva,
Raseva, Ketap, Chirpo and Kunashir (Tikhenev, 1914; Uspenskii,
1955). On the other hand, in 1912, some authors believed that otters
on the Kuril islands disappeared almost completely (Зиуогоу, 1912).
Information concerning the 30’s (Barabash-Nikiforov, 1947), actu-
ally points to the complete absence of otters in the archipelago,
except for a few in Shumshu and between Onekotan and Urup.
Difficulty of access to many parts of the archipelago, espe-
cially the small islands of its middle part, permitted the animals, at
least in very small numbers, to protect themselves until the 40’s of
our century. In the very first post-war years it was established that
sea otters inhabited Paramushir, Onekotan, Shiashkotan, the Musiru,
and probably several other small islands of the middle part of the
chain (Solov’ev, 1945; Sergeev, 1947; Kuznetsov, 1949).
By the middle of the 50’s and the beginning of the 60’s, 1.е.,
within 10-15 years of initiating protection, the number of kalans
in the Kuril islands grew strongly and their range broadened; the
cause of this may be only partially correlated, owing to better
censusing and inspection. In 1955, the population of Kuril kalans
was in any event not less than 1500 individuals, and they lived on
Paramushir, Antsiferov Island (Shirink), Onekotan, Shiashkotan,
Rasshua, Simushir and Urup. South of Urup, animals were not
recorded (Klumov, 1957). After 1955, growth of the Kuril sea
otter population and the expansion of their range continued, and in
the mid-60’s, they occupied the whole archipelago, were more com-
mon south towards Urup (B.G. Voronov), and especially
common on Urup.
At the southern extremity of Kamchatka, sea otters were dis-
tributed (information from the 40’s and 1960) not only at Cape
Lopatka; along the Okhotsk side, they lived to capes Kambal’ and
Kitov, along the ocean north to at least Gavryushin Kamen’ Island
(Cape П’уа, Cape Zheltyi, Cape Inkanyushin, Utashud Island,
Gavryushin Kamen’ Island; Barabash-Nikiforov, 1947; Gribkov,
1963). All these places are located in southernmost Kamchatka,
south of 52° N. lat. and are associated with Cape Lopatka. Kalans
1347
have now spread, however, and are considerably more to the
north. Thus, in 1960, one animal was found in the delta of the
Avacha river in Avachin Bay, and another (in winter) at the former
Ust’ Kamchatka region at 57° М. lat. i.e. north of this place, which
Steller (1753) referred to as the [range] limit. The latter animal
was caught in the river 20 km distance from the sea shore (Gribkov,
1963).
At the end of the 50’s, kalans were introduced to Moneron
Island, lying approximately 50 km west of the southwestern end of
Sakhalin. The animals at first survived, but later died (G.V.
Уогопоу).
Therefore, the present range of the kalan occupies the Com-
mander islands, the extreme southern end of Kamchatka (Cape
Lopatka and adjacent places) and all the Kuril islands. The rough
number of animals apparently exceeds 4000 individuals.
Geographic Range outside the Soviet Union
Along the Asiatic coast, kalans were encountered on the coast of
Hokkaido (Yeso), apparently, northern, eastern and southern and
at the northern end of Honshu (Hondo, Nippon). These may in part
be wandering animals (Temminck, 1847; reference to northern shore
of islands).
In the eastern part of the Pacific Ocean, sea otters were present
along the entire Aleutian Island chain, in the Pribilof islands and
along the American coast and adjacent islands from approximately
60° N. lat. (almost from the mouth of the Yukon—at least along
the shore of the Alaska Peninsula) southwards to the middle part
of the California Peninsula [Baja California] (Sebastian Vizcaino
Bay at Cedros Island, about the latitude 28° N. lat.; Hall and Kelson,
1959) and, perhaps, even Guadalupe Island. There is information
on their occurrence to the southern extremity of [Baja] California,
i.e. to the Tropic of Cancer.
In the beginning of our century, in all American sections of the
range, kalans were destroyed almost everywhere and were met
with in very small numbers only along the Aleutian chain and
perhaps individually in the San Francisco region* and in other
separate places. In San Francisco, where sea otters were consid-
ered to already have been extirpated long ago, they unexpectedly
*Actually, Monterrey and Big Sur, south of San Francisco Bay—Sci. Ed.
894 Fig. 334. Reconstructed range of the sea otter, ог kalan, Enhydra lutris L. оп
the coast of North America. V.G. Heptner.
appeared in the 30’s in quite considerable numbers (Bolin, 1938;
Fisher, 1939). Thanks to their protection, the population of otters
in America quickly grew; at the beginning of the 60’s they were
already quite significant. Consequently, their range has also wid-
ened. The question now concerns organizing utilization of the
animals (V.H.).
Geographic Variation
Geographic variation in the kalan is poorly understood. Collection
materials of skins in museums are very few because of their very
high price, and these skins are often old and their color has changed.
Conceptions of species variability are, therefore, based mainly on
craniological features and only partially on color and fur characters.
895
1349.
Within the USSR, two races may be distinguished’.
1. Commander sea otter, or kalan. Е. [. lutris Linnaeus, 1758
(syn. marina).
Fur color relatively light, with significant development of gray
hair and quite long guard hairs (30 mm long on the average, in
middle of back). Skull fairly large, moderately wide. Zygomatic
width constitutes 71-80% and mastoid, 71-79% of condylobasal
length.
Condylobasal length of skull of males (17) 15 130.5—M 136.5—
140.3 mm, of females (7), 120.0-M125.4-128.2 mm; zygomatic
width of males is 101.0-М 104.3-110.0 mm, of females, 90.0-М
94.3—102.0 mm; interorbital width of males is 40.2-М 43.4—46.5
mm, of females, 36.7-М 39.2-42.2 mm; mastoid width of males is
93.9-M 99.5-108.0 mm, of females, 88.0, М 91.4-95.3 mm; length
of upper toothrow in males is 46.2-М 48.3-51.4 mm, of females,
44.3, М 46.0-48.6 mm.
Greatest skull length is 144.2 шт on average (Barabash-
Nikiforov, 1947).
Commander Islands.
Outside the USSR, Aleutian and Pribilof islands, and on the
American coast, from the northernmost place of occurrence south
to Vancouver Island inclusive.
2. Kuril otter, or kalan. Е. [. gracilis Bechstein, 1799 (syn.
crientalis, stelleri, kamtschatica).
Fur relatively dark, with weakly-developed gray hair and a
relatively short guard hairs (on middle of back, about 25 mm long
on average).
Skull somewhat smaller, wider than in nominal form and some-
what flattened. Zygomatic width constitutes 85-87%, mastoid—
84-86% of condylobasal length.
Condylobasal length of skull (5 old males; Kamchatka) is
132.8-M 133.4-133.6 mm; zygomatic width is 113.3-M 114.9-
116.4 mm; interorbital width is 46.9-М 48.0-49.1 mm; mastoid
‘Characteristics of the races after Barabash-Nikiforov, 1947 and Stroganov, 1962.
Description of fur of different races by Barabash-Nikiforov, 1947 is, however, slightly
contradictory. In contrast to other authors, he indicates (p. 26) a “tawny sheen” on fur
of Kamchatka animals, giving fur even a “reddish” color, compared to fur of Com-
mander animals. On the other hand, he refers to a “darker tawny tint” of fur of
Kamchatka animals. The review of Stroganov (1962) does not give anything new and
only complicates the picture (it refers to the existence of “not less than five” races of
the species). From this work, if not with reservations, data on dimensions and propor-
tions are taken.
1350
width is 107.0-М 109.5—112.3 mm; the length of upper toothrow
is 46.2-М 46.6-47.0 mm.
Greatest length of skull is 137.3 mm on average (Barabash-
Nikiforov, 1947).
Kuril islands, Kamchatka, and in the past, in South Sakhalin.
Outside the USSR—in the past, northern Japan (Hokkaido,
and possibly northern Honshu).
The attempt to separate Kuril and Kamchatka sea otters into
separate races (Stroganov, 1962), must not be considered justified.
Those geographical considerations on which this was based do not
give any basis for this. There is no isolation between kalans inhab-
iting different parts of the chain; nor is the Kamchatka population
isolated from the Kuril. Sea otters not only drift with currents and
storms (to which even Steller, 1753 referred), but they themselves
undertake quite large migrations and undoubtedly move from one
island to another. Apparently, the populations of Cape Lopatka and
Shumshu Island communicate with each офег”. In particular, this
is inferred from information given above concerning the “disap-
pearance” and “restoration” of the Lopatka population during the
last ten years. It is also known that in the pre-war years, Japanese
kalan hunters tried more than once to “drive” Lopatka animals to
Shumshu using various means.
Kuril kalans were once considered the very best and were es-
pecially valuable. Kamchatka were also esteemed more highly than
Commander [otters]. This correlation of values corresponds to ra-
cial difference between Kuril-Kamchatka and Commander [otters]
and to a certain degree, indicates the isolation of the Commander
population.
Outside our country, only one well distinguished form is ap-
parently known: Е. [. nereis Merriam, 1904—the North American
coast south of Vancouver Island (V.H.).
Biology
Population. In the second half of the 18th century, populations of
kalans were still significant in some places. Concerning this one
may judge from the fact that on the Pribilof islands, in 1787, more
than 5000 kalans were caught. Later, their number greatly decreased,
‘Width of the First Kuril Strait, separating them “reaches 10 miles, but free
passage is reduced to 3.0-3.5 miles due to reefs present on both shores” (Sergeev,
1947).
896
135
from killing and in 1821 the harvest was prohibited, and by the
40’s they completely disappeared here. During 1763-1764 on the
Andreyanov islands, about 3000 kalans were taken, and during
1775-1780, on Urup (Kuril islands)—1170. After the strong earth-
quake of 1780 kalans disappeared here. With renewal of harvest-
ing on the Kuril islands, during 1828-1830, 2600 animals were
caught on only Urup and Simushir. For 1842-1860 on the Kuril
islands, 4510 individuals were taken overall, i.e. an average of
only 250 animals per year. In 1745, on Bering Island more than
1500 kalans were captured, and from 1747-1749, an additional
1500. As a result, for the period 1754-1755, a total of 5 kalans
were caught, and in 1756—not one. In 1762, 20 head were taken
and after that, the harvest was suspended for a long time in view
of the trifling results—kalans were very few. Therefore, on Bering
Island during the second half of the 18th century, the stock of
kalans was slightly more than 3000 head. With depletion of the
otter stock on Bering Island, the main harvest was shifted to Mednyi
Island. Here, in 1754, no less than 790 otters were caught, and
during 1760-1763, no more than 20. Here, probably, the popula-
tion was half that on Bering Island. Overall, apparently, only 4000-
5000 of these animals inhabited the Commander islands, since there
is information that the quantity of kalans on Mednyi Island
attained 2000. From these uncoordinated figures (complete data
for whole range are absent), the conclusion may be drawn, that at
the beginning of the 18th century, the kalan population consisted
of approximately 20-30 thousand (possibly slightly more) in the
whole range—from the northern coasts of Japan to the shores of
[Baja] California. In the latter area, during the past century, 7000
skins were bought from the native hunters in only one year.
In order not to undermine the basic stock, not more than 2000—
3000 animals could be harvested every year throughout the whole
range. In the period beginning in 1870, the world take of kalans
fluctuated within the range of 4000 to 8000 head, on average 5000
head per year. Therefore, the entire time harvest was always at the
partial expense of the basic stock, and the result was that after the
harvest of 8000 head in 1884, it steadily decreased; in 1900, it was
reduced to 1000 head. This decrease continued until the nearly
complete liquidation of legal harvest in the beginning of this cen-
tury. Within more than two centuries, otters were almost destroyed
due to their rapacious exploitation.
897
1352
On the Commander islands, after the period of almost com-
plete disappearance of kalans, they appeared again in the 70’s of
the previous century, but only on Mednyi. The main stock of Mednyi
Island settled mostly on the northern (northwestern) extremity of
the island. Disturbed by a take of living animals in 1937-1938, the
herd migrated to the southeastern extremity (П’1па, 1950). Ten years
later, kalans returned again to the northern extremity of
the island. If in 1931-1932, 282 kalans were counted in the waters
of Mednyi Island, then in 1952, 250 animals were recorded
(O. Danilov), and in 1955 about 350 (Marakov, 1964). Therefore,
despite the absence of harvesting due to some reason which were
difficult to eliminate, the Commander stock of kalans hardly grew
during the last 20 years.
In 1924, on both Commander Islands, 400 individuals were
counted; in 1939—800; and in 1957—500 (Nikolaev, 1958). Dur-
ing the last decades, the situation sharply changed, and the kalan
population began to grow quickly—in 1964, on Mednyi, the number
of adult animals alone reached 1000 (Marakov, 1964). As shown
above, the population of sea otters on Bering is very small; they
Fig. 335. Habitat of kalan on Mednyi Island. Photograph by S.V. Marakov.
1353
are encountered individually. It follows that growth in the number
of animals is correlated to their reproduction locally—the possibil-
ity of their immigration from outside is, in practice, very improb-
able (Marakov, 1964). It must be noted that recent methods of
counting still allow significant discrepancies between data of vari-
ous authors. Thus, in 1931, according the Commander state fur
farm put the number at 282, but from another source of data
(Barabash-Nikiforov, 1948)—about 500.
Kalans of the Kuril chain and Kamchatka constitute one popu-
lation, which is characterized by regular seasonal migrations
(Dobrovol’skii, 1936; Nikolaev, 1958). In the Kuril chain, there
were 200 in 1912; in 1924—600, and in 1939—800 kalans. By
autumn of 1958, north of Iturup there occurred about 1900-2200
animals living; and by 1961, the population increased to 2900—
3100 head. On the islands of the archipelago inhabited by kalans,
they were most numerous on Urup—938 animals in 1958, and
1700—in 1961, and on Paramushir where in 1958, 614 individuals
were recorded. In the remaining cases, the number of recorded
animals did not exceed 79 (1958). Relatively more animals were
observed in the waters of Shikotan, Ketoy and Simushir islands
(Nikolaev, 1958, 1964). On Kamchatka in 1910, about 306 sea
otters were censused, in 1924—400, and in 1943—again only 300
(Averin, 1948). The total number of our kalans at the beginning of
the 60’s, apparently, exceeded 4000.
On the islands of the Aleutian chain and the shores of the
Alaska Peninsula in 1912, there were about 200 otters; in 1924—
200—250; in 1958, thanks to rapid growth of the population, the
number had already reached several thousand. In the waters of the
California coast (36° parallel and south), in 1938 in all 94 were
censused; but in 1958, already there were about 500. According to
the most recent data, along the coast of the state of Alaska there
are 20-30 thousand otters. It is calculated that on the islands of the
Aleutian chain, in particular on the Rat, islands, the kalan popu-
lation has already reached its density limit (Kenyon, 1961). In
1962, with examination of six main sections of occurrence from
the air, 11000 kalans were counted (Brooks, 1963). Attempts to
settle these animals on the Pribilof island have still not succeeded.
Proceeding from the given data on America and paying atten-
tion to information on the animal’s population in the USSR
and in California, the total number of kalans may be determined as
898
898
1354
Fig. 336. Biotope of kalan. Babichii Pod’em Bay оп Mednyi Island. Photo-
graph by S.V. Marakov.
25—35 thousand head. Apparently, this species’ extinction, antici-
pated not long ago, no longer threatens it. Protection of the sea
otter is one of most remarkable in the field of nature protection.
Habitat. Habitat of the kalan is characterized by steep rocky
shores, barrier reefs, submerged and exposed rocks, continually
washed over by waves, as well as vast thickets of sea weed
(species of Alaria and Laminaria—“sea cabbage’’) extending as an
interrupted belt along the shore. The latter grow strongly in sum-
mer, but are destroyed by storms in winter. Everywhere where they
exist at the present time, kalans prefer the extremities of islands
and peninsulas. Here the animals have the possibility of moving
around to the lee side in stormy weather.
Condition of the water medium, feeding and climate are in
total more important for kalans than the character of the coastal
belt itself. Their inclination toward inaccessible coastal zones de-
899
1355
veloped as a result of savage pursuit. In structure and type of
adaptation, the animal is not only littoral but also pelagic, adapted
to living under conditions of the northern part of the Pacific Ocean.
This is a marine climate, with mild winter and cool summer, and
fog, frequent drizzle and entirely variable, but occasionally severe,
winds especially in winter. These winds, and especially storms,
are very unfavorable for kalans. The small difference
between air temperature and that of the upper layer of water seems
to be a positive factor, especially in summer. The difference in
water temperature during a year on the Commander islands is only
15°C in all (from—1.5°C in January to 13°C in August). In the
Kuril chain, water temperature fluctuates from—1.8°C in January
to 5—10°C in the northern islands and 10—20°C in the southern
islands. The lag of the period of greatest coolness at a depth of 20
m and more, has large significant in the ecology of marine animals
serving as food for kalans. During experiments on acclimatization
on the Murmansk coast (Reshotkin and Shidlovskaya, 1947), kalans
suffered some from summer heat and mosquitoes.
Food. On Mednyi island, two species of sea urchin serve as the
main food of kalans; bivalve mollusks are of significant impor-
tance—octopuses are less so. In numbers, crabs of several species
constitute 10%, but fishes—6.7%, of which cod, lumpfish
[Cyclopterus lumpus], Pacific capelin, sand eels [Ammodytes] and
sockey salmon predominate. Altogether, 28 food components were
established. When held in captivity, the kalan reveals great frugal-
ity in feeding. Under natural conditions, its food regime is fairly
monotonous (Barabash-Nikiforov, 1947).
The chief characteristics of the animal’s nutrition in the au-
tumn—winter period on the Commander islands consist of the fact
that at this time there is a sharp fall in the proportion of crabs
(from 24 to 4%) and fish (from 22 to 6%) which migrate at that
time from the shores into great depths of water. The proportion of
mollusks slightly decreases, from 50 to 45%. Sea urchins remain
the main food component (100-98%). On the Pribilof islands, sea
urchins also constitute 92% of the kalan’s food (Murie, 1940). This
is also confirmed by observations on otters in nature, on Mednyi
Island (Barabash-Nikiforov, 1947). On Urup (Kuril chain), in
February—March, kalans in the western (Okhotsk) coast feed on sea
urchins, polyplacophore, gastropod and cephalopod mollusks, crus-
taceans, starfish, fish and algae. Along the eastern coast at this
1356
=.
Fig. 337. Zapalata Вау on Mednyi Island—place of residence of kalans. Photograph
by S.V. Marakov.
time, crustaceans occupy first place, various mollusks—second,
and then—sea urchins and algae (Nikolaev, 1958).
In captivity, the daily [dietary] norm of the kalan constitutes
4.5—5.5 thousand calories (Reshotkin and Shidlovskaya, 1947).
According to other data, for the average sea otter, it is 6.5-7.5
thousand calories, but for large animals—even 10 thousand (II’ina,
1950). Proceeding from the fact that the quantity of sea urchins
eaten by a kalan in one day contains only a total 3 thousand large
calories, some (Il’ina, 1950) consider that, in freedom, fish must
predominate in the kalan’s food, and kalans cannot feed on sea
urchins alone. In captivity, food of the kalan consisted of fish
(66.8%) and sea urchins (33.1%). As concerns digestibility, the
fish proportion increases to 80.3%, and sea urchins drops to 19.6%
(Il’ina, 1950). It is impossible to consider that the question of
kalan nutrition has been sufficiently clarified.
