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Mustela sibirica (Carnivora: Mustelidae)

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Mustela sibirica Pallas, 1773, commonly known as the Siberian weasel, is a widely distributed Palearctic musteline with natural populations ranging from west of the Ural Mountains of Siberia to the Far East and south to Taiwan and the Himalayas. A key characteristic that distinguishes M. sibirica from most sympatric musteline species is the occurrence of a black mask on its face that surrounds the eyes, a white muzzle and chin, and the presence of a nearly completely monotone yellowish-brown coat. Although M. sibirica is hunted to make “kolinsky stable-hair” paintbrushes, populations remain stable and the species is currently listed as “Least Concern” by the International Union for Conservation and Nature and Natural Resources.
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109
MaMMalian SpecieS 50(966):109–118
Mustela sibirica (Carnivora: Mustelidae)
Chris J. Law www.mammalogy.org
Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, 130 McAllister Way, Santa Cruz, CA 95060,
USA; cjlaw@ucsc.edu
Abstract: Mustela sibirica Pallas, 1773, commonly known as the Siberian weasel, is a widely distributed Palearctic musteline with
natural populations ranging from west of the Ural Mountains of Siberia to the Far East and south to Taiwan and the Himalayas. A key
characteristic that distinguishes M. sibirica from most sympatric musteline species is the occurrence of a black mask on its face that
surrounds the eyes, a white muzzle and chin, and the presence of a nearly completely monotone yellowish-brown coat. Although
M. sibirica is hunted to make “kolinsky stable-hair” paintbrushes, populations remain stable and the species is currently listed as
“Least Concern” by the International Union for Conservation and Nature and Natural Resources.
Key words: kolinsky, kolonok, mustelid, musteline, Siberian weasel
Synonymy completed 19 February 2015
DOI: 10.1093/mspecies/sey013
Version of Record, first published online September 27, 2018, with fixed content and layout in compliance with Art. 8.1.3.2 ICZN
Nomenclatural statement.—A life science identifier (LSID) number was obtained for this publication: urn:lsid:zoobank.org:pub:
6625EDBE-8629-46FB-A3EB-29300D954CFF
Mustela sibirica Pallas, 1773
Siberian Weasel
Mustela sibirica: Pallas, 1773:701. Type locality: “Sibiriae
montanis, sylvis densissimis;” restricted to “Vorposten
Tigerazkoi, near Usstkomengorsk, W. Altai” (Oskemen,
Kazakhstan, 49.9833° N, 82.6167° E) by Pocock 1941. First
use of current name combination.
Viverra sibirica: Shaw, 1800:431. Name combination.
P[utorius] sibericus: Griffith, 1827:122. Name combination.
Mustela [Putorius] subhemachalana Hodgson, 1837:563. Type
locality “Nepal.”
M[ustela] canigula Hodgson, 1842:280. Type locality “Tibet.
[Mustela] humeralis Blyth, 1842:99. Type locality “Sikkim.
Mustela hodgsoni Gray, 1843:118. Type locality “India,
Himalaya.”
Mustela horsfieldii Gray, 1843:118. Type locality “Bhutan,
India.”
Vison sibirica: Gray, 1865:117. Name combination.
Putorius fontanierii Milne Edwards, 1871:205. Type locality
“la Chine;” description based on a specimen obtained by
M. Fontanier in China.
Putorius davidianus Milne Edwards, 1874:343. Type locality
“Kiang-si, [Moupin, Tibet].
Putorius moupinensis Milne Edwards, 1874:347. Type locality
“Moupin in Szechwan.
© The Author(s) 2018. Published by Oxford University Press on behalf of American Society of Mammalogists. All rights reserved.
For permissions, please e-mail: journals.permissions@oup.com
Fig. 1.—Adult Mustela sibirica from Longleat Safari Park, Britain.
Used with permission from photographer Clare Bambers.
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110 MAMMALIAN SPECIES 50(966)—Mustela sibirica
Putorius sibiricus miles Barrett-Hamilton, 1904:391. Type local-
ity “Dauria, Eastern Siberia.”
Putorius sibiricus noctis Barrett-Hamilton, 1904:391. Type
locality “San-yen-tze, China.”
Lutreola stegmanni Matschie, 1907:150. Type locality “Tsingtao,
Shantung.”
Lutreola quelpartis Thomas, 1908:53. Type locality “Island of
Quelpart, S. of Korea.
Lutreola major Hilzheimer, 1910:310. Type locality “Tibet.
Lutreola tafeli Hilzheimer, 1910:310. Type locality “Tibet.
Kolonokus sibiricus australis Satunin, 1911:266. Type locality
“Tyumen.”
M[ustela] manchurica Brass, 1911:262. Type locality
“Manchuria.”
Mustela [Lutreola] taivana Thomas, 1913:91. Type locality
“Formosa.
Kolonocus sibirica sibirica Satunin, 1914:124. Name
combination.
Mustela hamptoni Thomas, 1921:500. Type locality “Imaw
Bum.”
Kolonocus sibiricus coreanus Domaniewski, 1926:55. Type
locality “Seoul.”
Kolonocus sibiricus peninsulae Kishida, 1931:380. Type locality
unknown.
Mustela [Kolonocus] sibirica charbinensis Lowkashkin,
1934:49. Type locality “Manchuria.
