Animal Diversity Web

Ge­o­graphic Range

Black-capped mar­mots (Mar­mota camtschat­ica) are Palearc­tic, or Eurasian mam­mals that have a patchy dis­tri­b­u­tion through­out north­ern and east­ern Siberia, or the east­ern por­tion of the Russ­ian Fed­er­a­tion. There are cur­rently three rec­og­nized sub-species of black-capped mar­mots, and each sub-species is ge­o­graph­i­cally iso­lated (or oc­cu­pies a dif­fer­ent ge­o­graphic lo­ca­tion). (Boyesko­rov, et al., 1994; Bran­dler, et al., 2010; Er­ba­jeva and Alex­eeva, 2009; Se­menov, et al., 2001a; Se­menov, et al., 2001b)

The first sub­species of black-capped mar­mots, Kam­chatka mar­mots (M. c. camtschat­ica) in­habit the Kam­chatka, Mil’kovski area along the val­ley of the Yurtinaya River. The sec­ond sub­species, Bar­guzin mar­mots (M. c. dop­pel­may­eri) oc­cupy a por­tion of Bury­a­tia in the Sever­obaikal’sk area, or more specif­i­cally the north-east­ern por­tion of Pre­baikalia (Baikal Moun­tains) as well as the north­ern por­tion of Trans­baikalia (Bar­guzin Moun­tain Range). Lastly, the third sub­species, Yakut­ian mar­mots (M. c. bungei) are found along the east­ern side of the Lena River in the Kha­raulakhskii Moun­tain Range in Yaku­tia, Rus­sia. Yakut­ian mar­mots oc­cupy one of the most north­ern parts of Rus­sia, in­hab­it­ing the north-east­ern part of Yaku­tia. The range of Yakut­ian mar­mots ex­tend from the delta of the Lena River, or along the Kha­raulah ridge, south along the Mom­sky, Chersy, and Verk­hoy­an­sky Moun­tain ridges as well as along the lower part of the Yana River. (Boyesko­rov, et al., 1994; Bran­dler, et al., 2010; Er­ba­jeva and Alex­eeva, 2009; Se­menov, et al., 2001b)

• Biogeographic Regions
• palearctic
  • native

• Other Geographic Terms
• holarctic

Habi­tat

Black-capped mar­mots in­habit high el­e­va­tion alpine and sub-alpine re­gions. Yakut­ian mar­mots oc­cupy moun­tain slopes 20 to 1500 m above sea level. Black-capped mar­mots in Yaku­tia is typ­i­cally ob­served at al­ti­tudes of 1200 to 2000 m. Kam­chatka mar­mots typ­i­cally in­habit areas 600 to 1500 m above sea level. (Boyesko­rov, et al., 1994; Filonov, 1961; Le Berre and Ra­mousse, 1994; Mosolov and Tokarsky, 1994; Se­menov, et al., 2001b; Tokarsky and Va­lent­sev, 1994)

Habi­tat sites ap­pear to be se­lected based on al­ti­tude, plant com­po­si­tion, and sun ex­po­sure. Black-capped mar­mots pre­fer bare moun­tain slopes that are ex­posed to the max­i­mal amounts of sun­light, which equates to south or south-west fac­ing slopes. Black-capped mar­mots are often found above the tree­line of dwarf pine and alder. Grass­lands, steppes, and mixed rock/grass areas seem to be pre­ferred over forested areas and other closed en­vi­ron­ments. (Boyesko­rov, et al., 1994; Le Berre and Ra­mousse, 1994; Mosolov and Tokarsky, 1994; Se­menov, et al., 2001a; Se­menov, et al., 2001b; Tokarsky and Va­lent­sev, 1994; Vino­gradov and Ar­girop­ulo, 1968)

Sites in­hab­ited typ­i­cally have dry, well-drained, soft or fine soils. The soils may con­tain silt, but may also have water and glacial de­posits that in­clude large boul­ders, bro­ken rock, and finer de­posits. Win­ter bur­rows are often cre­ated in clay soils. The up­land, alpine tree­less areas of east­ern Siberia and Kam­chatka are un­der­lain by per­mafrost and cov­ered with rocky soil and a few grasses. The black-capped mar­mots on Kam­chatka oc­cupy sites with vol­canic de­posits in close prox­im­ity to an ocean, for the moun­tain ranges are right next to the Pa­cific Ocean. Typ­i­cally, these areas are cov­ered with large loose rocks in­ter­spersed with small alpine mead­ows and shrubs, dwarf birch, and soli­tary Japan­ese stone pine (Pinus pumila). (Barash, 1989; Mosolov and Tokarsky, 1994; Se­menov, et al., 2000; Vino­gradov and Ar­girop­ulo, 1968; Zim­ina and Gerasi­mov, 1973)