Home range. Kalans do not reveal an attachment to any sort of
definite place. Individual animals and groups of them appear
irregularly in first one place and then another; however, true
2357
900
Fig. 338. One of the groups of kalan of Mednyi Island. Photograph by $.У. Marakov.
migrations are only observed in separate individuals and pairs.
Periodic redistribution of the kalan populations in the region of
their permanent habitat which happens locally, depends first of all
on the weather. With a wind of velocity 4, several concentrations
of them are observed at the shore. This becomes very obvious with
wind velocity 7. In calm summer weather, separate kalan groups
stay permanently at a distance of 18 and more kilometers from
shore, where there are shallow places (banks), with beds of sea
cabbage [kelp]. Here they spend the night. In winter, kalans gather
themselves at the shores, since the kelp beds are destroyed by
storms. In the open sea, kalans may be met with at a distance of
18—27 km from shore, but they are capable of reaching consider-
ably greater distances. The appearance of killer whales* (which
usually cause otters to flee in panic), human activity and the ex-
haustion of main foods, are reasons for local regrouping of kalans;
such was the case in 1938, when on Mednyi Island sea urchins
almost completely died out following some sort of epizootic
(Barabash-Nikiforov, 1947).
*Russian name is “КозаЖа”, misspelled “kasatka”, or swallow, in Russian origi-
nal—Sci. Ed.
1358
Among kalans of the Kuril chain are distinguished local ones
which are permanently present in one and the same place as
migrating ones.
In summer, kalans here completely confine themselves in defi-
nite sections of the shore, although they sometimes move for tens
of kilometers in search of food and protection from storms.
Burrows and shelters. For resting, kalans utilize sections of
rocky laida, narrow coastal strips under steep coastal cliffs,
especially in times of winter storms. For the same purpose, they
very frequently use the emergent and submerged rocks (“hiding-
places*”’) at various distances from the shore. In summer, otters
also frequently use dense kelp beds, which are sometimes consid-
erably far away from the shore, for resting and protection from the
attack of killer whales.
Daily activity and behavior. The kalan is an animal preferably
active during the day. Its activity begins at dawn or shortly before
it, and ends at twilight. At night, the kalan is active only on rare
occasions, usually on moonlit nights. In winter time, the animals
901 spend the night on shore and eat the entire day without an interval
Fig. 339. Kalan lying in water, May 1960. Photograph by S.V. Marakov.
*Russian word is tainiki—Sci. Ed.
1359
in the activity for resting. In summer, they feed mainly in morning
and evening, resting from 11-12 to 15-16 o’clock. In this time,
the animals stay in the algae beds, gathering in herds of 20-120
head. Old otters, sometimes “twist”, wrapping themselves in algae
and appearing as if anchored (Fisher, 1939).
The degree of activity of the kalan depends upon the
provisioning of the given region with food, the weather conditions
and the state of the sex glands. Animals are more active in regions
with small food supply. Males also manifest increased activity in
search of females in estrus. Before storms, activity of kalans also
grows in connection with relocation to protected places. The kalan
feeds several times in the course of a day. After satiation, it cleans
itself, sometimes plays, and then rests in the water, lying on its
back. In the open sea near shore, the kalan may feed even with
wind strength of 6-7.
In the sea, the sea otter is agile, mobile and quite quick.
In searching for food, it moves at a speed of 5-6 km/h, but its
maximum speed equals 12-16 km/h (A.M. Nikolaev). Capture of
902 Fig. 340. A group of resting otters on the rocks of Glinka Bay, Mednyi Island,
December 1958. Photograph by S.V. Marakov.
902
1360
bottom and swimming fish is with its teeth and fore paws, then
clasping them to its chest. Coming to the surface, the animal lies
on its back with it and in such a position eats it. This pose is very
characteristic for the animal in feeding and resting. After taking
food, the kalan cleans itself of the remains by revolving around on
its [body] axis. After resting, kalans often gambol, play with each
other and somersault in the water: The kalan is a herd animal, and
peaceful for the most part with respect to each other. However,
fights between males for females occur in places. On land, kalans
move quite slowly, strongly bending their back while walking, like
the river otter. Droppings on land with abundant remains of sea
urchin tests and remains of crabs, mollusks and fish serve as a
characteristic feature of the kalan’s residence.
Seasonal migrations and transgressions. Until recent times, it
was considered that kalans do not perform significant and regular
migrations. This idea was based on observations of herds of the
Commander islands. In actuality, a significant part of the Kuril
(perhaps Kuril-Kamchatka) population gradually translocate them-
selves from wintering places on Iturup and Urup islands northward
to a place of spring residence on the middle islands of the chain—
Brouton, Onekotan—where the animals spend March and April.
Then they move farther to summer (May—October) on the northern
islands—Paramushir, Shumigu, Aland and perhaps to the
Kamchatka coast.
In autumn, a local reverse migration to the south takes place.
Autumn migration of kalans from the northern islands of the chain
is associated with decrease in water temperature toward —1.8°C,
which causes displacement of the bottom fauna to deeper places,
complicating food capture. Another cause leads us to think it is the
appearance of ice which hinders movement and food-catching. At
the end of January, cooling of water to —1.8°С and the appearance
of ice drive the kalans away from the shores of Iturup Island and
in March, also from the Urup coast. The coastal waters of the
middle islands remain free from ice in March—April.
In the time of migration, kalans swim singly and in pairs at
intervals of 10-15 min[utes]. During migration, all animals se-
curely hold to a definite direction, neither stopping nor paying
attention to local kalans. Nevertheless cases occur of single indi-
viduals (possibly young) who lose their way and return (Nikolaev,
1958).
903
1361
In addition to seasonal migrations which do not include the
entire kalan population, more than once there was observed trans-
location of a herd for a considerable distance, after which the
animals reappeared in the abandoned places, sometimes only after
a series of years. Such was the case when kalans disappeared
from Urup after the earthquake in 1780, and also a series of cases
when the disappearance of kalans in one region was accompanied
by sudden growth in the population of another. In the 40’s of the
previous century, otters disappeared from the shores of Simushir
Island after the eruption of Prevo volcano. At the same time, they
appeared on Shumshu, Rasshua and Ushishir islands, where they
os cis
Fig. 341. Females with young and pregnant female otters. Mednyi Island, Zapalata
Bay. 6 August 1962. Photograph by S.V. Marakov.
1362
of a Japanese ship. Finally, it is sometimes believed possible
(Barabash-Nikiforov, 1947), to connect the secondary appearance
of sea otters in the 70’s of the last century on the Commander
islands (after their complete or almost complete destruction there
by the end of the 18th century) with increase of the harvest at that
time on the Aleutian and Kuril islands, which might have caused
their resettling on the Commander islands. As shown above, this
idea is lacking in foundation. There is the suggestion that all kalan
populations, with the possible exception of the isolated California
group, constitute one chain, within the boundaries of which
displacements and regrouping can, apparently, occur.
Reproduction. The kalan, apparently, attains sexual maturity in
the third year of life, but this is not accurately established. The
kalan has no definite estrus (rut) period. Nuptial games, mating
and new-born young with the females are observed at any time of
the year. On Mednyi Island, nuptial play was somewhat more often
observed in spring—March—May, but it was also observed in July—
August. On the Kuril islands, the mass parturition proceeds in
Spring, but new-borns are also observed at other times of year
(Nikolaev, 1958).
Fig. 342. Female kalan with the new born on its chest. Mednyi Island. Photograph by
S.V. Marakov.
904
1363
The duration of pregnancy is determined as 8-9 months; in
one case, it lasted 238 days. The female, as a rule, bears one cub.
In very rare cases, two embryos were found; still rare—two cubs
with a female.
Mating takes place in water and lasts 3-9 min[utes]—parturi-
tion—on the shore or on rocks flooded with water. Delivery lasts
about two hours. In several cases, the cub of the previous litter,
already almost equal to her in dimensions, is still with the parturating
female. Clearly-defined [mated] pairs are absent among otters.
The annual increase of the kalan population on Mednyi Island
was determined in the 20’s—30’s of the present century as 7% or
less (Barabash-Nikiforov, 1947). These figures characterize the
tempo of growth of a herd strongly undermined by predatory har-
vest. Rate of annual growth may be judged by the percent of
yearlings in the population. According to data of the autumn count
of 1958, on Paramushir, the growth was 10.5%, Onekotan—6.6%,
Shiashkotan—14.0%, on Ketoi and Simushir 1$1ап9$— 8.5%,
Urup—7.8%. Therefore, the probable annual growth of the Kuril
Fig. 343. Otters going to water. Mednyi Island. May 1958. Photograph by
$.\У. Marakov.
905
1364
population fluctuates within limits of 6.6 to 14.0% (A.M. Nikolaev).
Average equaled 9.2%, which is somewhat higher than earlier
figures given. It is undoubted, that in the thriving period of the spe-
cies, natural increase is higher—within limits of 10 to 20-25%.
Growth, development and molt. Immediately after birth, body
length of new-born is 50-56 cm, and weight, 1.4-1.6 kg. Kalans
are born able to see, with juvenile fur and having 26 milk teeth.
In the first days after birth, they are barely active, and lie on their
backs on the shore or in water. First attempts to swim independ-
ently are made at the age of about 2 1/2 weeks, and by the third
week, they are already able to swim for short distances. The juvenile
(infant) fur is replaced by permanent at the age of about 6-7 months.
At this time, replacement of milk incisors and first premolars by
permanent is completed. Replacement of second and third premolars
begins, and the molar teeth erupt. Total number of teeth is already
32. Body weight of such a “koshlak” reaches 9-11 kg, body length,
1.10-1.15 т. At one year, live weight reaches 17-19 kg and body
length, 1.25—1.35 т. Replacement of premolar teeth is completed,
but skull crests are still weakly developed (Il’ina, 1950).
Enemies, diseases, parasites, mortality, competitors, and Dy-
namics of number. Among animals, the most dangerous and almost
the sole enemy of kalans is the killer whale which often manifests
itself in otter habitats. The polar shark is rare and is mainly found
at great depths. Old information, that Steller’s sea-lions and fur
seals also appear to be enemies of the sea otter, have not been
confirmed by more recent observations. The largha seal is a com-
petitor of the kalan as regards main food components, and in re-
spect to shelters, Steller’s sea-lions, fur-seals and other pinnipeds
are competitors. Several of the sea birds are shown to be competi-
tors, in small degree of the kalan. However, their activity has no
fundamental effect on food resources of the kalan. Natural mortal-
ity of kalans is very great. Causes show up as age limits, diseases,
injuries caused by killer whales and, possibly, sharks; rockfalls on
to shore laidas; the breaking of ice (Nikolaev, 1958), and in indi-
vidual cases, the effect of storms and birth pathologies. Age limit
of the kalan is not known. Judging by the relationship between the
intensity of reproduction, animal dimensions and duration of life,
in kalans it must be considerable.
There are no reliable data concerning population dynamics of
otters. For the last 200 years, the human role was so great that
other factors were overwhelmed and could not be estimated.
906
1365
Besides direct destruction, the movement of ships in the re-
gion of their habitat and frequent frights caused by other human
activities have an extremely negative effect on the numbers and of
kalans and their colonies on the coastal belt.
Practical Significance
The economic significance of the kalan is very substantial. This
now rare animal provides beautiful, warm and durable fur and is
considered the most valuable fur-bearing animal. Individual otter
hides have been sold at the London auction for a price of $2000.
The value of the fur is doubled by the rarity of the animal: for the
past 80-100 years, not more than 8000 hides were sold per year,
and in the past 30 years—a negligible quantity (single ones), and
more or less accidentally.
In the USSR, the harvest was prohibited in 1924. This prohi-
bition continues until the present with the aim of future growth
and dispersal of herds. Because of the low reproductive rate of
kalan and conditions not yet entirely favorable for them; at the
present time, conditions for restoration of initial population re-
quire sufficiently more time. At the present time, the question has
been raised concerning a possible catch of a limited number of
otters, especially the sick and old, for skins (in nature, mortality in
this species is relatively very high: Marakov, 1964). Along the
American coast, where the density of otter colonies locally has
already reached its limit, a test harvest was conducted in 1962 and
174 animals were caught (Amchitka Island); in the following year,
there was planned a harvest of up to 300 individuals for determin-
ing the possible price in the fur market (Brooks, 1963).
Several perspectives may be borne in mind concerning cage
rearing of otters; experiments to keep them in captivity have been
relatively successful. However, feeding the animals on their natu-
ral food in captivity is very complicated, since preparation of this
food which is needed in large quantities for each animal is diffi-
cult. The experiment of the keeping animals in open-air confine-
ment on the Murmansk coast of the Barents Sea, may be also
considered partially successful.
The main means of catching otters in the shutter net, and in
summer, the landing-net. At the epoch of abundance of this ani-
mal, harvesters caught it on land by hitting it on the head with a
stick (P.Yu.).
907
Supplement to the Order Carnivora
Superfamily of Wolves
Superfamilia Canoidea Simpson, 1931
Family of Raccoons
Familia Procyonidae Bonaparte, 1850
Predators of small, moderate and large dimensions, in part gener-
alized, and in part specialized and highly specialized types’.
Limbs are pentadactyl, plantigrade and semi-plantigrade; digits
separated to different degrees, the third—longest; claws in major-
ity of cases non-retractile, in a few, semi-retractile (Ailurus, panda;
Bassariscus, cacomistle). The lower surface of foot and hand usually
bare.
General form of skull variable—in some species facial region
elongated, in others, skull short and rounded; in still others, whole
skull strongly elongated. However, in the majority, braincase rela-
tively, sometimes very, voluminous. In majority of species, ali-
sphenoid canal is not developed, canal of carotid artery (can.
caroticus) located at inner side of auditory bulla and separated
from posterior lacerate foramen (for. lacerum posterius).
Bre ee
Dental formula in majority is LC hag 590% in some,
Bi 4a Е, ) В AA
ЕЕ (Ailurus, red panda) or а, ам Fee
и ре аи
(Potos, kinkajou ог monkey-tailed “bear’’), ог Г.С т Р 3 M Amal
(Ailuropoda, bamboo bears or giant panda).*
‘Characteristics of the family, and later of the genus, are given briefly.
*Now recognized as a true bear, not a procyonid—Sci. Ed.
908
1367
Form of cheek teeth and character of their masticatory зиг-
faces extremely variable. In one, they are powerful and broad with
a flat multicusped, bluntly tuberculate masticatory surfaces, typi-
cal of omnivorous and phytophagous carnivores (extreme form is
Ailuropoda, bamboo bear). Teeth in several are characteristic of
meat-eating carnivores, of carnassial form with cutting crests. The
fourth upper premolar and the first lower molar are transformed
into carnassial teeth, resembling teeth of species of wolf family,
Canidae (Bassariscus, cacomistles). In majority of species, cheek
teeth of intermediate type—relatively broad and flat. Incisors rather
weakly developed, the first premolar has tendency toward reduc-
tion—small or absent, sometimes disappearing with age. Canine
relatively small.
The os penis is present and well-developed. Anal region de-
void of glands and glandular areas.
As regards general appearance and size, species of the family
are quite varied—with a small number of species in the group,
differences are considerably larger than in other families of the
order. One of the general aspects and dimensions 1$ bear-like
(Ailuropoda, bamboo bear), some are similar to cats (Ailurus,
panda), others—fox and raccoon-dog (Procyon, raccoons) or wea-
sel (Bassariscus, cacomistles) and some are quite individual (Nasua,
coati; Potos, kinkajou or monkey “bears”).
In some species, muzzle is short and obtuse; in others, it is of
the fox or Arctic fox type, in still others, with very long pointed
facial portion; in the majority of cases, ears quite large, rarely
short and rounded; eyes usually quite large. In majority of species,
tail long; in some, longer than trunk, covered with dense luxuriant
hair; in one species, very short (as in bears), in another, strong,
muscular and prehensile like a monkey’s tail (Potos, kinkajou; a
rare exception among carnivores). Pelage quite long and dense; in
some, very dense, fluffy and long (raccoon, providing valuable fur).
Color entirely variable—from more or less monotone yellow
or brownish to very mottled and bright, consisting of a combina-
tion of bright red with black (Ailurus, panda). In a series of spe-
cies, dark transverse rings are well set off against the light general
tone of the long, fluffy tail.
Differences in dimensions very great—smallest forms have body
length of about 25-30 cm and about 1.25 kg in weight (Bassariscus,
910
1368
cacomistles), and in the largest, head and trunk length is about 180
cm and the weight is about 150-160 kg (Ailuropoda, bamboo bear).
Species of the family are mainly in tropical, in part with warm
temperate climates, and only one (raccoon) extends quite far into
temperate zone, and in the north of its range even falls into winter
sleep.
Some species ascend high into mountains and are even com-
pletely associated with montane regions (Asia—see below). All
are, to a great or less extent, forest animals or associated with
forest regions, the majority closely associated with forest, climb
trees well and often, some live in tree hollows, etc. A series of
species are mainly, arboreal animals (semi-retractile claws) or even
specialized in climbing (monkey-bear, Potos). Some species are
very fast and active, others are languid and slow in their move-
ment. The majority are solitary nocturnal animals, forming only
temporary family groups. One species is partly diurnal and
sometimes forms large groups (Nasua, coati). The majority are
omnivorous animals, chiefly “gatherers”, feeding in addition to plant
foods, on invertebrates and lower vertebrates, small rodents, birds,
etc. Some are more predatory, a few are mainly phytophagous or
specialize strictly on plants (bamboo bear, feeding on young shoots
of bamboo).
The range of the family is divided into two parts—a vast
American and relatively very small Asiatic. In America, the range
occupies a considerable part of North America, except its northern
part; Middle America; and a large part of South America, except
its southern part. The northern borders of the range extend from
the Gulf of Saint Lawrence, through southern Quebec, southern
Ontario, the south of Manitoba and Saskatchewan, Alberta and
through the south of British Columbia (for details, see below;
description of raccoon). The southern border runs from the Pacific
coast of northern Chile (at approximately 20° S. lat.) to southeast
to the mouth of the La Plata [river] on the Atlantic coast. The
range includes Vancouver Island and several small islands lying
directly along the shores of the southern part of North America,
some of the Bahamas and Lesser Antilles islands (see below, spe-
cies description) and Trinidad. In Asia, the range occupies the
Himalayas in Nepal and Sikkim, and farther to the east upper Burma,
Yunnan and north to the border of Gansu (an entirely montane
region).
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911
13
The family Procyonidae in all of the diversity of forms admit-
ted into it, according to the present point of view, represents a
systematically diagnosed and well-defined group, which is suffi-
ciently connected by common origin. Actually, this is, in substance,
the broadly accepted opinion. The diversity of forms within the
family must, apparently, be considered as an adaptive divergence,
associated with feeding characteristics, and different degrees and
forms of adaptations to climbing.
At the same time, diversity of structures within the family, led
and sometimes still leads, to strong fragmentation. Separate au-
thors divide it into 2, 3 and even 7 separate families. In the latter
case, each well-defined genus is separated into a distinct family.
There has also been no lack of division of the family into a series
of subfamilies.
For example, several authors not only separate the genera Potos
(kinkajou) and Bassariscus (cacomistle) from the family, putting
them in the rank of separate families, but also entirely rejected the
idea of close relationship with remaining Procyonidae. The sys-
tematic position of the genus Ailuropoda (bamboo bears) has been,
and is, frequently treated in various ways. It was related to bears,
separated into a special family (alone or together with the panda,
Ailurus), and only relatively recently, it was again recognized as a
member of the raccoon family.*
Having gone far, the subdivision of the raccoon group, as with
anything to such detail, does not bring utility and only obscures
the. general picture of family structure and its relation to others.