Context and Content. Order Carnivora, family Mustelidae,
subfamily Mustelinae. Twelve subspecies are currently recog-
nized, 11 listed by Wozencraft (2005) and M. sibirica taivana
proposed by Suzuki et al. (2013). A revision of subspecies tax-
onomy, however, is needed as up to 22 subspecies have been
proposed (Larivière and Jennings 2009).
M. s. canigula Hodgson, 1842:280. See above.
M. s. charbinensis Lowkashkin, 1934:49. See above.
M. s. coreanus Domaniewski, 1926:55. See above; peninsulae
Kishida, 1931:380 is a synonym.
M. s. davidiana Milne Edwards, 1874:343. See above; noctis
Barrett-Hamilton, 1904:391 is a synonym.
M. s. fontanierii Milne Edwards, 1874:205. See above; steg-
manni Matschie, 1907:150 is a synonym.
M. s. hodgsoni Gray, 1843:118. See above.
M. s. manchurica Brass, 1911:262. See above.
M. s. moupinensis Milne Edwards, 1874:347. See above; hamp-
toni Thomas, 1921:500, major Hilzheimer, 1910:310, and
tafeli Hilzheimer, 1910:310 are synonyms.
M. s. quelpartis Thomas 1908:53. See above.
M. s. sibirica Pallas, 1773:701. See above; australis Satunin,
1911:280 miles Barrett-Hamilton, 1904:391 are synonyms.
M. s. subhemachalana Hodgson, 1837:563. See above; hume-
ralis Blyth, 1842:99 and horsfieldii Gray, 1843:118. are
synonyms.
M. s. taivana Thomas, 1913:91. See above.
NoMenclatural NoteS. Mustela sibirica has been previ-
ously placed in the genus Viverra (Shaw 1800), genus Putorius
(Griffith 1827), genus Vison (Gray 1865), genus Lutreola
(Matschie 1907), genus Kolonokus (Satunin 1911), and genus
Kolonocus (Satunin 1914). In addition, M. sibirica has also been
placed under the subgenus Lutreola (Youngman 1982) and later
in the subgenus Kolonokus (Abramov 1999). Other vernacular
names include the kolonok and kolinsky (Novikov 1962).
DIAGNOSIS
Mustela sibirica occurs sympatrically with a variety of
mustelids including ferret-badgers, martens, otters, and weasels
and stoats (mustelines). Mustelines like M. sibirica can be dis-
tinguished from many other mustelids by their small sizes and
elongated bodies. In its natural ranges in Asia, M. sibirica can
be distinguished from most sympatric mustelines—mountain
weasel M. altaica, ermine M. erminea, yellow-bellied weasel
M. kathiah, least weasel M. nivalis, and the introduced American
mink Neovison vison—by the presence of a black mask on its
face that surrounds its eyes, a white muzzle and chin, and a nearly
completely monotone yellowish-brown coat in winter (Fig. 1).
The sympatric Steppe polecat M. eversmanii also exhibits a dark
mask that surrounds its eyes but the mask extends farther across
its face toward the cheeks. In addition, M. eversmanii exhibits
a white band between the ears and eyes that crosses its head
from cheek to cheek (Heptner et al. 2001; Larivière and Jennings
2009). Other characteristics that distinguish M. sibirica from
M. eversmanii are body size (M. eversmanii can attain a body
mass twice that of M. sibirica) and coat color (M. eversmanii
exhibits a coat with a combination of yellowish-white and dark
brown color, whereas M. sibirica exhibits a nearly completely
monotone yellowish-brown coat in winter and a dark brown coat
in summer—Heptner et al. 2001; Larivière and Jennings 2009).
On the Japanese islands of Honshu, Shikoku, and Kyushu,
introduced populations of M. sibirica occur sympatrically with
the Japanese weasel M. itatsi. Characteristics that distinguish
M. sibirica from M. itatsi include body size (M. sibirica is larger
than M. itatsi), the ratio of tail (T) length to body (HB) length
(T/HB ratio is > 50% in M. sibirica, whereas the T/HB ratio is
< 40% in M. itatsi), and coat color (M. sibirica exhibits a lighter
brown coat than M. itatsi in winter—Masuda et al. 2012).
GENERAL CHARACTERS
Mustela sibirica is sexually dimorphic and males are
almost twice as heavy as females (Larivière and Jennings
2009). Body weight is 650–820 g for males and 360–430 g for
females (Hunter 2011). Body length is 28–39 cm for males and
25–30.5 cm for females, and tail length is 15.5–21 cm for males
and 13.3–16.4 cm for females (Hunter 2011).
Like other mustelines, M. sibirica has a long, slender body
with short limbs. The summer pelage is characterized by short,
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50(966)—Mustela sibirica MAMMALIAN SPECIES 111
coarse hair with a dark brown color almost completely covering
the entire body and tail; the winter pelage is denser and pale,
yellowish-brown in color (Heptner et al. 2001; Larivière and
Jennings 2009). The face exhibits a dark mask around in front of
the eyes with a white muzzle and chin (Hunter 2011). Females
have 4 pairs of mammae (Pocock 1941).
The skull is characterized as long and narrow (Heptner et al.
2001; Fig. 2). Mean skull measurements (mm, with ranges in
parenthesis) for adult male and female M. s. sibirica in Russia,
respectively, were: condylobasal length, 61.7 (58.0–63.5), 52.8
(49.8–56.3); zygomatic breadth, 32.2 (28.7–35.7), 27.8 (26.4–
29.6); interorbital width, 11.7 (11.7–13.2), 11.0 (10.5–12.2);
mastoid width, 27.5 (26.8–28.7), 24.3 (23.0–26.1—Heptner
et al. 2001). The skulls of male M. sibirica are 16.25% larger
than the skulls of females (Law and Mehta 2018).