Black-capped mar­mots are a semi-fos­so­r­ial species. Per­mafrost and rock pre­vents bur­row­ing to ex­ten­sive depths. Bur­rows may only reach depths of 0.25 to 0.6 m below the sur­face, which equates to the depth that the ground thaws dur­ing the sum­mer. How­ever, some moun­tain slopes have thicker soils and thaw to a depth of 1 m, so bur­rows may reach a depth of 1 m. Fur­ther­more, mar­mot en­vi­ron­ments ap­pear dif­fer­ent from the sur­round­ing tun­dra be­cause bur­row­ing and for­ag­ing ac­tiv­i­ties of black-capped mar­mots alter the veg­e­ta­tion com­mu­nity. (Boyesko­rov, et al., 1994; Kapitonov, 1978; Le Berre and Ra­mousse, 1994; Mosolov and Tokarsky, 1994; Se­menov, et al., 2001a; Se­menov, et al., 2001b; Zim­ina and Gerasi­mov, 1973)

Am­bi­ent tem­per­a­tures are usu­ally very low in en­vi­ron­ments in­hab­ited by black-capped mar­mots. Am­bi­ent tem­per­a­tures along the Lena River in Yaku­tia typ­i­cally range from -34°C in the win­ter to an av­er­age of 12.8°C at the end of July. Sum­mer tem­per­a­tures across the black-capped mar­mot range may reach 25°C. In gen­eral, win­ters are long, have lit­tle snow, se­vere frost, and tem­per­a­tures that may fall as low as -70°C. (Boyesko­rov, et al., 1994; Se­menov, et al., 2001a; Se­menov, et al., 2001b; Se­menov, et al., 2000; Zim­ina and Gerasi­mov, 1973)

• Habitat Regions
• polar
• terrestrial

• Terrestrial Biomes
• tundra
• taiga
• mountains

• Range elevation

  20 to 2000 m

  65.62 to 6561.68 ft

• Range depth

  1 (high) m

  3.28 (high) ft

• Average depth

  0.25-0.6 m

  ft

Phys­i­cal De­scrip­tion

The top of black-capped mar­mots heads are black from the tip of the nose to be­hind the ears. The sides of their heads are black to about the level of the eyes, and then sandy yel­low mixed with gray below the eyes. A black line car­ries down the back of the neck to the shoul­ders. Their lips are black and throats are or­ange. Their ears are or­ange to sand col­ored. Their dor­sal guard hairs have three dis­tinct bands of color; dark at the top and bot­tom and light in the mid­dle. Guard hairs are typ­i­cally 48 mm long. Their dor­sal sur­face is lighter near the head, be­com­ing darker pos­te­ri­orly to­wards the tip of the tail. The dor­sal un­der­fur is soft, dark, and light-tipped. On the ven­tral sur­face there is no un­der­fur and guard hairs are a yel­low­ish-brown to brown­ish-red, again with a darker colour pre­sent at the base of the hair and po­ten­tially at the tip. (Hoff­mann, et al., 1979; Step­pan, et al., 1999; Vino­gradov and Ar­girop­ulo, 1968)

Bar­guzin mar­mots, at the south­ern por­tion of their range, tend to dis­play a browner cap and browner tips on the dor­sal guard hairs. The mid­dle por­tion of the dor­sal guard hairs is a light to dark yel­low­ish-beige (buff). The ven­tral guard hairs are brown to cin­na­mon in color. North­ern sub­species of black-capped mar­mots, such as Yakut­ian mar­mots, dis­play a darker pelage. The mid­dle of the guard hair is ivory to white-yel­low in color, and the ven­tral guard hairs are brown or a cin­na­mon to red­dish-brown. Kam­chatka mar­mots may dis­play yel­low-beige or ivory in the mid­dle of its dor­sal guard hairs. Ven­tral guard hairs for Kam­chatka mar­mots are shades of red-, or­ange-, or yel­low-brown in color. (Hoff­mann, et al., 1979)