Evidently, it is more natural to consider a single family,
Procyonidae, composed of two recent subfamilies (see below).
In the circle of families of the order, and the group Canoidea,
the family Procyonidae shows the clearest relationship to the bear
family, Ursidae (see characteristics of this family). Relationship to
the wolf family, Canidae, is regarded as much less close, although
some data also point to a certain relationship between these groups
(see characteristics of wolf family, Canidae, and of genus of rac-
coon dogs, Nyctereutes). Also, there appear to be no clear relation-
ships between the raccoon family and the marten family, Mustelidae.
In the series of families of the order, the raccoon family is usually
*Presently Ailuropoda is considered a bear—Sci. Ed.
912
1372
placed аз the immediate neighbor of bears and between bears,
Ursidae, and martens, Mustelidae.*
The raccoon family is less ancient than the wolves, Canidae,
and is associated, apparently, by its origin, with early forms of the
latter. The main—American—branch of the family split off in the
lower Miocene in the Americas, and throughout its entire history,
it was restricted, as at present, to South and North America, gen-
erally within the boundaries of the present range. The Asiatic group
is known from the upper Miocene, and throughout its entire his-
tory was much more widely distributed than in the present epoch,
and in particular, the range included not only several parts of Asia,
lying outside its present range, but also Europe. In the
zoogeographical sense, and partly also the systematic, the Asiatic
forms must be considered as relicts.
The family is divided into three subfamilies—one extinct
American (Cynarctinae) and two contemporary—Procyoniae, the
American raccoons and Ailurinae—the Asiatic raccoons or pan-
das. The total number of genera is 18, of which 8 are contempo-
rary. The number of described species of contemporary American
genera is quite great, however, most of them undoubtedly constitue only
races and the total number of species is, probably, only 11 or 12.
The subfamily of American raccoons, Procyoninae, includes
the genera: Bassariscus (“Jentinkia”**) (cacomistles; 2, possibly 3
species), Procyon (raccoons, 2 species), Nasua (coatis, 1 species),
Nasuella (mountain coatis, 1 species), Potos (kinkajou, or monkey
bears, 1 species), Bassaricyon (1, possibly 2, species).
The subfamily of Asiatic raccoons or pandas, Ailurinae, con-
sists of the genera—Ailurus (panda, or cat bear, 1 species) and
Ailuropoda (bamboo bear or giant panda, 1 species). Species of
the family constitute in all 5% of the total number of species of the
order. Equally with hyaena and bears, this family is one of the
least specious families of carnivores.
In practical terms, a series of species are innocuous, some may
be harmful to hunting economy, and individual species are consid-
ered valuable fur-bearing species or game animals (raccoons).
In the fauna of our country, indigenous representatives of the
family are absent. One representative of the raccoon genus, Procyon
has been acclimatized as a fur-bearing species (V.H.).
*Appears to contradict statement in previous sentence—Sci. Ed.
**Misspelled “Yentikia” in Russian original—Sci. Ed.
1373
Subfamily of American Raccoons
Subfamilia Procyoninae Gill, 1872
Genus of Raccoons
Genus Procyon Storr, 1780
1780. Procyon Storr. Prodromus meth. mamm., p. 37. Ursus lotor
Linnaeus.
Size moderate.
In general appearance and body structure, the animal resembles
the fox or, more closely, raccoon dog. Head broad, with sharp
muzzle, ears large, protruding from the fur and tapering, tail fluffy,
not prehensile, its length comprises about half, or slightly more, of
body length. Digits long, deeply separated, grasping.
Skull moderately elongated, with broad braincase and well-
defined crests. Hard palate extends far backwards—to 1/2 or 1/4
of total length of palate.
И 2
Dental formula I 3 С i р i M aa 40. Molars and last upper
premolar broad, more or less defined quadrate form, with flat
masticatory surface and low, sharp protuberances. Carnassial teeth
not developed.
Os penis present, to a greater or lesser degree curved.
Omnivorous predators, associated with forest regions, good
tree-climbers. Northern forms in north of range fall into winter sleep.
A characteristically American genus, distributed in North
America to the northern border of the range of the family (see
characteristics of family above, and description of species below),
and to the extreme, or almost extreme, southern limit of range of
the family in South America—to southwestern Brazil, Paraguay,
northwestern Argentina and southeastern Uruguay.
According to its systematic position within the limits of the
family, the genus is quite closely related to several others and is
not related to the number of those which show particularly sharp
deviation. Both morphologically and biologically it is considered a
generalized, apparently, a less specialized group of the family.
913
1374
Within the limits of the genus, quite а lot of species have been
described. Seven of them have been accepted at the present time
by several authors. However, it is quite obvious, that there are in
the genus 2 in all—the North and Central American P. lotor
Linnaeus, washer-raccoon, and the South and Central American P.
cancrivorus Cuvier, crab-eating raccoon. These sharply distin-
guished species are sometimes relegated to different subgenera
(Eupro-cyon—for the southern species).
One of the species (northern) belongs among the most impor-
tant fur-bearing animals of North America.
During the last decades, P. lotor was introduced into several
European countries and into the USSR for acclimatization as a
quality fur-bearing species (V.H.).
WASHER-RACCOON, OR AMERICAN RACCOON
Procyon (Procyon) lotor Linnaeus, 1758
1758. Ursus lotor. Linnaeus. Syst. Nat., ed. X, I, p. 48.
Pennsylvania’.
Diagnosis
Only species in the genus in the USSR.
Description
In general appearance—relatively short trunk, short legs,
generally stocky build and dimensions—and in part in several char-
acteristics of color, the raccoon, most of all resembles the rac-
coon dog, and, to a lesser extent, the red fox. Tail relatively short
and constitutes about half the body length. Head short and broad
with a short pointed muzzle and quite large eyes. Ears set wide
apart, large and protruding strongly from fur, with moderately
acute tips.
Winter fur long and fluffy, quite soft, with a thick underfur.
Tail covered with long erect hairs and appears thick and fluffy.
Upper surface of hands and feet clad in very short hairs—their
pelage differs greatly from that on legs. Callosities on lower sur-
face of paws bare.
Further synonyms—very numerous—represent nothing of interest to us and are
not provided.
914
1975
General color tone of dorsal winter fur dull gray оп the upper
side, with greater or lesser admixture of brown, sometimes reddish
or chestnut hairs (mainly on nape). Dark tips of hairs form slight
wavy pattern against general fur color. On lower surface, general
color tone slightly lighter; guard hairs here shorter and fewer and
underfur more obvious. General tone of underfur grayish-tawny.
Fore legs and thigh region have same color as lower body surface,
hind legs higher*; their heels are blackish. Hands and feet lighter,
almost white. On tail are 5 to 7 sharply defined black or brown
rings, and its tip is black. Color of parts between rings (they are
broader than rings) grayish or brownish. On lower surface of tail,
dark rings not so sharply defined, their color lighter than on dorsally,
and they are sometimes interrupted.
Top of head gray. On facial portion, black or nearly black
(brownish) “mask” sharply defined in form of large spot beginning
on cheeks and extending, becoming wider, across eye to muzzle.
Along upper surface of muzzle, it extends to lower part of nose
forming narrow projection, which also extends backwards to fore-
head. Sometimes, stripe extending along muzzle slightly separated
from eye spots by a slight brightening [see Fig. 346]. Facial mask
is demarcated by well-defined white fields bordering it above and
extending backwards under ears and to sides of neck. Lateral parts
of muzzle, lips, chin and lower surface of lower jaw white. On
throat, a transverse black or brown area, separated dark mask
pattern; narrow white stripe which extends backward from muzzle.
Ears clad with short grayish or brownish hairs and bear small black
spots of varying size and definition at base of posterior surface.
In addition to the raccoons of the described normal color,
black animals are met with (melanists).
There is one molt per year.
Facial region of skull short and quite wide. Braincase volumi-
nous, obviously swollen from behind, long and wide in
interzygomatic region with nearly parallel borders. Facial length of
skull less than cranial. Nasal bones short and quite broad. Line of
the upper skull profile convex—frontal region highest, from which
profile line gently descends both backward and forward.
Supraorbital processes very weakly defined, frontal surface
between them somewhat concave longitudinally. Mastoid processes
*Meaning unclear; hind legs longer?—Sci. Ed.
1376
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ИИ
913 Fig. 346. Washer-raccoon, or American raccoon, Procyon lotor L. (Sketch by
A.N. Komarov).
large; paroccipitals moderately developed. Hard palate extends
behind end of toothrow to approximately 1/3 of total palatine length.
Auditory bullae rounded in form, inflated. Sagittal crest weakly
defined; occipital—well [defined]. Canines relatively short, upper
almost straight, lower clearly curved. Fourth premolar and first
molar of upper jaw almost equal to each other (molar a little larger),
916
1377
second molar approximately twice less than first. First and second
lower molars almost equal in dimensions.
Body length of males is 435-545 mm, of females, 410-570
mm; tail length of males is 200—405 mm, of females, 192-340 mm;
length of hind foot (with the claw) of males is 96-138 mm, of
females, 83-129 mm.
Condylobasal length of male skull is 94.3-125.8 mm, of fe-
males, 89.4—115.9 mm; zygomatic width of males is 60.2-89.1
mm, of females, 58.3—81.2 mm.
Weight fluctuates very greatly, particularly in connection with
fat accumulation before winter hibernation—from 5.4 to 15.75 kg
and may even reach 22.2 kg? (V.H.).
Systematic Position
In comparison to the second species of the genus—P. cancrivorus,
the described form is sharply characterized. In some respects, it is,
apparently, more specialized (V.H.).
Geographic Distribution
Central and North America. Acclimatized in the USSR, Central
Europe and on several islands of the West Indies.
Geographic Range outside the Soviet Union
Natural range occupies North America and Central America, south-
wards to the Panama Canal zone. The northern border of the range
is quite complicated—starting at the lower reaches of the Saint
Lawrence River (near Quebec [city]), it extends along the south of
Quebec and Ontario (Canada) provinces to Lake Nipigon (a little
north of Lake Superior), starts along the shore of Superior, going
along it to its western extremity. Thence, it directs itself to the
southwestern extremity of Lake Winnipeg, and farther, rises in a
curved line northwards to the western part of Lake Athabaska.
Thence, the border sharply descends to the south-southwest—to
There is very little information on dimensions and weight of raccoons acclima-
tized in our [country], and they do not give a correct idea about variation in these
characters. Figures given above are from American sources and characterize variabil-
ity of the species in its native land in general. Size and maximum weight after Hall
and Kelson (1959), remaining data on weight after Burt and Grosenheider (1952).
1378
the southwestern corner of Alberta Province. Farther, the border
takes a completely curious form. It describes a large extension, a
loop as is were, to the south, bypassing a significant (eastern) part
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915 Fig. 347. Skull of washer-raccoon, or American raccoon, Procyon lotor L.
1379
of the Rocky Mountains and Great Basin and descends to 40° and
even 35° N. lat*. From this region, being drawn towards the Pa-
cific Ocean the border passes northward, sometimes near it (south-
ern), sometimes strongly distanced (northern), to the southeastern
corner of British Columbia and, rising somewhat to the northwest,
reaches the coast of the Pacific ocean at the level of the northern
extremity of Vancouver Island.
In the Pacific Ocean, the range includes Vancouver Island
and the small islands of Las-Tres-Marias along the Mexican coast.
In the Atlantic Ocean, it is encountered on small islands lying
directly off the coasts of South Carolina, Georgia and Florida.
916 Fig. 348. Natural range of Procyon lotor L. Somewhat simplified. Habitats in the
Archipelago of the Windward Islands are not shown (after Hall and Kelson, 1959).
*The species range now includes most of this region—Sci. Ed.
917
1380
Generally speaking, raccoons are absent on islands of the West
Indies; however, they are found on some of the Bahamas (in par-
ticular, New Providence near Nassau‘), on Cosumel Island near
Yucatan, and on the Lesser Antilles Islands, in particular, on
Guadeloupe and Barbados (possibly also others) from the Wind-
ward Islands.
The raccoon was acclimatized and lives in many parts of the
F[ederal] R[epublic] of G[erman] and the С[егтап] D[emocratic]
R[epublic]. It was not introduced in other countries. Transgres-
sions to Holland and northwestern France are known (Niethammer,
1963).
Geographic Range in the Soviet Union
The experiment in acclimatization of the raccoon in our country
began in 1936, and was repeated more than once in various parts
of the country. The total number of attempts (up to 1962) was 26
releases, for a total 1222 individuals. Of these, 64 were from the
farms and zoos (38 imported from western Europe), the remaining
were derived from a population which had been put together in the
Trans-Caucasus (Zakatalo-Nukhinsk valley in Azerbaijan;
Rukovskii, 1963). The animals were introduced in various places,
often far from each other, and they never formed a significant,
continuous range. Far from all of the transplantations (with respect
to the purely biological side of the matter) had the same results. In
several places, the animals died, in others, the results have not yet
been sufficiently determined or are unreliable, in several, the ani-
mals apparently survived well, obviously settled down and formed
quite numerous populations?.
In the Far East, raccoons (black) were introduced on Petrov
Island near Vladivostok and a series of places in Primor’e territory
(southern half), but all introductions were unsuccessful—the ani-
mals died (on the map, these places are not shown; Abramov,
“The suggestion that raccoons appeared (were introduced?) on the Bahama
Islands in 1932 (Niethammer, 1963) was based on a misunderstanding—P. maynardi
was described as early as 1898 from New Providence Island.
‘Concerning information given below on places and results of transplantation of
raccoon in our country are based on the most recent, in part unpublished, data and
describe the situation in 1962 and the beginning of 1963. They differ significantly
from Kolosov’s (1958) and Rukovskii’s (1960) communications, in which the success
of acclimatization was strongly overstated and already did not conform with the actual
situation.
1381
917
918
Fig. 349. Localities of introduction and occurrence of the washer-raccoon or American
raccoon, Procyon lotor L., in the USSR. Places where acclimatization was unsuccessful
or results unclear, are designated by crosses; places where the raccoon apparently
survived—by dots, and the districts in Byelorussia and in Azerbaijan, where more or less
stable populations were constituted, are designated by blotches (V.G. Heptner).
1963). In Middle Asia, raccoons were introduced into former
Dzhelalabad district of Kirghizia, in the nut and apple forests of
the southern slopes of the Tien Shan Mountain Range—facing
towards the Fergana valley—in Arslanbob (north of Dzhelalabad).
They were noted here in the Ach’region (between Arslanbob and
Dzhelalabad) and in the Bazar-Kurgan region (west-northwest of
Dzhelalabad; introduced in 1936 and 1952; Novikov, 1956). In
January 1963 here “raccoons are practically absent’®. Raccoons
were introduced into the mountain forests of the valley of the
Pskem River (near kishlak* Sidzhak northeast of Tashkent, be-
tween the Ugamsk and Pskem ranges—the extreme western spurs
of Tien Shan; Salikhbaev et al., 1963).
°P. Sagaraev, Board of Forestry and Nature Protection belonging to the Council
of Ministers of the Kirghiz Soviet Socialist Republic. Apparently, these introductions
(in Chavachin and Karaamin forest farms; their location is imprecise), and Yanushevich
(1963) also spoke of them, believing that these introductions were not successful,
recommending cessation of raccoon introduction in Kirghizia.
* Local word for village—Sci. Ed.
1382
According to one datum of Salikhbaev et al. (1963), the rac-
coon survived here, and has somewhat broadened the area of its
occurrence; according to other data—“the results...were unclear”,
and evaluation of its situation and its future fate is too optimistic
in the literature’. On the whole, attempts to acclimatize the
raccoon in Middle Asia have been unsuccessful or their results as
of 1963 are not clear and, apparently, they must be stopped.
In the eastern Trans-Caucasus (Azerbaijan), introduction of
the raccoon into the forest zone associated with the southern slope
of the Main Caucasus range, beginning in 1937, was successful
and led to the creation of a quite considerable and stable popula-
tion (annual catch of 1000-1500 individuals), occupying one quite
significant section, and isolated centers separated from it. The main
range extends as a strip from the region of Belokana City (a little
east of Lagodekha) along the so-called Zakatalo-Nukhinsk forest
(Zakatalo-Nukhinsk valley) to Nukha city, and farther, to the
Ismailla region, i.e. for an extent of more than 200 km. This region
is the largest focus of occurrence of the species in the USSR, from
whence is taken material for future settlements.
A separate section of occurrence is found in the so-called Kuba-
Khachmas forest massif—from Kuba toward the northeast and north
to Khachmas and Khudat on the railway line near the sea shore. At
the beginning of the 60’s, this section was apparently not yet con-
nected with Zakatalo-Nukhinsk. A small section of occurrence lies
in the Talysh and Lenkoran forests. On the whole, in 1963 in
Azerbaijan, the raccoon inhabited Belokansk, Zakatal’sk, Kakhsk,
Nukhinsk, Vartashensk, Kurtkashensk, Ismaillinsk, Khachmassk and
Lenkoransk regions (Ya.K. Aga-Zade, Nature Protection Board of
the Council of Ministers of the Azerbaijan SSR).
In the North Caucasus, the raccoon was introduced, and appar-
ently, survived, in the riparian forest of the Terek and along the
Sulak [river]? in the lowlands of Dagestan (south of the eastern
Cis-Caucasian lowland). Raccoons were introduced along the “left
tributary of the Kuban’” (Aliev, 1963), but clear data about them
are absent. The attempt to settle raccoons in Kabardin-Balkariya
(black raccoons) was unsuccessful.
As shown by an almost ten-year experiment, attempts to accli-
matize the raccoon in Byelorussia were, apparently, quite success-
7M. Isamukhamedov, Board of Forestry and Nature Protection of the Council of
Ministers of the Uzbek. SSR.
919
1383
ful, where, as a result of three introductions (52, 37 and 38
individuals in 1954 and 1958), a significant population was formed
(on 1 January 1963, about 700 individuals). It occupies a section
along the Pripyat’ River from Petrikov up almost to Pinsk (about
120 km), approximately 30 km in breadth along each bank
(Petrikovsk and Zhitkovichsk regions of Gomel’sk district, and
Stolinsk region of Brest district*). The situation of the population
has been evaluated as satisfactory.
Information in the foreign literature (Niethammer, 1963) con-
cerning the occurrence of raccoons in Latvia (in spring 1961, 7900
individuals!), is completely untrue. In Latvia, the raccoon-dog is
widely distributed, but the raccoon was not introduced at all.
The perspective on raccoon acclimatization in our country, if
the Caucasus—mainly the Trans-Caucasus—is not considered,
Trans-Caucasus, is very limited from the purely biological side,
although steps already undertaken have not yet been sufficiently
studied. The economic side has also not yet been analyzed (for
details, see section “Biology”) (V.H.).
Geographic Variation
American authors, even the most recent, believe that geographic
variation of the raccoon is very extensive. Some accept 30 races of
the species P. lotor, and another 4 species (insular) besides P.
cancrivorus, accepted here (Miller and Kellog, 1955); others rec-
ognize 24 races of P. lotor and another 5 species (one with two
races), besides P. cancrivorus (Hall and Kelson, 1959). These con-
siderations are, apparently, exaggerated.
It is not known which forms were introduced into our country.
As said, part were derived from the zoological parks and fur farms
(V.H.).