For mustelids in general, the degree of sexual dimorphism
in body mass and length can be strongly impacted by the food
supply for a cohort during growth, and dimorphism in body size
often exceeds that for teeth and jaws (King and Powell 2007).
M. sibirica does exhibit sexual dimorphism in craniodental size
but little in shape (Sheng 1987; Abramov and Puzachenko 2009;
Suzuki et al. 2011). Discriminant analyses using 45 craniodental
linear measurements found the following characters contributed
to larger skull size in males compared to females: relatively wide
viscerocranium; wide postorbital constriction; a slender, long,
and high neurocranium; short and wide auditory bullae; short
carnassials; and a long and high mandible (Suzuki et al. 2011).
The degree of sexual size dimorphism varies across the species’
geographic range (Sheng 1987; Abramov and Puzachenko 2009;
Suzuki et al. 2013). Abramov and Puzachenko (2009) found that
the subspecies M. s. manchurica of the Far East displays a greater
degree of sexual size dimorphism than M. s. sibirica of western
and central Siberia. In China, populations occurring in the river
plains near the Yangtze and Huai rivers are generally larger and
exhibit greater sexual size dimorphism than conspecifics occur-
ring in forest habitats of the Changbai Mountains (Sheng 1987).
In addition, male and female individuals of insular populations
exhibit smaller skull sizes; the subspecies M. s. taivana in Taiwan
exhibits significantly smaller skulls compared to M. s. davidiana
of southeast China (Suzuki et al. 2013). Similarly, populations
of M. s. coreana in Tsushima Island are slightly smaller than
populations of conspecifics found in South Korea (Suzuki et al.
2013).
The baculum is weakly curved; the distal tip is flattened
and bent upwards, forming a slight hook (Heptner et al. 2001;
Baryshnikov et al. 2003). Mean measurements (mm, ranges in
parenthesis) for adult males and juvenile males, respectively,
were: length, 33.9 (32.0–35.8), 32.2 (30.2 –34.2); width of base,
2.15 (0.6–3.7), 1.6 (0.5–2.7); height of base, 3.65 (2.0–5.3), 2.25
(1.5–3.0—Novikov 1962).
DISTRIBUTION
Mustela sibirica is widely distributed across Palearctic Asia,
with natural populations ranging from the western base of the
Ural Mountains of Siberia to the Far East and south to Taiwan
and the Himalayas (Abramov et al. 2016; Fig. 3). M. s. sibirica
occurs in all of Siberia, ranging from the Kostroma Oblast to
63°N in the Ural Mountains and the upper reaches of the Pur
River and down south to the northern border of Kazakhstan and
the Altai Mountains (Bakeev 1971; Kassal 2013; Abramov et al.
2016). The range continues eastward through the Zeya Basin and
Mongolia and ends at the western parts of northeastern China
(Manchuria—Heptner et al. 2001).
Both M. s. charbinensis and M. s. manchurica occur in
northeastern China (Manchuria—Heptner et al. 2001); however,
the exact ranges are unknown and the validity of these sepa-
rate subspecies remains untested. M. s. coreana is endemic to
the Korean Peninsula and to Tsushima, Japan (Sasaki and Ono
1994). M. s. fontanierii occurs in the northern parts of Anhui,
eastern parts of Gansu, southern parts of Hebei, Henan, northern
parts of Hubei, northern parts of Jiangsu, southern parts of Nei
Fig. 2.—Dorsal, ventral, and lateral views of cranium and lateral view
of mandible of an adult female Mustela sibirica. Photograph taken at
the California Academy of Sciences (CAS-MAM 11668). Total skull
length is 5.3 cm.
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112 MAMMALIAN SPECIES 50(966)—Mustela sibirica
Mongol, Shaanxi, Shandong, Shanghai, and Shanxi (China—
Allen 1929; Smith et al. 2010). M. s. quelpartis is endemic to
Jeju Island (formerly Quelpart Island), Japan (Abramov 2005).
Two subspecies occur at the southeast edge of the species’
geographic range: M. s. davidiana occurs in southeast China
(Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan,
Jiangxi, Shaanxi, Sichuan, and Zhejiang) and M. s. taivana
is endemic to Taiwan (Smith et al. 2010; Suzuki et al. 2013).
M. s. moupinensis occurs in the Chinese provinces of Gansu,
Guizhou, western parts of Hubei, southeastern parts of Qinghai,
southern parts of Shaanxi, Sichuan, Yunnan (Ellerman and
Morrison-Scott 1951; Smith et al. 2010).
Three subspecies occur around the Himalayas: M. s. cani-
gula occurs in Tibet (Hodgson 1842; Heptner et al. 2001);
M. s. subhemachalana occurs in Nepal to Bhutan (Ellerman and
Morrison-Scott 1951); and M. s. hodgsoni occurs in Kashmir
and the Western Himalayas from Kam to Garhwal (Gray 1843;
Heptner et al. 2001).
Mustela sibirica was released from fur farms in Hyogo and
has since spread to the Japanese islands of Honshu, Shikoku, and
Kyushu (Miyashita 1963; Sasaki et al. 2014). M. s. sibirica was
also reintroduced in the Semenov District of Nizhny Novgorod
Oblast, Russia in 1937 and in the Dzhetyoguz District of Issyk
Kul Province, Kyrgyzstan in 1941 (Heptner et al. 2001; Long
2003). No fossils are known for M. sibirica.
FORM AND FUNCTION
The dental formula for Mustela sibirica is i 3/3, c 1/1, p 3/3,
m 1/2, total 34 (Larivière and Jennings 2009). Comparison of the
craniodental morphology using 32 linear measurements found
subtle shape differences between 5 populations of M. sibirica
(southeast China, Korea, Tsushima, Honshu, and Taiwan—
Suzuki et al. 2013). Skulls from insular populations tend to be
smaller than continental specimens (Suzuki et al. 2013).
In northern Russia, the spring molt occurs toward the end
of February or the beginning of March; the winter guard hairs
shed and the pelage is quickly replaced with summer guard hairs
(Novikov 1962). The autumn molt occurs at the end of August or
the beginning of September (Novikov 1962). The winter guard
hairs grow out simultaneously with the loss of summer guard
hairs, and the winter pelage is completely grown out by early
November (Novikov 1962). In Heilongjiang, China, autumn
molt begins around October–November (Hua et al. 2010).
Increased in hair densities (hairs/mm2) from summer to winter
coats in males and females, respectively, were: 91.82 to 219.33
and 73.83 to 182.35 on the head; 121.93 to 263.98 and 105.99
to 205.50 on the back; 73.89 to 175.12 and 65.91 to 151.26 on
the belly; and 80.38 to 183.59 and 73.21 to 180.63 on the tail
(Hua et al. 2010). Increase in hair lengths (mm) from summer to
Fig. 3.—Geographic distribution of Mustela sibirica. Map redrawn from Abramov et al. (2016). Subspecies are: 1) M. s. sibirica; 2) M. s. charbi-
nensis and M. s. manchurica; 3) M. s. coreana; 4) M. s. fontanierii; 5) M. s. quelpartis; 6) M. s. davidiana; 7) M. s. taivana; 8) M. s. moupinensis;
9) M. s. canigula; 10) M. s. subhemachalana; and 11) M. s. hodgsoni. 12) Indicates invasive populations.
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50(966)—Mustela sibirica MAMMALIAN SPECIES 113
winter coats in males and females, respectively, were: 11.50 to
17.84 and 11.48 to 14.27 on the head; 20.82 to 27.01 and 18.90
to 25.18 on the back; 15.97 to 19.82 and 14.06 to 18.33 on the
belly; and 30.36 to 42.61 and 23.59 to 41.63 on the tail (Hua
et al. 2010).
In the Longkou Forest Farm of Tonghe in the Xiaoxingan
Mountain, China, mean measurements (ranges in parenthesis)
of the winter guard hairs from the mid-backs of 15 adult males
and 15 adult females, respectively, were: hair length, 33.50 mm
(32.00–36.00 mm), 28.85 mm (25.50–31.50 mm); hair follicle
length, 0.37 mm (0.23–0.40 mm), 0.27 mm (0.20–0.34 mm);
hair diameter, 126.6 µm (108.4–152 µm), 79.41 µm (98.5–
147.8 µm); and hair root diameter, 26.8 µm (19.7–39.4 µm),
22.5 µm (19.7–29.6 µm—Zhang et al. 2008). Mean measure-
ments (ranges in parenthesis) of winter upper-hairs from the
hind-toes of 15 adult males and 15 adult females, respectively,
were: hair length, 11.32 mm (9.31–14.28 mm), 10.45 mm
(9.10–11.59 mm); hair follicle length, 0.91 mm (0.46–1.33 mm),
0.79 mm (0.11–1.21 mm); hair diameter, 107.7 µm (91.6–
119.2 µm), 101.0 µm (88.7–108.4 µm); and hair root diameter,
86.0 µm (68.0–109.3 µm), 71.9 µm (59.1–88.7 µm—Zhang et al.
2008).
The anal gland contains 9 sulfur-based volatiles: 2,2-dimeth-
ylthietane, (E)-2,4-dimethylthietane, (E)-2,3-dimethylthietane,
2-ethylthietane, (E)-2-ethyl-3-methylthietanes, (Z)-2-ethyl-
3-methylthietanes, 2-propylthietane, 3,3-dimethyl-1,2-dithi-
acyclopentane, and (Z)-3,4-dimethyl-1,2-dithiacyclopentane;
(E)-2,2-dimethylthietane is the most abundant (Zhang et al.
2002). Volatile abundance differs between the sexes: (E)-2,4-
dimethylthietane and (E)-2,3-dimethylthietane are significantly
more abundant in females than in males, whereas 3,3-dimethyl-
1,2-dithiacyclopentane is significantly more abundant in males
than in females (Zhang et al. 2002, 2003). 2-Ethylthietane only
occurs in females and is undetected in males (Zhang et al. 2002,
2003). Laboratory experiments reveal that rice-field rats Rattus
rattoides exhibit self-anointing behavior when presented filter
paper scented with anal-gland secretions of M. sibirica (Xu
et al. 1995).