Biology
Population. Experiments in acclimatization of the American
raccoon first began in 1936, when the Uzbek Zoo introduced 22
raccoons in former Achin region of the Kirghiz SSR. In 1949, their
number here did not exceed 80-100 individuals, with a density
5—8 animals рег 1000 hectares.
8A. Abaturin, State Committee for the Nature Protection of the Council of Min-
isters of Byelorussian SSR.
1384
An experiment in introduction of two pairs of black raccoons
on Petrov Island in Primor’e territory in 1937 proved unsuccessful.
In 1941, 21 raccoons were introduced in Ismaillinsk region,
Azerbaijan SSR (Rukovskii, 1951). In 1950, it was established that
they already inhabited a series of regions in an area of 1300-1400
km2. The total number of raccoons reached 2000 individuals, with
density averaging 15.3 animals per 1000 hectares. In 1950, 44
raccoons were introduced into the Khuzakhsk and Khasav’ yurtsk
regions of Dagestan. In 1951, 28 raccoons were introduced 70-90
km to the south of Krasnodar city. In 1952, 33 raccoons were
introduced into the former Oktyabp’sk region of Kirghizia, and 79
raccoons into the Zakatal’sk region of Azerbaijan SSR. In 1953, in
former Khizinsk region of Azerbaijan, 17 raccoons were intro-
duced, while 43 raccoons were settled in former Bastandaksk
region of Kazakhstan, and 16 black raccoons—in Kabardinsk
A[utonomous] S[oviet] S[ocialist] R[epublic] south of Nal’chik in
the region of the Nal’chik and Belaya rivers. In 1954, 50 raccoons
were introduced in Petrikovsk region of Gomel’sk district of
B[yelorussia] SSR, 55 raccoons in former Budennovsk region of
Primor’e territory, and 104 raccoons—on Zelenchuk river in
Stavropol’ territory. In all, from 1936-1954, 534 raccoons were
introduced for acclimatization. From 1941, all introduced animals,
except 16 black raccoons, were caught in the Zakatalo-Nukhin
valley of Azerbaijan (N.N Rukovskii).
In 1954, 486 raccoons (Abramov, 1961) caught in the Trans-
Caucasus were introduced in Partizansk, Anuchinsk and Yakovlevsk
regions of Primor’e territory. The introductions were accomplished
in coniferous—broad-leaved forests in the valleys of montane taiga
rivers. In these forests there was extensive tree-felling and forest
fires, trees with hollows were few, and food resources were insuf-
ficient. The raccoons went off to the lowlands, but there, tree
hollows were also few, and the raccoons settled in burrows, clefts
of rocks and haystacks. They were not able to accumulate fat before
winter and they spent the winter badly (its duration is 120-150
days) with severe frosts. Mortality was recorded among them from
hunger and cold. In following years, the population everywhere
was less than in the year of introduction. Reproduction occurred
locally, but mortality was greater. In the 60’s, the raccoon became
extinct in Primor’e.
920
1385
In Krasnodarsk territory (Kotov and Ryabov, 1963), raccoons
settled in a narrow belt with an area of about 7000 km’, extending
from Novorossiisk to Lazarevskaya. Here, they were sporadically
encountered everywhere in low density. In better places—in
Goryache-Klyuchinsk region—3.5 animals per 1000 hectares; in
the Gelendzhiksk region—0.6; in Lazarevsk region—0.1. In more
favorable lands of the Ismaillinsk region, density reached more
than 20.0 animals per 1000 hectares (Rukovskii, 1951). In their
native land, in boggy lands of Wisconsin state (USA), density of
raccoon settlement fluctuated from 25.7 to 34.6 per 1000 hectares
(Dorney, 1954). Under more favorable conditions of the southern
states, raccoon populations may attain a very high density. Up to
1949, their highest density was considered to be established in
Illinois; in an area of 384 hectares—111.0 animals per 1000 hec-
tares, but in the state of Missouri, 102 raccoons were caught in an
area of 25 hectares; i.e. here their density was 408.0 (Twichell and
Dill, 1949). In the state of Ohio (Butterfield, 1944), in open hunt-
ing lands, were established densities of: 36.1; 142.0; 166.6; for
reserves—340.0 (66 raccoons on 193.5 hectares).
Habitat. In Kirghizia, raccoons inhabit montane forests domi-
nated by walnut trees. In the Trans-Caucasus, they prefer valleys
with broad-leaved forests consisting of oak, white-leafed poplar,
smooth-leaved elm, alder, walnut and chestnut. In these forests,
there are many lianas, wild almond-leafed willows, hops and hol-
low trees. Raccoons no less willingly settle in montane beech for-
ests almost devoid of undergrowth (N.N. Rukovskii).
In the USA (Nelson, 1930), raccoon—one of the few wild
animals, whose populations increases with cultivated appropriation
of territory. With free water and forest vegetation, raccoons
here settle also in the swamp lands (Dorney, 1954); the presence
of trees of large diameter with hollows are preferable but not
necessary.
Food. The American raccoon belongs to the group of omnivo-
rous predators. In composition its food includes a variety of both
animals as well as plants. Their combination, and the ratio of each,
depend upon the biotope, time of year, and the yield of each food
(see Table 71).
In the northern USA, in swamp lands of the state of Wisconsin
(Dorney, 1954), the raccoon’s main food consists of crustaceans,
fish, young muskrats, insects, grapes, fruits and berries (see Table
12).
921
1386
Table 71. Seasonal characteristics of raccoon nutrition in Texas (% of
occurrence. Becker, 1945)
Type of food Spring Summer Autumn Winter
Birds 4 — — —
Snakes — 2 — —
Fish 4 4 — 4
Freshwater crustaceans — — — —
Mollusks 4 3 — —
Wasps — — 23 —
Beetles 18 22 2 —
Crickets 3 16 10 —
Larvae and pupae of butterflies — _ — 1
Spiders — — — 7
Water scavengers, and diving — 9 — —
beetles
Water bugs — 16 —- —
Acorns 55 47 29 ИО
Wild grape — 38 24 —
Mulberry 14 2 6 —
Persimmon — 10 36 10
Hawthorn — 3 6 13
Table 72. Seasonal characteristics of raccoon nutrition in Wisconsin (%
occurrence)
Food type I-IV V-VI VI-VII IX-X X-I
Adult muskrats 20 6 2. 2 12
Young muskrats — 40 35 9 —
Cricetid rodents 10 11 2; 1 —
Adult ducks —- 1 — 1 12
Duck eggs — 7 3 1
Eggs of other birds — 4 1 — —
Fresh water 49 52 64 64 6
crustaceans
Fish 20 17 3 22 25
Frogs — 1 — — —
Insects 7 27 16 5 —
Maize 39 8 1 11 50
Acorns 2 — a 9 —
Grapes — — 8 60 —
Wild pear — — 10 14 —
Raspberry — — 19 — —
The frequency of young muskrats in the raccoon diet in vari-
ous swamplands fluctuates from 8-9 to 19% (Dorney, 1954). In
the state of Washington (Tyson, 1950), the occurrence of mollusks
in the diet of adult raccoons in summer was 85%, of crustaceans—
85%, of fish—65% and of neris worms*—50%. In the stomachs of
*Meaning not clear; probably refers to marine polychaetes—Sci. Ed.
922
1387
young raccoons, mollusks constituted 67%, crustaceans—100%,
fish—45%, worms—11%, milk—78% of the contents (20 stom-
achs of adults and 9 of young).
In the Trans-Caucasus (612 excrement samples; Rukovskii,
1951, 1957); the raccoon feeds mainly on the following kinds of
food (% of occurrence):
Mammals 5.5 Fish 2.3
Birds В? Crabs 2.6
Amphibia 52.6 Insects 82.4
Reptilia 2 Mollusks 12.9
Plants 63.9
Therefore, under new conditions, the raccoon shows itself to
be a typical polyphagous animal, like the badger as regards food
habits. Of mammals, it feeds on mice and voles; of birds—mainly
on small passerines and injured ducks. Of reptiles, raccoons eat
turtles, lizards, rarely snakes; of amphibians—Trans-Caucasian frog.
Among insects were encountered beetles, orthopterans,
lepidopterans, dragon flies and caddis flies, among them a signifi-
cant number fall among pests of agricultural plants—mole cricket,
locusts, crickets, long-horned beetles, bark beetles, click beetles,
leaf beetles, weevils and others. Raccoons feed also on plant foods:
in the Trans-Caucasus, most frequently wild grapes, fruits of lianas
(sasparilla), nuts and acorns. To a lesser extent, they eat pears,
apples, cherry plum and blackthorn. In spring and winter, raccoons
utilize more animal food, in summer and autumn—plants.
Home range. Dimensions of the home range of the adult rac-
coon are associated with the abundance of food within its borders
and the distance of feeding areas from shelters. In Azerbaijan
(Rukovskii, 1951), the area of such a home range fluctuates from
4 to 9 km’, with a radius of 1.0-1.5 km from shelter. A lactating
female does not travel more than 600 m from her litter. Conse-
quently, the area of her home range does not exceed 144 hectares.
In the state of Ohio (USA), during two years, no cases were
recorded where a raccoon traveled more than 1.5 km from the
place it was marked (Butterfield, 1944; according to recapture of
91 marked raccoons). On average, the animals were caught no
farther than 400 meters.
Burrows and shelters. In Azerbaijan, the raccoon settles in
hollows of white-leaved poplar trees, rarely oak (Rukovskii, 1951,
1388
1955). This is explained by the faci that hollows of size suitable
for raccoon, are here more frequent in poplars. Average height of
hollow trees inhabited by raccoons is about 21.0 m, their circum-
ference (at breast height) reaches 4.5 m. The living cavity is situ-
ated at a height of 10-12 т from the ground. The average diameter
of the entrance hole is 19 cm (not less than 12 cm). Amplitude of
daily temperature fluctuation inside such a cavity does not exceed
3.5°C, and in hot summer hours, the temperature is 10-12°С lower
than that of the outside air. Besides the main living hollow, within
the home range of the raccoon, there are also temporary shelters—
hollows, where the raccoon spends the day resting.
In North America, raccoon also settles in caves and among
rocks and clefts. In the absence of tree holes, it settles in earthen
burrows, muskrat huts, boxes for nesting ducks and other shelters
(Dorney, 1954). During the capture of 297 raccoons in the state of
Ohio, only 8% of them were hidden in tree holes, 47.5%—among
rocks and in burrows, the remainder—under roots of trees, on newly
fallen trees, etc. Of 316 raccoons, 46.4% occupied ground bur-
rows, 18.2%—tree hollows, 14.4% —shelters in the base of trees,
and 10.0%—among rocks. Raccoons occupied 22% of the artificial
bird nests of the box type (Butterfield, 1944).
Raccoons often occupy burrows of marmots, sometimes in open
places. Litters were also found in haystacks and heaps of brush-
wood. It is supposed that the raccoon was initially a species closely
associated with tree hollows. As a result of cutting down large-
trunked trees containing hollows, and the transformation of forest
areas into agricultural ones, the raccoon became an animal capable
of living under various conditions of cultivated landscapes.
Daily activity and behavior. The raccoon is a crepuscular-noc-
turnal animal. It emerges from its hole at sunset and sometimes
returns to it even at day break. It climbs trees well, but it gathers
its food mainly on land—along river banks, in swamps and other
similar places. In searching for food, it sometimes rambles through
shallow water and along sand bars.
A characteristic feature of the raccoon is that it rinses its
gathered food in water before eating it. Hence, the raccoon’s name
“poloskun”, (= to wash off).
Hibernation and winter sleep. In districts with snowy, cold
winters, the raccoon, as well as the brown bear and badger, have
a characteristic winter sleep. In the Trans-Caucasus (Rukovskii,
923
1389
1951), in association with warm winters and brief frost, raccoon
activity stops only during snowfall. It emerges when the soil is
partly free of snow. In Kirghizia and Canada, where snow lasts for
a longer time, raccoons leave their shelters earlier, before the snow
thaws in spring. In Canada, their sleep lasts for about four months
of the year. Here they are able to survive hollows while the tem-
perature of the outside air is down to —43°C.
Seasonal migrations and transgressions. After the introduc-
tion of 250 raccoons from Ohio into Arkansas, their appearance
was noted 37, 75 and 255 km from the place of introduction
(Butterfield, 1944).
Reproduction. In Canada, i.e. in the northern parts of the range
in America, estrus occurs is March. In yearling females, estrus
takes place in May-June. In the state of New York, raccoons mate
at the end of January—beginning of February (Mackeever, 1958).
In the state of Michigan, the majority of females mate in February—
beginning of March, and few of them—at the end of March; indi-
vidual cases of mating were observed in Virginia at the beginning
of June.
In the states of Florida and Georgia, the season of reproduc-
tion lasts from 10 February until 3 August: in February, 14% of the
females are mated, in March—41%, in May—12%, in June—12%,
in July—12% and in August 6% (Llewellyn, 1953*). After а preg-
nancy lasting for 63 days, raccoons bring their young into the light
in the period from April to the beginning of October. Practically
half of all [parturition] occurs in May. The number of embryos
averages 3.2, fluctuating from 2 to 5. In the state of Michigan, the
average number equals 4, with fluctuation from 3 to 7.
In the Trans-Caucasus, young appear in April—beginning of
May. The quantity of newborns fluctuates from 3 to 8, more
frequently 4 (Rukovskii, 1951).
Growth, development and molt. On the 21st day of pregnancy,
length of the embryo reaches 24 mm, on the 35th day—45 mm,
and on the 46th day—65 mm (Llewellyn, 1953)**. Young are born
blind, and eyes open on the 20th day. At 1.5 months of age, they
cut all teeth. At the age of 2.0-2.5 months, they already accom-
pany their mother while searching for food, and at 4—5 months old,
they switch over independent foraging. In September—October,
*Not in Lit. Cit—Sci. Ed.
**Not in Lit. Cit.—Sci. Ed.
1390
young raccoons do not yield, in terms of dimensions, to adults,
but are markedly lagging behind them in weight.
Raccoons attain sexual maturity at the age of about one year.
Sex ratios in populations are different: in the states of Ohio and
Illinois, the ratio favored females—46:54 and 40:60 (among hunted
animals, even 14:86), in the state of Missouri—(62:38 and 61:39,
i.e. the ratio favored males).
Out of 102 raccoons caught in an area of 25 hectares, 32 were
alone, and in 12 cases caught in pairs; in 6 cases the raccoons were
obtained in trios. Only one time were groups of 4, 5 and 6 rac-
coons were caught. The animals were caught in winter during winter
sleep (Twichell and Dill, 1949).
Enemies, diseases, parasites, mortality, competitors and popu-
lation dynamics. Wolf, lynx and eagle-owl are regarded as enemies
of the raccoon. In places inhabited by raccoons in the USSR, these
predators are rare and do not cause great losses to it. There are no
data on diseases and mortality of raccoons. In Trans-Caucasus,
competitors of the raccoon in obtaining food are shown to be brown
bear, badger, otter, pine and stone martens, jackal, cats, wild pig,
red deer and fat dormouse (Rukovskii, 1951, 1953).
There are no data on population dynamics. In Kirghizia, the
raccoon population increased 5-fold within 13 years, and in
Azerbaijan—100-fold within 9 years.
Field characteristics. The crepuscular-nocturnal activity of the
raccoon makes it very difficult to carry on direct observations.
Their presence may be known by the characteristics of inhabited
hollow (claws marks on bark, polishing of entrance into tree-hole,
etc.) and by the tracks of their paws on damp soil along the banks
of water bodies. In the track of the anterior paws, prints of five
long clawed digits are clearly seen. Tracks of the hind paws resem-
ble, in strongly reduced aspect, tracks of the hind paws of brown
bear. They are more extended in length than in the badger
(P.Yu.).
Practical Significance
The American raccoon is a valuable fur-bearing animal. Its hide is
mainly used in the manufacturing of men’s winter fur coats. In
view of the fact that the districts of its probable distribution should,
apparently, be restricted to the zones of mixed and broad-leaved
924
1391
forests, the practical significance of raccoon as а fur-bearing ani-
mal cannot be great. Its practical importance may be more con-
siderable in the southern regions of the USSR, where objects for
fur manufacture are few. With occupation by raccoons of marshy
lands, they may cause losses to young muskrats. Raccoons do not
pose any danger to adult muskrats and aquatic birds since they
only feed on injured individuals. Raccoons are useful as they de-
stroy insects harmful to forests and agriculture. (P.Yu.).
Family of Mustelids
Family Mustelidae Swainson, 1835
AMERICAN MINK'!
Mustela (Mustela) vison Schreber, 1777
1777.2 Mustela vison Schreber*. Die Sdugethiere .... pl. 27.
Eastern Canada.
1949. Mustela vison tatarica Popov V.A. Tr. Kazansk. Filiala
AN SSSR. Seriya biol. i s-kh. nauk, 2, p. 135. Tatari.
1958. _Lutreola vison altaica. Ternovski. Biologiya i akklimatiz-
atsiya amerikanskoi norki (Lutreola vison Brisson) na
Altae (Biology and acclimatization of the American mink
... in Altai), p. 15. Middle course of Sary-Koksha River,
Altai. (V.H.).
Diagnosis
Color and skull are same as in Russian mink, Mustela (Mustela)
lutreola, but upper lip has same color as top of head (not white)
and postorbital constriction of skull is sharper—its width is less
than width of interorbital area. Dimensions somewhat larger (V.H.).
‘In our literature, this species is sometimes called the “eastern mink”. There is
no basis for the introduction of this new, completely artificial name. This name also
is confusing—the range of the American mink, with respect to the European, lies
nearer to the west, in the western hemisphere.
2Some of our monographers (Novikov, 1939, 1956; У. Popov, 1949; Ternovskii,
1958) refer to Brisson as the author of this name with reference to 1756, and they
even try to designate the priority of this assignment to Schreber. This is a clear
misunderstanding. Zoological nomenclature begins in 1758 (publication of the 10th
Edition of “Systema Naturae” by Linnaeus) and all names given earlier are generally
not accepted.
*Mispelled Schreder in Russian original—Sci. Ed.
925
1392
Description’
In general appearance and construction, the American mink is
completely like the Russian; however, the tail is on average
somewhat longer, and reaches or almost reaches half the body
length (from 38 to 46 and even 51%).
Winter fur is as in the Russian mink, but denser, close-fitting,
longer, more luxuriant and softer. As fur, its value is higher than
fur of the European mink. Length of contour hairs (Altai) on back
is 27.7 mm, on belly is 22.2 mm, guard hairs are 22.7 and 16.6
mm, underfur is 14.8 and 10.6, respectively. Number of hairs on
1 сп? on sacrum is 19,530; on shoulder, 21,783 and on the belly
17,320 (Tserevitinov, 1958).
According to other data, in Altai mink (4 specimens)
(Ternovskii, 1958), number of hairs on 1 cm? in middle of back
ranges between 22,664 and 37,100, consisting on average about
22,458. For one guard hair, there are from 31 to 34 underfur hairs.
Length of guard hairs on back is 20.9 mm, of underfur hairs, 12.3
mm.
There is one pair of inguinal teats, and three pairs of abdomi-
nal, in all eight; as an exception, six.
General tone of winter fur tawny of various intensity, from
very dark blackish-tawny to light-tawny. Color is equally distrib-
uted over all of body, lower side only slightly lighter than upper.
Guard hairs, bright, dark-tawny, on spine often approaching
black, straight and elastic, but not coarse. Underfur hairs on dorsal
part very wavy, grayish-tawny with a bluish tint. Base of underfur
is light bluish-gray. Tail darker than trunk, and closer to tip, inten-
sity of dark tone strengthens, and sometimes end of tail has pure
black color. Sometimes, along the back, a broad wash of somewhat
darker stripe is displayed. Chin and lower lip white—white color
does not spread onto upper lip as a rule.
Color exhibits quite significant individual variability, which
may be somewhat different in populations comprising various parts
of our country. This applies to both qualitative and quantitative
aspects (for cause, see section “Geographic Variation”). Percent-
age ratio of different saturated color types varies in different
ЗАП data given below, especially where no individuals are preserved, belong to
populations from natural conditions within the USSR. Best known are populations in
the Tatar Republic (V. Popov, 1949); Altai (Ternovskii, 1958); in others data are very
scarce and fragmentary.
1393
populations, but dark and very dark animals predominate. Thus, in
Tatariya, tawny-black animals constitute 48%, dark-tawny 36%,
tawny 10% and the light-tawny 6%. Frequently on skins (in Tatariya,
about 24%) white guard hairs (“hoary”) are encountered and some-
times the tail tip is white (in Tatariya, about 4%).
In some individuals, underfur is white in color and may
occupy from 40 to 80% of skin area, or they may even be distrib-
uted on the entire body surface. In addition to white lower lip and
chin, very often there are irregular white marks noted on the lower
surface of the body—on throat, neck, chest, belly and groin. They
lie along the mid-line of the animal venter in the form of patches
and sometimes stripes. The number of patches and the area they
occupy vary. In the Altai, 29.2% of them were on the lower lip and
throat, in 30.4% they extended from lower lip to fore limbs, in
35.1% they extended from lower lip to anus, in 4% they were
distributed on the groin, in 0.1% between fore paws, in 1.2% on
lower lip, throat and groin, and in 0.1% on chest and groin. It is
noteworthy that the white marks may sometimes (3.3%) not occur
925 Fig. 350. American mink, Mustela (Mustela) vison Schreb. Sketch by А.М. Komarov.
926
1394
at all (data from 727 skins). As а rule, the upper Пр in the Ameri-
can mink is never white. However, in the Altai, in 36.4% (213
skins of 585) white patches occurred on the upper lip. Their di-
mension varies from weakly defined to those occupying more than
half the lip. In Tatariya, this is absent. In the Altai, the white
marks are, apparently, more strongly developed than in other re-
gions of our species range. A broad white patch (as in marten) on
the neck and chest, is rarely encountered in the Russian mink M.
(M.) lutreola; in our American mink, it has not been mentioned.
Considerable development of white patches is characteristic of
cage minks. It is very interesting that in Tatariya, with naturaliza-
tion of the animals, within 10-11 years, the number of skins with
white patches on chest, belly and groin, decreased sharply. At the
end of this period (1938-1948), only animals with small white
patches and only on lower lip, chin and throat were found (V.
Popov, 1949). Apparently, there was a gradual elimination of this
feature.
The animals in which the ends of the guard hairs are twisted,
similar to singed hairs, are rarely met with. This “singing” is a
mutational deviation known from caged animals in the Altai (7.8%).
On the whole, color variation of American minks inhabiting the
USSR is much greater than in Russian [mink].
Sexual differences in color and in development of patches and
hoariness are absent. The latter is more often developed, either in
males (Tatariya) or in females (Altai, 94.6%). White underfur
apparently occurs more often in females. Hair length in females is
approximately 2 mm less than in males (middle of back, V.A.
Popov, 1949).
Summer fur of our [American] mink has not been described in
detail. Its color differs little from winter, but is shorter, sparser and
dull. Length of summer guard hairs on dorsum is 17.3 mm, and
that of underfur—9.8 mm; on 1 cm?’ of dorsum there are about
17,450 hairs on average, for 16-22 underfur hairs for each one
guard hair (Altai).
The skull is similar to that of the European mink, but it is more
massive. It is narrower and less elongated, cranium is wider, shorter
and more flattened. Postorbital constriction is more sharply de-
fined—in narrowest part, its width is greater than width of skull
between orbits. Zygomatic arches, especially in middle portion, are
thinner and weaker than in Russian mink, but diverge somewhat
1595
more widely. Mastoid processes well-developed and fuse with lower
part of occipital crest. Auditory bullae are flat and broad; in
posterior part they are set apart from each other not so far as in
European species. Ratio of their length to their breadth approxi-
mately 1.0:1.5. Protuberances, crests, etc. of skull are generally
more sharply-defined than in Russian mink; occipital crest devel-
oped significantly more strongly; sagittal crest—well[-developed].
Upper molar larger and more massive than in Russian mink, its
area, in frontal projection, no smaller or only slightly smaller than
area of upper carnassial tooth; longitudinal diameter of its well
developed inner lobe more than logitudinal diameter of outer lobe.
Posterior end of second upper premolar more or less strongly trans-
gresses into notch found at anterior end of carnassial tooth. Inner
surface of main cusp of lower carnassial tooth with well-developed
sharp rib. On the whole, in its general appearance, specialization
of skull as that of a predator skull more strongly developed than
in Russian mink, and as regards degree of specialization itself,
somewhat reminiscent of polecat skull.
Skulls of females, besides general dimensions, distinguished
by somewhat less development of crests, protuberances, etc. and
are somewhat lighter. Skulls of young are characterized by the
more rounded and expanded braincase and weakly developed
facial portion.
Geographic variation of the skull within the borders of our
country not expressed (see beyond).
The os penis of the American mink, having a structure typical
of the genus, differs considerably from that characteristic of the
European mink, not only in size, but also in form. If one looks at
the bone from below, its terminal part is not deflected to the right
and the bone is curved in one plane. Its base is laterally com-
pressed and is significantly wider along the vertical. Basal third is
also flattened. Along middle of sides, there is a short groove, and
on lower surface of the anterior part, there is a deep groove
(Novikov, 1956; V. Popov, 1949). There are 14 thoracic vertebrae,
6 lumbar, 3 sacral and 18-21 caudal (40 individuals). Weight of
heart in males (16) is 5.3 M8.4—-11.5 g, in females (9), 4.5 M5.4—
6.18 g; cardiac index is 7.25-М9.71-12.79. Length of intestine in
males is 1635-М1806-2140 mm, in females, 1220 М1480-1690
mm (Altai). Number of caudal vertebrae in Tatariyan mink is
19-22.
1396
Maye are ©
SANA ee
WS
Naz an St ee (5)
И ь. ГА а 4
Wh, NS
Sez W
sau A \
>
eS SN
A WW:
АТ, 1 24
Pe Е oe
а
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927
Fig. 351. Skull of American mink, Mustela (Mustela) vison Schreb.
928
1397
Diploid number of chromosomes is 30.
Body length of males is 340-450 mm, of females, 310-375
mm; tail length of males is 156-247 mm, of females, 148—215 mm;
length of hind foot in males is 51-70 mm, in females, 40-58 mm;
length of ear in males is 20-29 mm, in females, 19—26 mm (in 110
males and females from Tatariya and 41 males and 31 females
from Altai). :
Weight of males in winter (Altai and Tatariya) is 500-1580 с,
of females, 400—780 g.
Condylobasal length of male skull (32; Altai*) is 62.1-M66.4—
72.6 mm, of females (14), 57.0-M59.3-61.6 mm; zygomatic width
of males is 35.0-М37.1-41.4 mm, of females, 31.2-M32.8—34.4
mm; mastoid width of males is 30.3-МЗ32.8-36.8 mm, of females,
27.6-М28.5-30.5 mm.
Os penis length (24) is 35.9-M40.5—45.4 mm. Weight is 0.110-
МО.229-0.580 g. Age differences in dimensions and particularly
weight of os penis are very significant. Length.of bone in young
(30; Tatariya) is 37.9-М41.7-45.2 mm, weight is 0.130—M0.167-—
0.210 g; in adults (30; Tatariya), length is 39.9-M44.6-49.7 mm,
weight is 0.240-МО.336-0.450 р (V.A. Popov, 1943). Difference
in weight is double.
Sexual dimorphism in body measurements and weight is very
significant. Measurements of females (Tatariya) constitute 82-85%
on average of measurements of males, while weight is 60-80%.
Weight is very strongly variable according to season, reaching its
maximum in autumn. At that time, males in Tatariya weigh about
1000 g, maximum 1580 g, females about 600 g, maximum 780 g.
Least weight of males happens from April to June, females from
June to August (nursing young). Average weight of the animals,
and in part measurements, obviously change by year (V.H.).
Systematic Position
According to morphological characteristics, the American mink is
closest to the European. These concern its main ecological and
ethological characteristics. Some authors (among the most recent
examples, K. Zimmermann, 1959; Gaffrey, 1961) even consider
it as only a subspecies of the European, and suggest (Gaffrey,
1961), that in Middle Europe, remaining natural populations of the
“For skull dimensions of Tatariyan mink, see section “Geographic Variation”
929
1398
European mink disappears (extirpated) due to its hybridization with
the American, dissolving it so to speak, into the population of the
latter.
According to our data, however, uniting the two minks into
one species, cannot be accepted. They are close (“paired”), but
independent species. This is indicated not only by their morpho-
logical, and most of all, craniological, characteristics (color—white
marks on lips, a feature which is not very stable), but also by the
absence of hybridization in nature here, in part a result of
asynchrony of the reproductive cycles. It is possible that there is
also a lack of compatibility in genital systems also occurs (see
structure and size of os penis). Sharp antagonism exists in nature
between these species—not only competition, in which the larger
and stronger M. (M.) vison crowds out the European mink, usually
simply destroying it (V. Popov, 1949; V. Popov et al., 1954).
Accurate material data on hybridization of the two mink spe-
cies in nature are absent, apparently not only in our country, where
this question was especially studied, but also in Central Europe.
Their experimental crossing in captivity was also unsuccessful
(Schmidt, 1933; V. Popov, 1949). At the same time, the Russian
mink yields hybrids with the black polecat [M. putorius] (see above
and below).
The American mink itself represents a sort of further develop-
ment of the European mink—a further stage in the specialization
towards predatoriness. This is indicated by the structure of its skull,
with constricted interzygomatic area of braincase, more strongly
developed protuberances and crests of skull, stronger dentition,
etc. In comparison with the skull of the American mink, the skull
of the European bears several infantile features.
The “paired” mink species show a quite remarkable parallel-
ism with the “paired” species of polecats—black M. (P.) putorius
and white M. (P.) eversmanni. The European mink corresponds to
the first, and the American—to the second. This parallelism is
manifested in a series of features—first of all, in skull structure
and its interzygomatic region, relatively wide in the less special-
ized black polecat and constricted in the white. Features of paral-
lelism in skulls of polecats with both pairs of “mink-polecat” are,
however, developed at a higher level of specialization—mink do
not attain such a height as each corresponding polecat species.
This is the parallelism of a pair of terrestrial species, in part of
1399
even desert-steppe species [polecats], and a pair of species associ-
ated with the aquatic environment. In this situation, the level of
specialization found in the polecat cannot be attained, possibly
related to smaller and weaker prey [of mink].
The significant quantity of shared characters with polecats, on
the one hand (especially American mink) and on the other—with
kolonok does not allow us to split off mink as a separate subgenus,
much less a separate genus, and also separate polecats (“Putorius’’)
at a generic level from remaining species of genus Mustela. It is
also hardly acceptable to separate mink from the genus Mustela
and to unite them, as a subgenus, with an independent genus of
polecats—Putorius (Stroganov, 1962). Mink serve as a connecting
link between polecats and other species of the genus Mustela,
especially the pair European mink—black polecat. They also pro-
duce hybrids. The American mink itself represents a branch of this
trunk, with white polecat—its summit. (У.Н.).
Geographic Distribution
Beginning with 1933, the American mink, usually in fairly large
groups, were introduced into very different parts of our country,
both in its European part, in Siberia, in the Far East, and also in
the Caucasus. At the start, captive mink from fur farms were in-
troduced, but quickly for further establishment, wild animals from
populations naturalized to various places, began to be used, in
particular, in the Altai. After the first years, introduction of ani-
mals within the range of Russian mink was stopped, or almost
stopped, and new groups of mink were directed mainly into Siberia
and the Far East where the Russian mink is absent. At the begin-
ning of the 60’s, and still earlier in some places (Altai), fairly
considerable areas occupied by American mink were formed.
However, in the majority of cases they were isolated from each
other and one continuous range of this species is still absent in our
country. There is a basis for considering that, on account of the
new introductions and the natural dispersal, it will spread over
time, although some territories are poorly suited as habitat for this
species.
As a result of absence of sufficient data on distribution of the
mink in separate locales, we will tabulate below only districts where
it was introduced, irrespective of how many times and in which
931
932
1400
parts of these regions the animals were introduced. Also,
intraregional distribution of local populations is not considered.
Starting in 1933, American minks were introduced into the
European part of the USSR in Voronezh district (first experiment—
Voronezh preserve). From 1933 to the beginning of 1963, the mink
was introduced in various quantities to Murmansk and Arkhangel’sk
districts, Karelia, in Kalininsk, Gork’ovsk, Volgogradsk, Chelyabi-
nsk and Voronezhsk districts, and into Tatarsk, Bashkirsk, Mariisk
as well as Lithuanian and Byelorussian Republics. Beyond the Urals,
animals were introduced in Sverdlovsk, Tyumensk, Omsk,
Kemerovsk, Novosibirsk, Chitinsk and Irkutsk districts, in Altai
and Krasnoyarsk territories, in Tuvinsk, Buryatsk, and Yakutsk
Autonomous Republics, into Magadansk, Kamchatsk and Amursk
districts, into Khabarovsk and Primorsk territories, into Chukotsk
national regions and in several other places, among them Sakhalin
and Urup Island in the Kurils.
In the Caucasus, introductions were in northern Osetiya, in
Georgia and Azerbaijan, and in Middle Asia—in the Tien Shan on
territory of Kirgiziya.
The most northerly place of introduction—Kola Peninsula,
lower Severnaya Dvina, mouth of Ob’ River, mouth of Nizhnaya
Tunguska, Podkamennaya Tunguska (middle course), Vilyui, up-
per Kolyma and lower Anadyr’. In the expanse between the Pacific
Ocean and upper and middle courses of the Ob’, mink are estab-
lished generally southward, extending or almost extending to the
state border. In western Siberia and in the European part of coun-
try, it extends southwards to the steppe zone. On the whole the
range is not continuous, and separate places of introduction and
separate regions of acclimatization, one may say are still isolated
from each other, in some places by great distances. Regions of
introduction in the Caucasus and Middle Asia are particularly
remote from other places.°
In a series of places, small groups of animals were introduced
in a few spots. These introductions have not yet given results in
the sense of creating sufficiently significant populations from the
perspective of being considered permanent. This applies to the
‘Materials used in the construction of the map and the considerations given were
obtained from the Main Board of Game Farm, of the Council of Ministers of RSFSR
(Chief N.B. Eliseev; L.S. Tamantseva). Some data were obtained from V.A. Popov
and G.A. Voronov).
‘1э34эН “OA “ASSN Py) Ul '4э242$ мозлм (р]215пИ/) 612151! ‘ини цеопэшу JO попопроцит JO зэпиезот ‘558 “sly 056
08
C
931 Fig. 353. Natural species distribution area of the American mink, Mustela (Mustela)
vison Schreb. (from Hall and Kelson, 1959).
Extreme North, northwestern and in part western parts of the
European USSR, as well as the Caucasus and Middle Asia. More
positive results were apparently obtained in Byelorussia. Quite
complete and numerous populations were created in the eastern
European territories of the country—in Tatariya, in part in
Bashkiriya and other adjacent places. Here populations are
of exploitable number and density. Everywhere, in regions of
habitation of the indigenous mink, i.e. to the west of the Ural, the
American mink has crowded out and exterminated it.
In Siberia and the Far East, where in several districts living
conditions are favorable for the introduced species, two quite sig-
1403
nificant populations, both in numbers and in territory occupied,
already have spread out. One occupies the Ussuri territory, includ-
ing Priamur’e, and the other—Altai and adjacent places. In Trans-
Urals (western Siberian lowland) where living conditions are less
favorable for the American mink, numerous attempted introduc-
tions did not succeed. It is evident that a significant number of
introductions into Siberia, in particular in the Extreme North and
northwest, did not lead to the formation of stable and entirely
naturalized populations.
Geographic Range outside the Soviet Union (natural).
The mink occupies the greater part of North America. The north-
ern border extends along the northern shore of the continent, ex-
cluding, however, the Ungava Peninsula. Moreover, in the expanse
from Hudson Bay to Amundsen Gulf in the Arctic Ocean, the
northern border extends approximately along the southern limit of
the tundra from Eskimo Point at Hudson Bay to its intersection of
the northern coast of the continent at 125° west long.
The western border is formed by the Pacific Coast southwards
to San Francisco and the eastern—by the Atlantic Coast, except
the eastern extremity of the Nova Scotia peninsula. The southern
border extends along the coast of the Gulf of Mexico, approxi-
mately to Corpus Christi, and thence is directed along an irregular
line across the states of Texas, New Mexico Utah, Nevada and
northern California passes to the Pacific Ocean. Mink are absent
on the Aleutian and other islands of the Bering Sea, on Kodiak, on
the Queen Charlotte islands and several other small islands along
the Pacific Coast of America. In the east, the mink is absent in
Newfoundland and the islands of the Gulf of Saint Lawrence.
In Europe the acclimatized mink is found in France, in Central
Europe (F[ederal] R[epublic] G[ermany] and the G[erman]
D[emocratic] R[epublic]), is widely distributed in the Scandinavian
Peninsula. In Cent[ral] and in part in west[ern] Europe, it has
replaced and destroyed the European mink (У.Н.).
Geographic Variation
Very many geographic forms of mink have been described from
the natural range in North America. Recent authors accept 14 races
933
1404
of the species М. vison, and moreover, а separate species
M. macrodon extinct (exterminated) about 1860 and inhabiting,
apparently, an extremely limited territory in the northeastern United
States (New Brunswick and state of Maine). The actual number of
races of the species in America, apparently, is less, and M.
macrodon—differing only in somewhat larger dimensions hardly
deserves to be split off as a separate species.
In the beginning, the various parts of our country were settled
by captive-born mink from various large fur farms. The greater
part of the material consisted of animals born in captivity in our
farms. Animals directly imported from outside our borders consti-
tuted only a minority. The origin, i.e. the races to which this material
belonged, even if we assume that at least a part of them were wild
mink, is unknown. However, they were all, apparently, captives.
Later, introduction of mink in the USSR proceeded by means of
animals captured by us from the wild.
During study of the results of acclimatization, several real
characteristics of the Tatarsk and Altaisk mink were established,
and separate subspecies were described—tatarica and altaica (see
synonymy). Difference between both “forms” lies in dimensions
(tatarica is larger) and degree of development of white marks (in
altaica, they are more, and may even appear on upper lip) and
density of pelage (in altaica, it is slightly denser). The relationship
of these “forms” to American mink is unclear.
The separation of the mentioned “subspecies” has no basis, in
that what is signified by the word “form” does not correspond to
the concept of subspecies. The word applies to very young
populations, only yet being established from genetically heterog-
enous material (tatarica—mainly from Pushkino near Moscow,
1934; altaica—Kola fur farm, 1937). At the time of their study,
they had existed in all for 14-15 years. Both populations them-
selves bore features of their “domestic” origin, in the form of
occasional mutations that are unknown in wild mink, or are ex-
tremely rare (albinism, variant of underfur color, “scorched”).