ONTOGENY AND REPRODUCTION
Little is known about the reproduction of Mustela sibirica. In
Siberia, the breeding season occurs at the beginning of February
to the end of March (Heptner et al. 2001). Captive M. sibirica in
Novosibirsk, Russia, however, bred from April to August, with
peak breeding activity occurring in late April (Ternovsky 1977,
not seen, cited in Amstislavsky and Ternovskaya 2000:572). This
variation in timing may be due to differences in environmental
conditions, including those imposed by captivity. Copulation lasts
from 27 min to up to 2 h and 40 min (Ternovsky and Ternovskaya
1994). M. sibirica has the shortest gestation period (32–35 days;
mean 33.5 days) of all studied mustelids (Ternovsky 1977, not
seen, cited in Amstislavsky and Ternovskaya 2000:572). Liter
size ranges 2–12 kits (mean 6.2 kits) (Ternovsky 1977, not seen,
cited in Amstislavsky and Ternovskaya 2000:572). M. sibirica
does not exhibit delayed implantation (Mead 1989).
Young are born blind and almost naked with only sparse
white fur (Heptner et al. 2001). Young open their eyes for the
1st time by 28–30 days, and weaning ends at the end of the
2nd month (Heptner et al. 2001). Young born in April become
independent toward the end of summer, usually by August
(Novikov 1962).
ECOLOGY
Population characteristics.—The range of Mustela
sibirica is extensive across Palearctic Asia, with natural popu-
lations ranging from west of the Ural Mountains of Siberia to
the Far East and south to Taiwan and the Himalayas (Abramov
et al. 2016). Food abundance is hypothesized to determine the
population and distribution of M. sibirica, and Siberia and
northeast China are believed to contain the highest densities of
M. sibirica because of large densities of several rodent species
(Heptner et al. 2001).
Mustela sibirica is a common game species in western
Siberia, and records of population censuses are largely based on
fur trapping records (Bakeev 1971). Long-term records reveal
great annual and multi-annual fluctuations in population den-
sity. Increases in population densities were preceded by large
increases in rodent abundance (Bakeev 1971). The mean total
number of M. sibirica trapped during the early to mid-1900s are
25 in the Kostroma Oblast, 437 in the Orenburg Oblast, 422 in
the Republic of Tatarstan, 525 in the Sverdlovsk Oblast, and 86 in
the Tyumen Oblast (Bakeev 1971). Since the 1950s, the decline
in successful fur trappings suggested that population densities
in several regions decreased, which may be attributed to the
combination of deforestation and reduction in rodent abundance
(Bakeev 1971). Low fur prices also may reduce number of indi-
viduals trapped (Abramov et al. 2016). In the Sverdlovsk Oblast,
Russia, M. sibirica experienced a 39–71% decline in total popu-
lation abundance from 1987 to 2011 (Monakhov 2011a).
In Kyushu, Japan, population density of introduced
M. sibirica is 4–15 individuals per km (Sasaki et al. 2014). The
mean longevity for wild M. sibirica is calculated to be 2.1 years
(Miyagi and Shiraishi 1978).
Space use.Mustela sibirica is found in a wide variety
of habitats including dense primary and secondary deciduous,
coniferous, and mixed forests; woodlands; open grasslands;
and river valleys (Heptner et al. 2001; Abramov et al. 2016).
M. sibirica prefers regions near lakes and swamps covered
with bushes and fallen trees where small rodents are abundant
(Novikov 1962). M. sibirica is well documented at high eleva-
tion: 1,400–1,700 m in the secondary forests of Guandaushi
Forest, Taiwan (Wu 1999); 2,700–3,700 m in the primary for-
ests of the Tawu Mountains, Taiwan (Chiang et al. 2012); >
3,000 m in Nepal (Ghimirey et al. 2014); 1,500–4,800 m in
Bhutan (Abramov et al. 2016); and up to 5,000 m in China
(Abramov et al. 2016).
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114 MAMMALIAN SPECIES 50(966)—Mustela sibirica
Mustela sibirica uses fallen logs, empty stumps, and brush-
wood piles as shelters and nests (Heptner et al. 2001). Individuals
also inhabit the burrows of their prey, such as voles, mice, and
pikas (Heptner et al. 2001). Near Lake Baikal in Russia, bur-
rows ranged from 0.6 to 4.2 m in length and from 0.2 to 1.3 m
in depth, and the nesting chamber located in the middle of the
burrow was lined with feathers or fur from prey (Fetisoff 1936).
Each individual usually has 1 primary burrow as well as many
secondary refuges across its range, which may extend for several
kilometers (Fetisoff 1936).
Diet.Mustela sibirica exhibits a mesocarnivorous
(50–70% vertebrate prey) to hypercarnivorous (> 70% verte-
brate prey) diet that is largely dependent on the habitat and
location. Small voles, mice, and pikas constitute the basic
diet of M. sibirica in most locations (Fetisoff 1936; Novikov
1962; Heptner et al. 2001). Larger sized rodents such as chip-
munks, invasive muskrats, and other squirrels are also preyed
upon (Heptner et al. 2001). Birds, amphibians, fish, eggs, ber-
ries, and nuts are consumed when rodents are not available
(Novikov 1962).
On the Tsushima Islands of Japan, scat analyses (n = 218)
reveal that M. sibirica exhibits a mesocarnivorous diet: small
mammals (35%, average percentage of relative occurrence),
insects (20%), berries and seeds (13%), birds (10%), other plant
material (10%), earthworms (7%), and amphibians and reptiles
(5%—Tatara and Doi 1994). The Shannon–Weaver’s diversity
index (H) of the prey items was 1.869 (Tatara and Doi 1994).