From published material (V. Popov, 1949; Ternovskii, 1958) it
is clear that populations were subjected to the action of intensive
selection and were rapidly changed—general dimensions and skull
measurement changed, as well as frequency of occurrence and size
of white marks, etc. All of this is entirely natural for animal groups
with a heredity “uncoupled” during cage raising; it cannot be used
1405
аз a basis to accept the above-mentioned populations as estab-
lished geographic races in the present sense of this word. One can
only speak about differences in these artificially created populations.
In the meantime, it is entirely unclear how they will form them-
selves, even after several decades, especially in case of union of
ranges, not to mention the time needed for elaboration of actual
subspecies.
Dimensions of mink from Tatariya (110 males and females) at
the end of the 1940’s (У. Popov, 1949), and from the Altai (41
males and 31 females), at beginning of the 50’s (Ternovskii, 1958)
are the following: body length of males from Tatariya is 375-
M417-450 mm, of females, 330-М358-371 mm; the same from
Altai—of males is 340-M395-450 mm, of females, 310-М345-—
375 mm; tail length of males from Tatariya is 195-M224—247 mm,
of females, 178—М195-215 mm; tail length of males from Altai is
156-М184-205 mm, of females, 148—M158-173 mm; length of
hind foot of males from Tatariya is 61.0-М65.0-70.0 mm, of fe-
males, 50.0-М54.5-58.0 mm; the same of males from Altai is 51-
M60-67, of females, 40-М50-55 mm; length of ear of males from
Tatariya is 22-М26-29 mm, of females, 21.4-М22.8-26.0 mm;
the same of males from Altai is 20-М22-25 mm, of females,
19- M21-23 mm.
Average measurements of mink skulls from Tatariya:
condylobasal length of males (20) is 68.95 + 0.47 mm, of females
(19), 60.65 + 0.31 mm; zygomatic width of males is 39.75 + 0.40
mm, of females, 33.9 + 0.23 mm; interorbital width of males is
15.38 + 0.31 mm, of females, 13.2 + 0.17 mm (for craniological
data on Altai mink, see above in section “Description”) (V.H.).
Biology
Population. From 1933 to 1948, about 4000 mink were introduced
into the territory of the USSR, distributed on 50 sites in 22 oblasts.
To 1964, the total population of animals introduced into the Euro-
pean part of the USSR exceeded 2000, and in Siberia and the Far
East, 14000 (V. Popov, 1964).
Concerning the results of these introductions one may judge
that in 1957—1959, in the RSFSR (excluding Yakutiya) from 26.8
to 35.7 thousand wild minks of both species were caught. Of this
number, American minks did not exceed 20-25%; i.e., the number
1406
934 Fig. 354. Arkhara river—a place of introduction and occurrence of the
American mink in Arkharinsk region of Amursk district. June 1956. Photograph by
A.G. Pankrat’ev.
of American minks caught per year was from 5350-6450 to 7150-
8925. Separate count of the yield of each species were not made;
therefore, the ratio of their number in regions where both species
existed can only be estimated very approximately. Moreover, at the
present time, the figures on skin preparation reflect not just the
934 population numbers of the species but the numbers of hunters,
their skill, the time they spend in this trade, etc. Due to the strength
of a series of causes, mink, in many districts, are hunted less than
would be allowed by their population.
More than 80% of the American minks are obtained from
Siberia and the Far East, mainly from the Altai, Krasnoyarsk and
Khabarovsk territories. From a series of indirect data, it can be
estimated that the total number of the American mink in our coun-
try apparently does not exceed 30—40 thousand.
935
1407
Habitat. With respect to choice of habitat, there are no differ-
ences between the American and European mink. In Georgia, in
the Alazan’ valley (Ekvitimishvili, 1951), mink stations* are areas
around forest creeks 1.0-1.5 m wide and 15-100 cm deep, with
steep banks freezing only along the sides. In the Altai (Ternovskii,
1955), lands favorable for mink are rich in fish, and have a good
network of spaces under the shallow ice. Banks are well protected,
steep and covered with trees containing a large amount of fallen
limbs and branches, good undergrowth, with an abundance of
mouse-like rodents. There, 3—5 mink are usually found along 1 km
of shore-line.. Satisfactory lands can be distinguished from less
favorable conditions for nesting and protection; banks are weakly
forested and with poor cover. On one km of shore-line, only 1-2
mink are encountered in such an area. Unfavorable areas are poor
in fish, or if rich, then there are no under-ice cavities and few
open water areas. In one km of shore-line in these conditions, no
more than one mink may be met with. Surroundings of the first
and second types predominate in the lower courses of small riv-
ers, and the third—in the upper reaches of rivers.
Food. Of greatest significance in foods of mink in Tatariya (V.
Popov, 1941, 1949) are voles (36% occurrence), fish (28.8%),
crustaceans (26.7%), frogs (17%) and aquatic insects (19.8%).
Importance of each food changes noticeably according to season.
In winter, foods obtained from water prevail: fish (45.5%), crusta-
ceans (32.2%) and frogs (17%); the percentage of voles decreases
to 10%. In spring, the significance of terrestrial animals increases;
occurrence of voles increases to 55.6%, of insects—to 20.7%, birds
comprise 6% and occurrence of crustaceans decreases to 18%; in
food remains, small hares are encountered. In summer, a greater
diversity of food is characteristic. Vole occurrence is high (40.6%),
crustaceans (30%), insects (29.6%), frogs (21.2%), birds (mainly
in the form of nestling) and their eggs—16.7%. In autumn, voles
occur in 50% of data, fish in 15.5-38.7%, and crustaceans in 22.4%.
Within the boundaries of montane Altai (Ternovskii, 1955), in
the diet of the American mink, mammals constituted 58.5% (occur-
rence); rodents 53.8% (including voles 50.2%), and shrews and
moles 4.7%. Birds were met with in 3.3%. Reptiles constituted
1.0%, amphibians—5.8%, fish—54.9%. Among birds, 11 different
species were recorded—most frequently, dippers [Cinclus cinclus]
*The Russian word is statsiya; stantsiya might have been meant—Sci. Ed.
1408
and pine grosbeak [Pinicola enucleator]. Among fish, small ones
prevail: minnow, gudgeon [Gobio gobio] and wide-headed sculpin
[Cottocomephorus], in all, 11 species of the 26 species known in
Altai. Mollusks constituted only 0.05%, insects 25.6% (mainly from
stomachs of fishes). Here also (Berger, 1947), in the earlier period
of acclimatization, the occurrence of insectivores in the food of
American mink constituted 3.9%, hares—5.1%, squirrels—2.6%,
mouse-like rodents 43.6%, water voles—14.1%, fish 7.7% and
insects—10.2%.
In Sverdlovsk district (L.M. Tsetsevinskii), mouse-like rodents
constituted—41.8% of mink food, water voles—25%, birds—12.5%,
fish—20.9% and insects—12.5%. In Irkutsk district (Vladimirov, 1940),
murid rodents constituted 20.5%, water voles—30.1%, birds—16.9%,
fish—15.7%, amphibians—1.2%, and insects—14.4%.
In connection with differences in the composition of aquatic
and bank fauna, a well-defined geographic variation is observed in
the foods of American mink. The absence or scarcity of river crus-
taceans in many of the water bodies of our country is particularly
evident. In the Far East, in the food of American mink, crustaceans
are replaced by fresh-water amphipods which are eaten in great
quantities (Yu.A. Salmin and V.D. Shamykin). In the first years
after introduction of American mink, living for a series of genera-
tions on farms, food obtained on land predominated, but feeding
characteristics quickly changed character, and approached that of
wild mink in type.
In North America (V. Popov, 1949), warm-blooded animals
have greater significance in the food of mink, than in the USSR
(mouse-like rodents, muskrat).
The mink eats 4-9 times daily. The quantity of food eaten
during this time constitutes 20-25% of the live weight of the ani-
mal. In Tatariya (V. Popov, 1949) and in the Altai, the American
mink makes a store of food. The size of the store reaches 1200 g
(6 frogs, 3 water voles, 1 tundra vole, 1 viper, 9 minnows and 1
wide-headed sculpin) (Ternovskii, 1955).
Home range. Dimensions of the home range of each individual
is subject to seasonal change. In mink having a settled mode of
life, area of the home range in southern Tatariya on average equaled
about 16 hectares, fluctuating from 0.1 to 40.0 hectares (V. Popov,
1949). Dimensions of daily range of activity in winter fluctuate
from 0.1 to 2.1 hectares. In summer, when food yield increases,
936
936
1409
Fig. 355. Brood shelter of American mink. Arkhara river in Arkharinsk region, Amursk
district. June 1956. Photograph by A.G. Pankrat’ev.
the home range decreases; usually only a small part of the range
is exploited, sometimes, only a radius of 50-80 т from the bur-
row. In the Altai, in winter mink live in an area of 200-300 т, not
wandering more than 100 m from the bank, and are confined to
open water (polynia) (Ternovskii, 1955). In the Far East (N.M.
Berger), 0.8—1.0 mink may patrol on 1 km of the shore-line, in
eastern Siberia—0.3—1.0; in Krasnoyarsk territory—0.04—0.10; in
Kuznetsk Alatau—0.7—0.8. In spring, the animals undertake move-
ments along the river banks to a distance of up to 20-30 km; in
autumn—up to 7-10 km.
In America (USA) in an area of 440 hectares (280 hectares of
which is dry land), 17 minks were captured and tagged (14 males
937
1410
and 3 females); 5 of them (29.4%) were caught а second time in
the same winter. During the following 4 years (winters of 1945/48)
in the same area, only 6 new minks were caught, out of which 2
were repeatedly caught (4 times). Males were considerably more
active than females and the area of 440 hectares was insufficient
for them; one male, in the course of three years moved on to the
area every month, making a round trip of about 1.5 km in one and
the same route and then returned. Females were less active—re-
peated hunting was achieved within a distance of from 25 to 400
m from the place of marking (McCabe, 1949).
Burrows and shelters. In the Volga-Kama territory, there are 4
types of burrows of American mink (У. Popov, 1949). They are _
constructed frequently in the hollows in root masses of growing
trees, and rarely—in hollows of fallen ones. The holes are often
in the root masses of hummocks (“koblakh’)* of the alder which
are typical for flood-land alder swamps. These holes are charac-
terized by a large number of entrances and twisting passages.
Rarer are earth holes of structure similar to that of the European
mink. The number of exits of the mink burrow varies from 1 to
8. In the breeding burrow, exits are often stuffed with grass so
young cannot crawl away. Usually, the breeding burrows are situ-
ated not more than 20 m from the bank of the water body. Bur-
rows of single animals are also encountered (considerably more
rarely) at a distance of up to 50-60 т. The bedding of the nesting
chamber consists of dry grass and leaves, wool and underfur of
consumed animals and wood dust.
Out of 24 mink shelters investigated in the Altai (Ternovskii,
1955), 54% were found in hollows of fallen trees, 34%—in intraroot
hollows among the roots of trees, 4%—in hummocks and 4%— т
openings among rocks and soil layers. Mink shelters, in the latter
case, are found in sections of floodlands which are not submerged
with water, at 2-10 m from the water level, but no more than 50
m from it. The bedding in the nesting chamber usually consists of
sedge, Carex pediformis (“puzzle grass’).
Daily activity and behavior. The mink is active at various times
of day. The rhythm of diel activity changes in different seasons in
relation to security of food, prevailing means of obtaining it, weather
conditions, etc. Thus, in the rut period, animals are highly active
*Local word?—Sci. Ed.
1411
throughout entire 24 hr period. For the period of pregnancy, daily
activity typically shows two peaks. In the period of nursing young,
the female is active most of the 24 hr period. In summer, the litters
are active mainly at twilight. Later in autumn, the activity during
the day light hours again increases (Ternovskii, 1955).
In Tatariya, in American mink in winter (January) three peri-
ods of activity are observed with an interval between the peaks of
approximately 8 hours (V. Popov, 1949). The first—a daylight
period, lasts from 10 until 14 hr and is associated with obtaining
food in water bodies. The second—at twilight, begins at 18 hr,
somewhat before nightfall. The third occurs in the second half of
the night and the pre-dawn period. During severe frosts, activity
decreases. The mink is maximally active in July-September, and
minimally in November—February. Males are more active in March
and less—from the end of rut until August. Females are maximally
active in July-September. In winter, in a 24-hr period, the mink
moves from some tens of meters to 4-5 kilometers.
In the Altai, the mink is most active morning and evening, and
also at the end of the night (Ternovskii, 1955). In winter and
summer, activity is less than in autumn and spring. Rut and disper-
sion occur in spring, and in autumn, dispersion of young and adults,
searching for home ranges, and then mink are maximally active. In
cloudy calm weather, activity grows. In winter, with air tempera-
ture from —20 to —37°C, the mink is active for not more than 51-
64 minutes during the 24 hr period. In December—January, it leads
a hidden, subnival mode of life. If in October and the first half of
November, the length of the 24 hr trail reaches 800-1000 т, then
during the period of deep snow, the daily trail decreases by 3-6
times and does not exceed 100 m.
During swimming, undulating movements of the trunk play the
main role. The swimming web is no more developed than in er-
mine, weasel and kolonok. In warm water (24°C), mink can swim
for more than 3 hours without stopping. In cold water, animals
died within 27 min. During this time, body temperature fell from
40°C to 19-20°С. With water temperature up to 8°C, the body of
the mink cooled down under water for 118 minutes, while in er-
mine and forest polecat—in 26-28 minutes (Ternovskii, 1955).*
Seasonal migrations and transgressions. In winter, the charac-
ter of activity in mink depends upon the abundance of food and its
*The mink body is better insulated in water—Sci. Ed.
1412
938 Fig. 356. Tracks of American mink on fresh new-fallen snow. Fudzin river, western
slope of Sikhote-Alin’. December 1956 (Photograph by A.G. Pankrat’ev).
availability. Complete freezing of water bodies, absence or small
number of polynias,* air holes and other opportunities to penetrate
under ice call forth emigrations and sometimes considerable con-
centrations of mink in those places where food is more abundant
and available. In areas of the meadow type with little food, some
minks lead a migratory mode of life, performing movements of
2-3 or up to 4—5 km in a 24-hr period in an area of floodland of
up to 60 hectares (in the presence of abundant food, minks do not
move away from the burrow more than 50-100 m). Migratory
animals restrict themselves to one place for several days, and then
perform a movement to settle down in a new place, again for
several more days. Young mink, dispersing, also undertake migra-
tions. More significant translocations were observed т
the first period after introduction of mink, especially in cases of
*Ice-free water surface—Sci. Ed.
938
1413
unsuccessful choices of places. Mink moved from the introduc-
tion site a distance of 15 km (Tatariya) to 40-60 km in Siberia,
and for a distance of up to 100-120 km in the Far East.
Reproduction. In our country, estrus in American mink pro-
ceeds from the end of February to the beginning of April. For the
most part, mating takes place in mid-March (Kler, 1941); (V. Popov,
1941, 1949), i.e. a month earlier than Russian mink. Estrus in
separate individuals lasts one month, with 6-8 day intervals. The
percentage of fertilized females equals 95-100. Duration of preg-
nancy is 51.3 days on average, fluctuating from 40 to 65 days
(Kler, 1941). Differences depends on duration of the latent period
in embryonic development, which lasts from 2—3 weeks to 1 month.
After formation of the placenta, development goes on for 25-26
days. Embryonic mortality is great and reaches 60-70%. It is most
of all determined by magnitude of litter and depends upon external
conditions of environment which control condition of the female
organism. Birth takes place during the period from 4 to 22 May.
In the Altai, the first signs of spermatogenesis were noticed in
January. Already by 10-15 March, all stages of spermatogenesis,
and sperm were in the tubules of the epididymis. On 1 February,
pre-estrus proliferation of the uterus was observed in the female
(Ternovskii, 1955). The first signs of rut were noticed at the end
of February. It proceeded throughout March and the first half of
April; peak of rut—the second ten days of March. Its general
duration here is 45 days.
The number of young in the litter fluctuates from 1 to 9, and
in exceptional cases, reaches 11 (Tatariya) and 16 (USA). In south-
ern Tatariya, average litter size fluctuated in various years from
3.2 to 5.8 (V. Popov, 1941).
Growth, development and molt. Weight of new-borns fluctu-
ates from 6 to 11 g. They are born blind, but develop rapidly, and
eyes open at age of 30-31 days. Differences in increase in weight
and dimensions between males and females are clearly noticeable
only after 40 days.
The lactating period lasts for 2-2.5 months. Mink’s milk con-
tains 3.8% lipids, 6.2% protein, 4.6% sugar and 10.66% mineral
salts. Already at the age of 20-25 days, young minks begin to try
food brought by the mother. In autumn, they attain adult dimen-
sions.
939
1414
On farms, minks live 7-10 years, but are usually killed for
their skins before reaching their age limit.
Enemies, diseases, parasites, mortality, competitors, and popu-
lation dynamics. The larger, stronger American mink ousts and
depresses the European.
There are little data concerning population dynamics, and these
data are not always reliable. The history of establishment of mink
in the Altai gives some conception of growth rate of the trans-
planted populations. Acclimatization was started in 1937. In 1937-
1941, 248 minks were introduced. During the period 1940-1953,
431 minks were transferred from one region to another, to accel-
erate the transplantation process. As a result, during a period of 19
years, the American mink settled in hundreds of rivers and streams
and occupied a territory of 79,400 km*. By the beginning of the
1960’s, they numbered several thousands (Ternovskii, 1955).
Field characteristics. It is not always easy to distinguish be-
tween the tracks of American mink and European. Jumping with
paired paw prints is a typical gait for the European mink. Length
of the jump is 25-35 cm. In American mink, tracks on the snow
most often form a group of 3—4 paw prints. Paired tracks and the
unique pace forming a chain of tracks are rarely observed. Length
of a jump is significantly greater—55—60 cm.
In the Altai, maximum jump length of the American mink is
1200 cm, while running, the mink can jump to a height of 50 cm.
It can attain a speed of 20 km per hour for short distances; at a
moderate walk—8-12 km, and in slow movement—3-5 km. The
mink cannot run in loose snow deeper than 15 cm.
Tracks of the paws of males are always larger than females;
the male track is equal to 4 x 3.5 cm on average, but the female
track is 3.5 x 3.0 cm. While moving slowly, the length of the
female jump is always shorter than the male. Sex can also be
determined by the characteristics of urination on the snow. The
weight load on 1 cm? of foot surface is large, 15-31 g (average of
20 samples—20 g). Weak fur covering of the foot prevents snow
freezing on it. “Sliding hills” and grooves in mink trails, as in
otters, are explained by their need to dry the fur in winter after
coming out of water (Tarnovskii, 1955).
Identification of tracks and burrows of mink may also be de-
termined by contents of excrement and food remains near entrances
to burrows (P.Yu.).
1415
Practical Significance
Acclimatization of the American mink in the USSR achieved its
colonization in water bodies where it was previously absent for
some reason or where it was destroyed. The American mink, as
an object of the fur trade, gives a larger and higher quality skin
than the Russian mink and its settling in water bodies, mainly in
montane regions of southern Siberia and Far East, may increase
the commercial yield to us of valuable “colored” furs.
Trade in American mink in the USSR began in Tatariya in
1938. At the present time, the animal is mainly used for further
settling by means of its capture in live traps.
In some parts of North America, the mink destroys muskrats in
quite significant quantities, especially the young. There are analo-
gous complaints in Siberia.
In past years, intensive breeding of the American mink in farms,
both abroad and in the USSR, has pushed hunting of wild mink to
a secondary position (P.Yu.).
940-
982
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990
Index of Latin Names of Taxa*!
A
aberrans cf. Meles
abyssinica cf. Mellivora
admirata cf. Mustela
adustus cf. Canis
Aeluroidae 54
aestiva cf. Mustela
Ailuridae 54
Ailurinae 911
Ailuropoda 54, 402, 907, 908, 909, 911
Ailuropodidae 54
Ailurus 54, 402, 907, 908, 909, 911
alhperakyi cf. Vormela
Alopex 62, 63, 68, 99, 194, 194, 195, 196,
268
— beringensis 196, 207
beringianus, 196
— fuliginosus 208
— groenlandicus 208
— halensis 208
— innuitus 208
— lagopus 56, 63, 65, 195, 196, 196, 197,
199, 202, 205, 206, 207, 208
— pribilofensis 208
— semenovi 196, 207
— spitzbergenensis 196, 206, 208
— ungava 208
alpinus cf. Mustela, Kolonokus
altaica cf. Mustela
Amblonyx 498, 853
— cinerea 853
americana cf. Martes
Amphicyoninae 62
Amphicynodontinae 62, 402
amurensis cf. Meles
anakuma cf. Meles
Anurocuon 383
Anurocyon 383
— clamitans 383
Aonyx 492, 498, 854, 883
— capensis 853, 854
— cinerea 851, 853, 854
— hessica 883
Arcticonus 404
Arctocyonidae 54
Arctogale 636, 640
— erminea 663
— tobolica 663
Arctoidea 54
Arctonyx 493, 498, 817
arctos cf. Ursus
arenarius cf. Meles
Artiodactyla 10, 11, 54
asiatica cf. Martes
aureus cf. Canis, Mustela
aurobrunnea cf. Lutra
australis cf. Dusicyon, Mustela
B
baicalensis cf. Ursus
barang cf. Lutra
Bassariscus 69, 907, 908, 911
Bassaricyon 911
beringiana cf. Ursus
beringianus cf. Ursus
birulai cf. Mustela, Kolonokus
boccamela cf. Mustela
Borophaginae 62
bosniaca cf. Martes
brachyura cf. Martes
*Reproduced from the Russian original. Page numbers of the Russian origi-
nal appear in the left-hand margin in the text—General Editor.
‘Page numbers with description are given in bold and those with illustrations, in
italics.
991
1540
Broiliana 817
brokmani cf. Mellivora
С
Caenolestoidea 9
canescens cf. Meles
Conepatus 498
Canidae 54, 55, 57, 58, 60, 6/, 62, 69, 272,
402, 907, 911
Caninae 62, 63, 66
Canis 53, 62, 63, 68, 96, 96, 98, 99, 132,
194, 268
— adustus 63, 99
— albus 123, 136
— altaicus 139, 139, 142
— alopex 318
— algirensis 108
— alpinus 63, 383, 386
— arabs 145
— argunensis 124, 142
— arundinaceus 123, 139
— aureus 63, 64, 96, 100, 100, 101, 102,
105, 107, 108, 108
— australis 63
— australis = antarcticus 53
— bactrianus 141
— baileyi 146
— beringiana 319
— bernardi 137, 146
— campestris 123, 139, 140, 142
— caragan 318
— caucasica 100
— caucasicus 108
— chanco 123, 141, 142
— communis 123, 137
— coreanus 124
— corsac 265
— crucigara 319
— cruesemanni 108
— cubanensis 124, 140
— deitanus 145
— desertorum 123, 141, 142
— dingo 52, 53, 61, 98, 99
— dorogostaiskii 124, 142
— dukhunensis 383
— dybowskii 124, 136
— ecsedensis 108
— familiaris 53, 56, 96, 100
— gregoryi 146
— hajastanicus 124, 140
— hattai 124, 143
— hodophylax 145
— hungaricus 108
— indicus 108
— italicus 145
— kamtschaticus 124, 136
— kurjak 145
— lagopus 194, 196
— laniger 123, 141
— lanka 108
— latrans 63, 100
— lepturus 383
— lupaster 108
— lupus 56, 59, 63, 64, 96, 97, 100, 123,
123, 124, 125, 129, 132, 133, 135, 136,
137, 137, 138, 139, 139, 140, 141, 141,
142, 142, 143, 145, 146
— lycaon 146
— mackenzii 145
— manningi 137, 146
— marocanus 108
— melanotus 319
— mesomelas 63, 99, 108
— minor 108
— montana 319
— moreotica 100, 108
— naria 108
— niger 97, 99, 146
— orientalis 124
— pallipes 123, 141, 145
— pambasileus 145
— primaevus 383
— procyonoides 69
— rex 124, 143
— rufus 146
— schensi 53
— simensis 63
— soudanicus 108
— signatus 145
— syriacus 108
— typicus 100, 108
— tschiliensis 142, 145
— tundrarum 146
— turuchanensis 124, 136
— vulpes 265, 318, 319
Canoidea 54, 55, 58, 402, 497, 498, 907,
911
caparioccoi cf. Mustela
capensis cf. Mellivora
Capreolus capreolus 837
Carnivora 10, 11, 49, 52, 53, 60, 907
caucasica cf. Mustela
caucasicus cf. Meles, Ursus
cavifrons cf. Ursus
Cetacea 10, 11
Cephalogale 402
Cervus elaphus 837
chamus cf. Fennecus
chanco cf. Canis
Charronia 498, 502, 505, 508, 604
chinensis cf. Lutra
Chiroptera 10, 11
Chrysaeus 383
Chrysocyon 62
— brachyurus 59, 62, 63
cinerea cf. Amblonyx
cinereo-argentatus cf. Dusicyon
Clethrionomys rufocanus 521
— rutilus 521
collaris cf. Ursus
colore griseo cf. Vulpes
Conepatus 493, 498
consica cf. Mellivora
coreana cf. Mustela
coreensis cf. Martes
Creodonta 54
crucigera cf. Vulpes
culpaeus cf. Dusicyon
Cuon 60, 62, 63, 383, 383, 384, 386
— adustus 392
— alpinus 60, 63, 384, 385, 386, 386, 388,
390, 391, 392, 392
— dukhuneusis 392
— fumosus 392
— hesperius 392
— infuscus 392
— jason 386, 392
— javanicus 386, 392
— laniger 392
— lepturus 392
— primaevus 392
— sumatrensis 392
Cuoninae 62
cylipena cf. Mustela
Cynalopex 265
Cynarctinae 911
Cynodictis 60
Cynodon 402
Cynodontidae 402
Cynaonyx 854
D
Dermoptera 10
Desmostyliformes 23
desertorum cf. Canis
1541
— dinniki cf. Ursus
desertorum — pallipes cf. Canis
Dinocyon 402
Diprotodontia 9
dolichocrania cf. Vulpes
Dugong 17, 23, 24
— dugong 20, 21, 24
Dugongidae 17, 23, 24
Dusicyon 62, 68, 268
— australis 63
— cinereo-argentatus 63, 67
— culpaeus 63
— hagenbecki 63
E
— ecsedensis cf. Canis
Edentata 10
Eira 498
Embrithopoda 23
Enhydra 50, 492, 493, 498, 499, 854, 882,
882, 883
— gracilis 895
— kamtschatica 884, 895
— lutris 854, 863, 883, 884, 885, 889, 891,
894, 894, 895
— marina 894
— nereis 895
— orientalis 895
— reevi 883
— stelleri 895
Enhydriodon 854
Enhydris 882
992 Eomellivora 803
Eotheroides 24
Euarctos 403, 404, 406
Eumetopias 53
Eumustela 637
Euprocyon 912
Eutheria 10
euxina cf. Vormela
eversmanni cf. Mustela
Eyra 493
F
Felidae 54, 55, 57
Felis catus 56
— libyca 56
— manul 50
— sylvestris 56
1542
Feloidea 54, 55, 497
Fennecus 62, 68, 268
— chamus 63
— pallidus 63
— riippeli 63
— zerda 59
— zerdus 63
Ferae 10, 49
ferghanae cf. Mustela
Ferungulata 10, 54
Fissipedia 53, 54, 60
flavescens cf. Vulpes
Foetorius 636
— foetorius 745
Гоша cf. Martes
fontanieri cf. Mustela
formosanus cf. Ursus
fuliginosus cf. Alopex
G
Gale 636
Galictis 493, 498
gedrosianus cf. Ursus
Glires 10
glogeri cf. Mustela
Grammogale 498
gregoryi cf. Canis
griffithi cf. Vulpes
Grisonella 498
groenlandicus cf. Alopex, Ursus
Gulo 493, 498, 500, 501, 614, 614, 616,
617, 641, 803
— albus 629
— arcticus 618, 628
— arctos 618, 628
— biedermanni 618, 628
— borealis 618, 628
— gulo 58, 617, 618, 618, 619, 622, 624,
628, 628, 629, 629
— kamtschaticus 618, 629
— katschemakensis 629
— luscus 618, 629
— luteus 629
— schlosseri 616, 618
— sibiricus 615, 618, 628
— vancouverensis 629
— vulgaris 618, 628
— wachei 618, 628
Guloninae 641
gwatkinsi cf. Martes
gyas cf. Ursus
H
hagenbecki cf. Dusicyon
Halicore 25
Halitheriinae 23
Halyanassa 42
hamgyensis cf. Martes
Helarctos 397, 398, 400, 403
— malayanus 403, 483
Helictis 493, 498, 817
heptneri cf. Meles, Mustela
Herpestes auropunctatus 52
— javanicus 52
hoole cf. Vulpes
huli cf. Vulpes
hungarica cf. Mustela
hungaricus cf. Canis
Hyaenidae 54, 55, 57, 60
Hyaenodontidae 54
Hyracoidea 10, 22, 54
Hydrictis 853
Hydrodamalidae 17, 23, 24, 25
Hydrodamalis 18, 23, 25, 25
— gigas 25, 27
— stelleri 25
Hydromustela 637
I
Ictis 636
— boccamela 644
— kamtschatica 644
— nikolskii 644
— nivalis 644
Ictonyx 498, 783
inaurita cf. Mellivora
indica cf. Mellivora
indochinensis cf. Martes
mesomelas cf. Canis
Insectivora 10, 11, 54
isabellinus-syriacus cf. Lutra
itatsi cf. Mustela
intermedia cf. Martes, Vormela
J
japonicus cf. Ursus
jessoensis cf. Ursus
K
kalmykorum cf. Vulpes
katschemakensis cf. Gulo
1543
Kolonokus 673, 640, 690, 691, 723 — gracilis 884
— alpinus 664, 686 — isabellinus-syriacus 868
— australis 700 — incarum 853
— birulai 664, 686 — kamtschatica 855, 867
— coreanus 700 — kutab 855, 869
— raddei 686 — lutra 853, 854, 855, 856, 858, 859, 862,
— sibiricus 700 865, 866, 866, 867, 868, 869
kolymensis cf. Ursus — maculicollis 853
koreana cf. Martes — marina 884
koshevnikovi cf. Vormela — meridionalis 867
kuatunensis cf. Martes — minor 718
kutab cf. Lutra — mira 853
— monticola 855, 868
993 L — nair 865, 869
— nudipes 854
— oxiana 855, 867
— perspicillata 853, 865
— platensis 853
— provocax 853
— seistanica 855, 867
— stejnegeri 855, 867
— stelleri 884
labradorensis cf. Ursus
Lagomorpha 10, 11
Lamprogale 502, 604
laniger cf. Ursus, Lupus
larvata cf. Mustela
lasiotus cf. Ursus
Е — sumatrana 853
Leptarctinae 497
Leptonyx 785 — vulgaris 854, 855, 867
3
leptorhynchus cf. Meles О tz 7
— 1
Leucocyon 194 a es aes
й — borealis 719
leuconota cf. МеШуога
— europaea 718
leuconyx cf. Ursus
— lutreola 719
leucurus cf. Meles у
— vison 924
libyca cf. Poecilictis
lineiventer cf. Mustela
Lupulella 99
Lupus 96
— altaicus 124
Lutrinae 497, 498, 817, 851, 853
Lutrogale 498, 853
Lutronectes whitelyi 855, 863
Lycalopex 62, 68
; — microtis 62
— filchneri 124
karanorensis 124 Ве
р Гусаоп 58, 60, 62, 63, 386
— laniger 123 ae
УЕ — microtis 63
— tschiliensis 124 р
— pictus 59, 60, 63
luscus cf. Gulo
— thous 63
luteus cf. Gulo
Lutra 493, 498, 500, 500, 501, 851, 851,
852, 853, 854, 854, 882, 883
— amurensis 855, 867
— angustifrons 868
Lycaoninae 62
lymani cf. Mustela
Lyncodon 493, 498
— annectens 853 M
— aurobrunnea 855, 869
— baicalensis 855, 867 mackenzii cf. Canis
— barang 866, 869 Mammalia 10
— canadensis 853 Manati balaenurus 25
— chinensis 868 — gigas 25
— enudris 853 manchurica cf. Mustela
— felina 853 mandchuricus cf. Ursus
1544
marinus cf. Ursus — milleri 598
maritimus cf. Ursus — nehringi 587, 597
Marsupialia 10 — notialis 596
Martes 493, 498, 50/, 501, 502, 502, 503, — obscura 508, 532
504, 505, 507, 516, 590, 595, 615, 616, | — осшасеа 507
637, 641 — ognevi 585, 597
— abietum 553, 568 — pennanti 503, 505, 516, 590
— alba 507, 596 — peninsularis 611
— altaica 508, 530, 585, 597 — princeps 531
— americana 504, 505, 516, 590 — rosanovi 585, 587, 596
— angarensis 508, 531 — ruthena 553, 586
— arsenjevi 508, 532 — saba 611
— asiatica 507 — sabaneevi 553, 568
— aterrima 610 — sahalinensis 508, 532
— averini 530, 580 — schantaricus 532
— baicalensis 531 — sajanensis 508, 531
— brachyura 533, 533 — sylvatica 568
— borealis 553, 568 — sylvestris 568
— bosnica 585, 598 — syriaca 598
— brachyura 532 — szetchuensis 611
— bunites 598 — tomensis 508, 530
— chrysospila 611 — toufoeus 598
— coreensis 508 — szetchuensis 611
— domestica 502, 596 — tsuensis 533
— fagorum 596 — tungussensis 508, 531
— ferruginea 507 — uralensis 553, 569
— flavigula 503, 505, 506, 590, 604, 605, — vitimensis 508, 532
607, 609, 610, 610, 611 — vulgaris 553
— foina 505, 506, 506, 507, 585, 586, 587, | — yeniseensis 508
589, 593, 596, 597, 598, 602 — yuenshanensis 611
— fusco-flavescens 507 — zibellina 50/1, 505, 506, 506, 507, 507,
— gwatkinsii 505, 611 508, 513, 518, 523, 528, 530, 531, 532,
— hamgyensis 508 533, 533
— ilimpiensis 508, 531 Martinae 505, 541
— indochinensis 611 Megalotis 268
— intermedia 597 Melodon 817
— jacutensis 508, 532 Melanarctos 404
— jeniseensis 531 — cavifrons 404, 408
— kamtschadalica 533 Meledes 816
— kamtchatica 533 — leptorhynchus 816
— koreana 611 Meles 493, 494, 498, 500, 500, 803, 816,
— koreensis 533 817, 818, 818
— kozlovi 598 — aberrans 819, 833
— kuatunensis 611 — altaicus 819, 833
— latinorum 569 — amurensis 817, 819, 836, 837
— leucolachnea 597 — anakuma 817, 819, 836, 837
— lorenzi 553, 569 — arcalus 817, 837
— martes 505, 506, 507, 517, 553, 553, — arenarius 818, 819, 834, 835
554, 557, 562, 563, 567, 568, 569, 569 — atavus 818
— maculata 507 — blanfordi 819, 837
— mediterranea 598 — canescens 819, 833, 836, 837
994 — melampus 505, 533 — caucasicus 819, 832
— melli 611 — danicus 837
Meles europaeus 819
— gennevauxi 818
— heptneri 819, 832
— leptorhynchus 819, 830, 833, 837
— leucurus 817, 837
— marianensis 837
— melanogenys 819, 837
— meles 493, 817, 8/8, 818, 819, 819, 820,
823, 826, 830, 832, 833, 833, 834, 835,
835, 836, 836, 837
— minor 819, 833
— ponticus 819, 837
— raddei 819
— rhodius 837
— schrenkii 819, 836
— severzovi 819, 833, 833, 835
— sibiricus 819, 833
— talassicus 819, 835, 836
— tauricus 819, 832
— taxipater 818
— taxus 819
— thordi 818
— tianschanensis 819, 835, 836
— typicus 819
meles cf. Meles
Melinae 498, 802, 803, 816, 851, 854,
Melogale 498, 817
melli cf. Martes
Mellivora 493, 498, 499, 500, 801, 801,
802, 802, 803
— abyssinica 812
— broekmani 812
Mellivora buchanani 812
— capensis 804, 804, 805, 807, 809, 812,
812
— consica 812
— cottoni 812
— inaurita 812
— indica 812
— leuconota 812
— makwelli 812
— pumilio 812
— га 804
— signata 812
— sigulata 812
— vernayi 812
— wilsoni 804, 812
Mellivorinae 498, 803
Мешгзи$ 50, 399, 403
— ursinus 403, 483
Mephitinae 498, 803
Mephitis 493, 498
meridionalis cf. Lutra, Ursus
1545
Mesaxonia 10
Mesonychidae 54
Metatheria 10
Miacidae 54
Miacoidea 54
michnoi cf. Mustela
Microaonyx 853
microdon cf. Paraonyx
middendorffii cf. Ursus
macrotis cf. Lycalopex
minor cf. Canis
minuta cf. Mustela
Miosireniae 23
mongolica cf. Mustela
Monotremata 10
montanus cf. Vulpes
monticola cf. Lutra, Vulpes
moreotica cf. Canis
mosanensis cf. Mustela
mupinensis cf. Ursus
Mustela 493, 498, 501, 505, 639, 636, 636,
641, 643, 848, 690, 691, 704, 705, 723,
724, 781, 783, 787, 929
— aestiva 663, 673
— admirata 774
— alascensis 677
— alba 585
— albica 719, 731
— algirica 671, 676
— alleghaniensis 657
— alpina 696
— alpinus 691
995 — altaica 641, 643, 648, 686, 686, 689,
690, 692, 694, 695, 696, 697, 697
— amurensis 508, 773
— angliae 749
— anguinae 676
— arctica 675, 675, 676
— asaii 712
— asiatica 507
— astutus 697
— aterrima 604
— aurea 771
— aureola 749
— aureus 748, 770
— australis 710
— baicalensis 508
— balkarica 664, 673
— bangsi 676
— baturini 664, 675
— biedermanni 732
— binominata 719, 730
— birulai 664, 674, 695
1546
Mustela boccamela 644, 650, 654 — itatsi 700, 706, 711
— borealis 553, 604, 719, 730 — kadiacensis 676
— brachyura 507 — kamtschatica 508, 652, 675
— budina 719, 731 — kaneii 663, 675
— caledoniae 749 — kanei 675
— campestris 657 — karaftensis 644, 652
— canigula 712 — karaginensis 664, 675
— caparioccoi 697 — katian 641, 690, 697
— caucasica 655, 730 — kerulenica 644, 653
— caucasicus 719 — koreensis 508
— celenda 677 — larvata 770, 773
— charbinensis 701, 712 — leucolachnea 585
— cicognanii 679 — lineiventer 754, 772
— coreana 712 — longstaffi 686, 697
— cylipena 718, 731 — lutra 854
— davidiana 712 — lutreola 637, 641, 643, 691, 718, 718,
— dauricus 772 719, 720, 722, 726, 728, 729, 730, 731,
— digna 664, 675 731, 732, 924, 925
— dinniki 655 — lutris 882, 884
— dombrowskii 644 — lymani 667, 674, 676
— domestica 585 — macrodon 932
— erminea 636, 637, 641, 642, 648, 663, — manchurica 700, 710, 711, 712
663, 664, 665, 666, 669, 671, 672, 673, | — martes 553, 585
674, 674, 675, 676, 676, 677, 737 — martinoi 664, 674
— eskimo 657 — melampus 507