M. sibirica exhibits seasonal differences in diets. Caterpillars and
beetles are common in spring and summer (24.4–31.8%), and
earthworms (19.8%) are consumed during the autumn (Tatara
and Doi 1994). During winter, tetrapods comprise nearly 80%
of the diet, including an increase in bird consumption (24.5%).
Small mammals remain the most common prey throughout the
year (22.6–48.9%), and a large portion consists of mainly house
mice, Mus musculus, and wood mice, the large Japanese field
mouse Apodemus speciosus and the small Japanese field mouse
Apodemus argenteus (Tatara and Doi 1994). Surprisingly, plant
materials also occur throughout the year (12.8–28.6%—Tatara
and Doi 1994).
Scat (n = 115) from the grasslands of Aoshima, Japan, also
reveal a mesocarnivorous diet; insects (68.7%, average percent-
age of absolute occurrence), mammals (48.7%), amphibians
(13.0%), fish (12.2%), and reptiles (9.6%) are the dominate prey
items (Sasaki and Ono 1994).
In the Guandaushi Forest of Taiwan, scat analyses (n = 157)
reveal that arthropods (43.6%, average percentage of relative
occurrence), small mammals (26.0%), and earthworms (17.6%)
are the dominant prey items in this region (Wu 1999). The
Chinese white-toothed shrew Crocidura kurodai and the lesser
Taiwanese shrew Chodsigoa sodalis are the most important
mammalian prey, occurring in one-third of all analyzed scats
(Wu 1999). On the other hand, in high elevation alpine grass-
lands in Taiwan, M. sibirica exhibits a hypercarnivorous diet
where small mammals, particular rodents including the Oldfield
white-bellied rat Niviventer culturatus, the Taiwan field mouse
Apodemus semotus, Kikuchi’s field vole Microtus kikuchii, and
Père David’s vole Eothenomys melanogaster (92.0%, average
percentage of relative occurrence), are the most dominant prey
(Ma 1990).
Diseases and parasites.—In Hokkaido, Japan, intestinal
parasitic worms found in Mustela sibirica include 3 nema-
todes Capillaria putorii, Strongyloides, and Spiruidea (larva);
1 trematode Echinostoma hortense; and 1 acanthocephalan
Centrorhynchus elongatus (juvenile) (Kamiya and Ishigaki
1972). In addition, the nematode Filaroides martis was found in
the lungs (Kamiya and Ishigaki 1972). In the Tohoku region, the
intestinal fluke E. hortense was found in 2 M. sibirica individuals
(Sato et al. 1999). Lung fluke infections caused by Paragonimus
miyazakii and P. ohirai were found in animals from the Miyazaki
Prefecture, Japan (Ashizawa et al. 1980). Other parasites found
in Japanese M. sibirica populations include the trematodes
Clonorchis sinensis, Echinostoma trigonocephala, Heterophyes
heterophyes, Isthmiophora melis, and Paragonimus wester-
mani; the tapeworms Sparganum mansoni and Dipylidium cani-
num; the nematode Gnathostoma spinigerum; the roundworm
Dioctophyme renale and Trichinella; and the acanthocephalan
Centrorhynchus itatsinis (Yoshida et al. 1932).
Intestinal parasitic worms found in M. sibirica in Taiwan
include 7 nematodes Filaroides (94.4%, frequency of occur-
rence from 16 individuals), Ancylostoma (77.4%), Uncinaria
(35.5%), Trichuris species 1 (35.5%), Trichuris species 2
(19.3%), Capillaria (6.5%), and Physaloptera (3.2%); 1
trematode Platynosomum (74.1%); and 1 acanthocephalan
Macracanthorhynchus (10%—Chen 2003). Two species of ticks
were also observed: Ixodes ovatus and Haemaphysalis (Chen
2003).
In Hoengseonggun, South Korea, the tapeworm Spirometra
erinaceieuropaei was found in 1 M. sibirica individual (Lee
et al. 2013). The nematode Gnathostoma nipponicum has also
been found in populations in Jejudo, South Korea (Woo et al.
2011). Some South Korean populations of M. sibirica have also
been identified to carry vector-borne diseases through infections
from Ehrlichia and Anaplasma species (Chae et al. 2003).
Parasites found in M. sibirica in Russia include mites
Ixodes persulcatus and Dermacentor caina and the nematodes
Agamospirura, Filaroides orientalis, Scriabingulus nasicola
(Romanov 1960; Kontrimavichus and Kazakov 1966; Heptner
et al. 2001).
The 1st reported cases of neoplasia in M. sibirica occurred in
2 nonwild individuals in Lower Saxony, Germany game reserve
(Zöller et al. 2008). The male specimen exhibited an intersti-
tial cell tumor in the right testicle, and subsequent necropsy
revealed tumor lesions within the abdominal cavity and spleen
(Zöller et al. 2008). The female specimen exhibited a fibro-
sarcoma on the upper left hind limb; the tumor had developed
multiple times after removal of the original tumor (Zöller et al.
2008). M. sibirica can also be infected by canine distemper virus
(Kameo et al. 2012).