— eversmanni 638, 640, 641, 642, 705, — michnoi 772
719, 742, 743, 744, 753, 753, 754, 755, | — miles 700, 710
758, 760, 761, 763, 768, 771, 772, 773, °— minima 676
774, 781, 929 — minor 729
— fagorum 585 — minuta 650
— fallenda 677 — mongolica 677
— ferghanae 674, 676 — mosanensis 644, 657
— flavigula 502, 533, 604, 719 — mosquensis 737, 749
— foetidus 749 — moupinensis 712
— foina 585, 587 — murica 676
— fontanieri 712 — namiyei 644, 652, 657
— frenata 640, 641 — naumovi 664, 675
— furo 56, 736, 743, 749 — nehringi 585
— gale 644, 654 — nigripes 641, 761, 769, 770, 774
— galinthias 657 — nikolskii 644, 655
— glogeri 719, 731 — nippon 663, 676
— gulo 614, 618 — nivalis 51, 636, 637, 640, 641, 642, 643,
— gulpsa 677 643, 644, 645, 647, 649, 650, 651, 652,
— heptapotamica 771 652, 653, 653, 654, 655, 655, 656, 656,
— heptneri 644, 654, 659 657, 662, 719
— herminea 663 — nobilis 771
— hibernica 676 — novikovi 719, 730, 713
— hodgsoni 712 — nudipes 641, 690
— hungarica 719, 731, 754, 771 — numidica 657, 671
— iberica 657 — occidentalis 771
— initis 677 — ognevi 664, 674, 737, 749
— intermedia 585, 587 — olympica 677
— invicta 677 — orientalis 663, 675, 737, 749
Mustela pallida 656, 771
— peregusna 781, 783
— polaris 676
— princeps 507, 531
— punctata 644, 653
— putorius 56, 501, 636, 641, 642, 719,
721, 724, 736, 737, 738, 740, 743, 744,
748, 749, 749, 754, 760, 761, 929
— pygmaea 51, 644, 652, 653, 656
— quelpartis 712
— raddei 691, 696
— richardsonii 676
— ricinae 676
996 — пхоза 641, 644, 650, 657
— robusta 754
— rothschildi 737, 749
— rupestris 507
— russeliana 657
— sacana 695, 698
— salva 677
— sarmatica 781, 783
— schnitnikovi 664, 676
— seclusa 677
— semplei 676
— sho 712
— sibirica 637, 641, 643, 675, 690, 700,
700, 701, 701, 703, 706, 706, 709, 710,
710, 711, 711, 712, 724, 754, 837
— sibiricus 772
— stabilis 676
— stantschinskii 749
— stolizkana 644, 657
— streatori 677
— strigidrosa 638, 641
— subpalmata 657
— sylvatica 553
— sylvestris 553
— talassica 772
— tatarica 924
— teberdina 664, 673
— temon 686, 697
— tiarata 774
— tobolica 674
— tonkinensis 657
— transbaicalica 663, 675
— transsylvanica 719, 731
— trettaui 644, 655
— turovi 730
— tuvinicus 772
— varina 719, 731
— vison 56, 641, 643, 723, 724, 924, 925,
928, 928, 930, 931, 932
— vulgaris 636, 637, 644, 650, 655, 656,
1547
749
— wyborgensis 718, 729
— yesoidsuno 644
— zibellina 56, 502, 507, 508
Mustelidae 54, 55, 57, 58, 491, 496, 497,
500, 785, 853, 911, 924
Mustelinae 497, 498, 502, 637, 641, 801,
803, 817
Mutica 10
Mydaus 817
Mylarctos 404
Myopus schisticolor 521
Myrmarctos 404
— eversmanni 404, 408
Mystacoceti 9
N
nair cf. Lutra
namijei cf. Mustela
Nandinia 49
Nasua 908, 911
Nasuella 911
Nectogale 497
negans cf. Vormela
Nepus 25
— stelleri 25
niger cf. Canis
nigricans cf. Vulpes
nivalis cf. Mustela
Nyctereutes 60, 67, 63, 66, 67, 68, 68, 69,
911
— amurensis 69, 75
— kalininensis 69, 75
— koreensis 75
— orestes 75
— procyonoides 59, 63, 64, 69, 69, 70, 71,
74, 75, 75
— ussuriensis 75
— viverrinus 66, 75
O
obscura cf. Vormela
Odontoceti 9
ognevi cf. Martes, Mustela, Putorius
orientalis cf. Canis, Mustela
ornata cf. Vormela
Otariidae 53
Otocyon 50, 58, 60, 62, 63
— megalotis 63
Otocyoninae 62
997
1548
Oxyaenidae 54
Oxygous 96
Р
Paenungulata 10, 15, 23, 24
Paleomeles 817
pallidior cf. Vormela
pallidus cf. Fennecus
pallipes cf. Canis
pambasileus cf. Canis
pamirensis cf. Ursus
Paralutra 854
Parameles 817
Paraonyx 498, 853, 854, 883
— congica 853
— microdon 854
— philippsi 854
Parataxidea 817, 818
Paraxonia 10
peregusna cf. Vormela
Perissodactyla 10, 11, 54
peninsularis cf. Martes
philippsi cf. Paraonyx
Pholidota 10
Pinnipedia 10, 11, 53, 54
Pteronura 498
piriformis cf. Putorius
piscator cf. Ursus
planicola cf. Vulpes
Plesiogale 640
Plesiogulo 616
— brachygnatus 616
— monspessulanum 616
Plesiomeles 817
Poecilictis 498, 783, 785
— libica 783
Poecilogale 498, 783, 785
Polyprotodontia 9
Potamotherium 854, 883
Potos 907, 908, 911
Prosimiae 9
Primates 10
Proboscidea 10, 23, 24, 54
Procynodictis 60
Procyon 69, 908, 910, 911, 912, 912
— cancrivorus 912, 914, 919
— юг 56, 912, 913, 913, 915, 916, 917,
919
Procyonidae 54, 55, 57, 60, 69, 402, 497,
907, 909, 911
Procyoninae 911, 912
procyonoides cf. Nyctereutes
Prorastomidae 23
Protosirenidae 23
Proteles 51
Prototheria 10
Protungulata 10
pruinosus cf. Ursus
Pteronura 853, 854, 883
— brasiliensis 494, 853
Pusa 882
— orientalis 884
pusilla cf. Vulpes
Putorius 636, 640, 641, 705, 723, 724, 736
742, 783, 929
— alpinus 686
— amurensis 754
— arcticus 663
— astutus 686
— aureus 754, 761, 772
— caucasicus 644
— dauricus 754
— dinniki 644
— erminea 663
— eversmanni 753, 754
— ferghanae 663
— foetidus 736
— fontanieri 700
— furo 743
— heptapotamicus 754
— kamtschatica 663
— kaneii 663
— kanei 663
— larvatus 753, 770
— michnoi 753
— nivalis 644
— nobilis 754
— occidentalis 754
— ognevi 737
— orientalis 737
— pallidus 644, 754
— piriformis 737
— putorius 736, 737, 739, 754, 761, 767,
772
— pygmaeus 644
— satunini 754
— sibirica 663
— sibiricus 754
— stantschinskii 736, 739, 767
— talassicus 754
— tuvinicus 754
— verus 736
— vulgaris 736
putorius cf. Mustela
?
pygmaea cf. Mustela
R
rixosa cf. Mustela
Rodentia 10, 11
Rohrwolf 140
rothschildi cf. Mustela
rufus cf. Canis
riippeli cf. Fennecus
Rytina 25
Rytina borealis 25
— cetacea 25
Rytiodinae 23
5
Sacalius 96
— aureus 96
sacana cf. Mustela
Schaeffia 99
seistanica cf. Lutra
Selenarctos 403, 404, 406, 414, 478
— ussuricus 478
severzovi cf. Meles
Speothos 62
— venaticus 63
sibirica cf. Mustela
sibiricus cf. Gulo, Meles, Ursus
signata cf. Mellivora
Simenia 62, 68, 99
— simensis 62
Simiae 9
Simocyoniae 62, 63, 383
Sirenia 10, 11, 15, 2/, 22, 54
seistanica cf. Lutra
Sivaonyx 854
Smilodon 50
Speothoninae 62
Speothos 58, 60, 62
— venaticus 59, 63
Spilogale 493, 498
spitzbergenensis cf. Alopex
Stellerus 25
— borealis 25
Stenoplesictinae 497
Stromeriella 817
Subungulata 22
syriaca cf. Vormela
syriacus cf. Ursus
szetchuensis cf. Martes
т
talassica cf. Mustela
Tamias sibiricus 521
tatarica cf. Mustela
Taxidea 493, 494, 498, 817, 818, 819
Taxodon 817
Taxus 816
Tayra barbara 616
teberdina cf. Mustela
temon cf. Mustela
Thalarctos 403, 404, 406, 455
— polaris 404
Thalassarctos 404
— jenaensis 455
— spitzbergenis 455
Thalassiarchus 404
Thos 96, 99
— vulgaris 96
thous cf. Lycalopex
tianschanensis cf. Meles
tiarata cf. Mustela
tuvinicus cf. Mustela
tobolica cf. Mustela
Tomarctos 268
Theria 10
998 Tarsioidea 9
transsylvanica cf. Mustela
Tremarctos 399, 403, 404, 406
— omatus 401, 403, 406
trettaui cf. Mustela
Trichechidae 15, 17, 23, 24
Trichechiformes 23, 25
Trichechus 15, 17, 18, 23, 24
— borealis 25
— inuguis 16, 20, 24
— manatus 20, 24, 25
— senegalensis 20, 24
tschiliensis cf. Canis
Tubulidentata 10, 54
turcmenicus cf. Vulpes
tundrarum cf. Canis
turovi cf. Mustela
U
ungava cf. Alopex
Unguiculata 10
ungavensis cf. Ursus
Ursarctos 404
Urocyon 59, 62
1549
1550
Urocyon cinereo-argentatus 63
Ursavus 402
Ursi campestres 419
Ursidae 54, 55, 56, 57, 60, 397, 401, 402,
911
Ursus 398, 401, 403, 404, 404, 406, 407
— albus 618
— americanus 399, 400, 404, 405, 406
— arctos 399, 404, 406, 407, 407, 408, 411,
412, 415, 422, 424, 428, 429, 430, 431,
431, 432, 433, 433, 434, 435, 436, 436,
437, 439, 451, 453, 459
— aureus 407
— baicalensis 408, 429
— beringiana 407, 436
— beringianus 435, 436
— brunneus 407, 436
— cadaverinus 407
— caucasicus 408, 431
— cavifrons 436
— collaris 407, 429, 431
— crowtheri 428, 437
— dinniki 431
— eogroenlandicus 467
— formicarius 407
— formosanus 485
— gedrosianus 485
— grandis 408
— griseus 407
— groenlandicus 469
— gulo 615, 618
— gyas 428, 436
— hypernefes 408
— indicus 804
— isabellinus 407, 434, 435, 439, 453
— japonicus 485
— jenaensis 467
— jessoensis 406
— kamtschatica 407
— kolymensis 408, 429
— labradorensis 469
— lagomyarius 408
— laniger 485
— lasiotus 404, 408, 436, 436
— lasistanicus 408, 433
— leuconyx 408, 434
— longirostris 407
— lotor 912, 913
— luscus 618
— mandchuricus 408
— marinus 455, 460, 467, 468
— maritimus 404, 406, 407, 4/11, 455, 455,
456, 458, 461, 465, 467, 468
— meles 816, 819
— melli 485
— meridionalis 407, 431, 431, 432, 433,
434, 459
— minor 407
— middendorffii 428, 436
— mupinensis 485
— myrmecophagus 407
— niger 407
— normalis 407, 408
— ornatus 404
— pamirensis 408, 434, 434
— persicus 408
— piscator 407, 435, 436, 437, 459
— pruinosus 404, 407, 434, 435, 437
— polaris 455, 467
— rossicus 408
— scandinavicus 408
— sibiricus 408
— smimovi 408, 431
— spitzbergenensis 467
— stenorostis 408
— syriacus 407, 431, 432, 433, 434, 435,
459
— thibetanus 404, 411, 478, 479, 481, 483,
484, 485, 485
Ursus tibetanus 399, 403, 404, 406, 407,
414, 437
— torquatus 478
— tyrannus 460
— ungavensis 469
— ussuricus 485
— yeniseensis 408, 429, 432, 433, 436,
437, 459
— yessoensis 404, 408
ussuricus cf. Ursus
ussuriensis cf. Nyctereutes
У
vancouverensis cf. Gulo
Vetularctos 406
Vishnuonyx 854
Viverra aterrima 884
— capensis 801, 804
— lutreola 636, 718
— ratel 801
Viverridae 54, 55, 497
viverrinus cf. Nyctereutes
Vormela 498, 501, 501, 781, 781, 782, 783
— alpherakyi 794
— chinensis 798
1551
Vormela euxina 784, 794 — flavescens 66, 319, 333, 334, 337, 340,
— intermedia 794 340, 360
— koshevnikovi 783, 794, 795, 796, 797 — fulva 327, 341
— negans 784, 796, 797 — griffithi 341
— obscura 784, 794 — harrimani 341
— ornata 784 — hoole 341
— pallidior 784, 795, 796 — Вий 320
— peregusna 782, 783, 783, 784, 784,785, | — шаша 341
788, 790, 793, 194, 795, 796, 797, 798 — ischnusae 341
— sarmatica 784, 794 — jacutensis 320, 339
— syriaca 784, 797 — japonica 341
999 — tedschenica 783, 794 — kalmykorum 273, 280
Vulgaris cf. Mustela — kamtschadensis 319, 339
Vulpes 62, 63, 68, 69, 99, 194, 196, 265, — kamtschatica 319, 339
265, 268, 277 — karagan 320
— abietorum 341 — kenaiensis 341
— aegyptiaca 341 — kiyomassai 320, 339
— alascenis 341 — krymea-montana 319
— alopex 319 — krymensis 338
— alpherakyi 319, 338, 338 — kurdistanica 319, 338
— alticola 320, 338 — lagopus 196
— anadyrensis 319, 339 — bengalensis 63
— anatolica 341 — leucopus 319
— arabica 341 — macroura 341
— atlantica 341 — melanotus 339
— bangsi 341 — montana 341
— barbara 341 — monticola 341
— bengalensis 268, 272, 327 — necator 341
— beringiana 339, 339, 341 — nigra 273, 281
— beringianus 196 — nigricans 269
— cana 63, 66, 266, 268, 269, 269, 269, — ochroxantha 320, 340
270, 271, 272, 327 — palestinae 341
— caragan 336, 339, 340 — pamirensis 320, 340
— cascadensis 341 — peculiosa 341
— caucasica 319, 338 — planicola 341
— chama 268 — pusilla 341
— cinerascens 319, 340 — pusillus 319
— communis 265 — regalis 341
— coloregriseo 319 — rubricosa 341
— corsac 63, 66, 266, 268, 273, 274, 275, — riippeli 268
276, 279, 280, 281, 315, 327 — schrenkii 320, 339, 341
— crucigera 319, 320, 337, 341 — scorodumovi 273, 281
— crymea-montana 338 — septentrionalis 319, 336
— crymensis 319 — sibirica 339
— daurica 320, 339, 339 — sibiricus 319
— deletrix 341 — silacea 341
— diluta 320, 337, 337, 339, 340 — splendens 319, 340, 340
— dolichocrania 320, 325, 336, 339 — splendidissima 320, 339, 341
— eckloni 281 — stepensis 319, 337, 338
— ferganensis 320, 333, 340 — tarimensis 319
— ferrilata 268, 272, 327 — tobolica 320, 339, 340
— ferrilatus 63, 281 — tschiliensis 319
1552
Vulpes turcmenica 273, 280, 315
— ussuriensis 319, 339
— velox 63
— vulgaris 319
— vulpes 56, 58, 63, 64, 66, 66, 266, 268,
318, 319, 320, 321, 326, 327, 329, 330,
333, 334, 336, 336, 337, 337, 337, 338,
339, 339, 340, 340, 341, 360
Vulpicanis 96
W
wilsoni cf. Mellivora
Y
yeniseensis cf. Martes, Ursus
yuenschanensis cf. Martes
Z
zerdus cf. Fennecus
zibellina cf. Martes
Zibellina 502
Zorilla libyca 783
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Other Books of Interest is Е eae ee
~MAMMALS OF THE SOVIET UNION
Volume II, Part 2 :
This volume, is part of a three-volume monograph, and is a continuation,
of Volume II, Part 1, which was devoted to sea cows and carnivores. It
contains species descriptions of terrestrial carnivores and detailed
information on their external morphology, skull, body measurements and
other data, affinities with other species, geographic distribution in the
historic past and today, geographic variation, practical significance, and
biology. Descriptions are presented for orders, families, and genera, and
keys given for their identification.
The book is richly illustrated with photographs, sketches, and colored
illustrations by the famous wild life painter,.A.N. Komarov, ue the
zoologist-artist, N.N. Kondakov. .
The results of original scientific studies are published here for the first
time, providing readers a vast wealth of material heretofore unknown. This
work is intended for teachers and students of faculties of biology and
geography in universities, as well as pedagogic, agricultural, and forest
institutes, similar organizations, workers engaged in game, fur, forest and
fish trades; those interested in the conservation of nature, and all persons
interested in zoology and nature study.
MAMMALS OF THE SOVIET UNION
Volume II, Part 3
This volume is fourth book of Mammals of the Soviet Union, representing the
third part of the second volume; it is devoted to descriptions of the orders
of Soviet aquatic mammals—pinnipeds (Pinnipedia) and, in part,
cetaceans (Cetacea), toothed whales (Odontoceti). In the sequence of
descriptions from the “higher” to the “lower” orders adopted in this series,
pinnipeds should have preceded carnivores, 1.е., should have appeared in
the second book. The grouping of the orders at a higher level is given after
G.G. Simpson (1945). The sequence of genera and species within the orders
has been retained as before, i.e., from the less specialized to the
more specialized.
The order of pinnipeds or seals (Pinnipedia) is described in this volume.
The cohort of ungulates and carnivores represented in soviet fauna by
orders of artiodactyls, and perissodactyls, sirenians, carnivores, and
pinnipeds thus came to an end and the cohort of whales (Mutica) com-
mences. The toothed whales are described in this volume.
While it has not always been possible to maintain a totally uniform
description of the genera and species as in the volumes already published,
in spite of every effort to do so, the sequence has been adhered to, with
some exceptions, in the case of Pinnipedia. But, it was impossible to use
same format in describing the toothed whales (Odontoceti). Only a brief
morphological description has been given for many species, which is more
or less adequate for identifying the species. General information on their
distribution and fragmentary biological data are also given. In some cases
the total absence of such information is indicated.
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