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50(966)—Mustela sibirica MAMMALIAN SPECIES 115
Interspecific interactions.Mustela sibirica occurs sym-
patrically with other carnivorans including felids, canids, and
other mustelids such as martens, ferret-badgers, weasels, and
polecats (Shaposhnikov 1956; Novikov 1962; Bakeev 1971;
Tatara and Doi 1994; Wu 1999; Chiang et al. 2012). Spatial,
dietary, and temporal variation in resource use have been sug-
gested to limit competition among these carnivores, but no
study to date has truly investigated interspecific interactions
between M. sibirica and other carnivorans
Mustela sibirica exhibits great dietary overlap with the
yellow-throated marten Martes flavigula chrysospila in Tawu
Mountain Nature Reserve, Taiwan, suggesting interspecific
competition for food (Chiang et al. 2012). Despite the similar
diets, yellow-throated martens exhibit almost exclusively diurnal
activity patterns, whereas M. sibirica is almost exclusively noc-
turnal, thus suggesting that the 2 mustelids limit competition by
avoiding each other temporally (Chiang et al. 2012).
Similarly, although sympatric M. sibirica and Chinese fer-
ret-badgers Melogale moschata have substantial dietary overlap
in the Guandaushi Forests of Taiwan, the relative abundance of
prey items for each differed significantly (Wu 1999). In addition,
M. sibirica occur at higher elevations (1,400–1,700 m) charac-
terized by secondary forest and flat terrain, whereas Chinese
ferret-badgers occur at lower elevations (850–1,400 m) charac-
terized by primary forests (Wu 1999).
In the Tsushima Islands of Japan, M. sibirica occurs sympat-
rically with 2 other species of carnivores: the Tsushima leopard
cat Felis bengalensis euptilura and the Tsushima marten Martes
melampus tsuensis (Tatara and Doi 1994). Scat analyses reveal
that the 3 carnivores do not compete for food: martens are the
most hypocarnivorous and consumed mainly fruits and berries,
whereas leopard cats are the most hypercarnivorous and con-
sumed small mammals and birds. M. sibirica exhibits an inter-
mediate, mesocarnivorous diet (Tatara and Doi 1994).
Mustela sibirica co-occurs with the sable Martes zibel-
lina in the taiga forests of the Altai, the Far East, and eastern
Siberia (Bakeev 1971). Sables generally eat small mammals,
birds, and vegetation matter such as nuts and berries (Monakhov
2011b). However, during periods of poor vegetation growth,
sables directly compete with M. sibirica for small rodents
(Shaposhnikov 1956). In this situation sables display agnostic
behaviors toward M. sibirica and often drive them into open
habitats which results in decreased populations of weasels
(Shaposhnikov 1956; Bakeev 1971). Fur of M. sibirica is occa-
sionally found in sable excrement (Shaposhnikov 1956).
Mustela sibirica is sympatric to 3 other mustelines in the
Baraba steppe of Western Siberia: M. erminea, M. nivalis, and
M. eversmanii (Abramov and Puzachenko 2012). Gut-content
analyses found that rodents comprised of 100% of each of these
4 mustelines, with great overlap in the consumption of smaller
rodent species such as mice and voles (Ternovsky and Danilov
1965). However, analyses on cranial traits suggest that these 4
mustelines occupy different regions of cranial morphospace and
thus may utilize different resources (Abramov and Puzachenko
2012). A more comprehensive study is needed to understand
patterns of resource partitioning between these 4 Mustela spe-
cies. Natural hybridization between M. sibirica and M. eversma-
nii is observed, resulting in hybrids known as “giant kolonoks”
that are much larger than typical M. sibirica individuals (Heptner
et al. 2001).
Invasive populations of M. sibirica in Japan are now sym-
patric with some populations of M. itatsi, and some researchers
have postulated that the 2 weasels compete for resources (Sasaki
et al. 2014). No comprehensive study has tested this hypothe-
sis. Recent distribution studies suggest that M. itatsi occurs
in grasslands and plantations and avoids urban areas, whereas
M. sibirica is more abundant in locations with greater human
activity (Sasaki et al. 2014). Sasaki et al. (2014) speculates that
the presence of M. itatsi prevents range expansion of M. sibirica.
M. sibirica primarily occurs in western Japan, and although the
distribution is slowly expanding eastward, the range cannot
expand past the Aichi Prefecture where M. itatsi is dominant
(Sasaki et al. 2014). Known predators of M. sibirica include
foxes and large falconiformes (Novikov 1962).
HUSBANDRY
Mustela sibirica is rarely held in captivity because of the
difficulty in raising them. M. sibirica is currently found in
only a few zoos such as the Longleat Safari Park in Britain,
the Poznan Zoo in Poland, and the Dresden Zoo in Germany.
The Experimental Research Station in Novosibirsk, Russia, is
the most successful in breeding M. sibirica (Ternovsky and
Ternovskaya 1994). Captive breeding can result in interspecies
hybrids between M. sibirica with M. eversmanii, the European
mink M. lutreola, and the European polecat M. putorius
(Ternovsky and Ternovskaya 1994). However, whether these
hybrids are reproductively viable is unknown. Individuals of
M. s. coreana from South Korea were kept in Japanese fur farms
in the late 1920s to early 1930s (Long 2003; Sasaki 2009).
M. sibirica are kept in Chinese fur farms (European Society of
Dog and Animal Welfare 2015).
Mustela sibirica and kohhosiks (hybrids between M. sibirica
and M. eversmanii) are kept as pets in Russia (Russian Ferret
Society 2007). The oldest known captive individual lived for
8 years and 10 months (Jones 1982).
BEHAVIOR
The behavior of Mustela sibirica is not well studied com-
pared to other Palearctic mustelines. M. sibirica is typically cre-
puscular or nocturnal (Heptner et al. 2001; Chiang et al. 2012).
Nocturnal activities are believed to limit resource competition
with other carnivorans such as yellow-throated martens (Chiang
et al. 2012). M. sibirica is solitary with the exception of females
raising young (Nowak 2005). Males do not assist in raising kits
(Novikov 1962).
Like other mustelines, M. sibirica uses anal glands for scent
communication, marking territory, and defense (Pocock 1941).
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116 MAMMALIAN SPECIES 50(966)—Mustela sibirica
The presence of sex-specific compounds in the anal glands sug-
gests that chemical secretion could be used to code for informa-
tion between males and females (Zhang et al. 2003). Individuals
caught in traps exhibit ear-piercing screams and anal-gland
secretion (Pocock 1941).
Both daily movements and seasonal migrations are depend-
ent on fluctuations of prey population (Heptner et al. 2001).
M. sibirica can move up to 8 km in a single night (Nowak 2005).
M. sibirica is reportedly a good swimmer and climber and is
able to pursue water voles in lakes and chase squirrels in trees
(Novikov 1962).
GENETICS
Mustela sibirica has a diploid number (2n) of 38 chromo-
somes and a fundamental number (FN) of 58 (Kurose et al.
2000). The karyotype consists of 7 pairs of metacentrics, 4 pairs
of submetacentrics, 7 pairs of acrocentrics, and 2 sex chromo-
somes (Kurose et al. 2000).
Mustela itatsi was once classified as a subspecies of
M. sibirica. Mitochondrial DNA data have since validated
the separation of the 2 Mustela species with divergence times
approximately 1.70–2.40 million years ago (Masuda and Yoshida
1994). More recent phylogenetic analyses using mitochondrial
and nuclear DNA demonstrate that M. sibirica and M. itatsi are
distinct species (Sato et al. 2012). M. sibirica is sister to a clade
comprised of M. lutreola, M. nigripes, M. putorius, and M. ever-
smanii (Koepfli et al. 2008; Sato et al. 2012; Law et al. 2018).
Genetic analyses using mitochondria DNA (cytochrome-
b gene) from 5 native populations—Transbaikalia (Russia),
Ural Mountains (Russia), Taiwan, South Korea, and Tsushima
Island (Japan)—as well as non-native populations on mainland
Japan indicated 4 distinct lineages: 1) a Korean lineage includ-
ing the introduced Japanese population, 2) a Tsushima lineage,
3) a Russian lineage, and 4) a Taiwanese lineage (Masuda et al.
2012). A separate study also using the cytochrome-b gene found
M. s. coreanus from the Korean Peninsula and M. s. quelpar-
tis from Jejudo are not distinctly different, suggesting that these
populations may not be 2 distinct subspecies (Koh et al. 2012).
Mustela sibirica is sympatric to M. eversmanii in the Baraba
steppe of western Siberia, and natural hybridization occurs
between these 2 species (Heptner et al. 2001). In addition, captive
breeding can result in interspecies hybrids between M. sibirica
with M. eversmanni, M. lutreola, and M. putorius (Ternovsky
and Ternovskaya 1994).
CONSERVATION
Mustela sibirica is listed as “Least Concern” by the
International Union for Conservation of Nature and Natural
Resources since 2008 (Abramov et al. 2016). Populations
of M. sibirica are protected under the Appendix III of the
Convention on International Trade in Endangered Species of
Wild Fauna and Flora (Convention for the International Trade of
Endangered Species of Wild Fauna and Flora 2015) in India, the
Tibet wildlife protection list, and the species is listed as “Near
Threatened” under the China Red List (Abramov et al. 2016).
The species’ wide distribution leads conservationists to presume
a large population with stable population sizes (Abramov et al.
2016), but no demographic censuses have been undertaken.
Currently, there are no major threats to M. sibirica (Abramov
et al. 2016). Historically, M. sibirica was considered a valua-
ble furbearer in Siberia and China and used to make “kolinsky
stable-hair” paintbrushes as well as ink brushes (Heptner et al.
2001). However, hunting levels are currently low because of low
commercial value of pelts (Abramov et al. 2016). In addition,
the placement of M. sibirica under Appendix III of CITES has
placed restrictions on the importation of kolinsky brushes to
some countries such as the United States (Shaw 2014).
ACKNOWLEDGMENTS
I am grateful to C. Bambers for providing the photograph
of Mustela sibirica as well as to M. Flannery and L. Wilkinson
for allowing me to take photographs of the skull specimen at
the California Academy of Sciences. I thank M. Hamilton,
D. Zegers, and 2 anonymous reviewers for valuable comments
on an earlier draft of this species account. Financial support was
provided by an UCSC Department of Ecology and Evolutionary
Biology Graduate Research Award and a National Science
Foundation Graduate Research Fellowship.
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Associate Editor was david a. zegerS. Editor was Meredith
j. haMilton.
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... The Siberian weasel, Mustela sibirica (Pallas, 1773), is widely distributed from Russia to Japan (Sasaki, 2015) and currently classified into 12 subspecies based on morphological characters (Law, 2018). Abramov et al. (2018) quantified morphometric variation in cranial characters of M. sibirica across its entire distributional range, except for Taiwan, and delineated three morphological groups: G1 (in Nepal and India), G2a (mainly in northern and north-western Asia) and G2b (mainly in north-eastern and south-eastern Asia). ...
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