As black-capped mar­mots get older, their color fades. Ju­ve­nile mar­mots go through three pelage stages. First, ju­ve­niles have a soft, dense un­der­fur with a brown­ish-black to black cap and sim­i­lar-col­ored dor­sal guard hairs. The ini­tial guard hairs are shed, re­sult­ing in the sec­ond pelage stage. Dur­ing the third pelage stage, ju­ve­niles grow a pelage that more closely re­sem­bles the adults of its species. (Hoff­mann, et al., 1979)

The pelage of Alaska mar­mots is very sim­i­lar to that of black-capped mar­mots with the same black cap and dor­sal guard hairs. How­ever, the ven­tral sur­face of Alaska mar­mots has a more gray ap­pear­ance, be­cause of the pat­tern cre­ated by the ven­tral guard hairs with their dark tips and bases and light cen­ter. Hoary mar­mots have white fa­cial mark­ings and coarser dor­sal guard hairs com­pared to black-capped mar­mots. Black-capped mar­mots moult around early to mid-sum­mer, with hair loss be­gin­ning on their rump and pro­gress­ing an­te­ri­orly and ven­trally. While most of the fur is shed and re­placed every year, the fur on the rump to the end of the tail may re­main, and in fact may not be shed for an ad­di­tional year or more. (Hoff­mann, et al., 1979; Le Berre and Ra­mousse, 1994; Step­pan, et al., 1999)

Mar­mots have six pads on the soles of their hind­feet, but the shape of these pads dif­fers for each species. The pos­te­rior pair of foot pads in black-capped mar­mot is elon­gated com­pared to Hoary mar­mots, which have a pos­te­rior pair of foot­pads that are round. Alaska mar­mots have foot pads that re­sem­ble black-capped mar­mot. The length of the hind­foot for black-capped mar­mot is 73 to 85 mm. Feet are pen­tadactyl and dig­its have large claws that are used for dig­ging. (Hoff­mann, et al., 1979; Vino­gradov and Ar­girop­ulo, 1968)

Black-capped mar­mots may have five or six pairs of mam­mae. Those mar­mots that only have five pairs of mam­mae often also have one un­paired teat. Black-capped mar­mots are sex­u­ally di­mor­phic with males larger than fe­males. The av­er­age head-body length of male and fe­male black-capped mar­mots is 473.3 mm and 458.4 mm re­spec­tively. (Car­dini, 2004; Hoff­mann, et al., 1979; Vino­gradov and Ar­girop­ulo, 1968)

Mar­mot size, both mass and body length, varies across the ge­o­graphic range with smaller black-capped mar­mots noted in the south­ern part of the range and larger mar­mots noted in the north­ern part of the range. The head-body length of a male and fe­male Bar­guzin mar­mots were 470 mm and 440 mm re­spec­tively. Bar­guzin mar­mot males have a tail length of 150 mm while the fe­males have a tail length of 140 mm. Kam­chatka mar­mot males and fe­males have av­er­aged head-body lengths of 508.1 mm and 496.3 mm, re­spec­tively; av­er­age lengths in­clud­ing tails are 162.4 mm and 153.4 mm, re­spec­tively. Yakut­ian mar­mot males av­er­aged a head-body length length of 460.0 mm and 133.3 mm in­clud­ing tails. Fe­male mem­bers of this sub­species av­er­aged a head-body length of 438.5 mm and a tail length of 124.6 mm. (Hoff­mann, et al., 1979; Vino­gradov and Ar­girop­ulo, 1968)

The mass of black-capped mar­mots ranges from 2 to 7.5 kg, with greater masses noted just be­fore hi­ber­na­tion and lower masses noted after hi­ber­na­tion. Adult Yakut­ian mar­mots weigh ap­prox­i­mately 2 to 4 kg. Bar­guzin mar­mots weigh around 3 kg, and Kam­chatka mar­mots have an av­er­age mass of about 4.5 kg. ("Squir­rels", 2006; Bibikov, 1994; Boyesko­rov, et al., 1994; Hoff­mann, et al., 1979; Mosolov and Tokarsky, 1994; Nowak, 1999; Se­menov, et al., 2001a)

Skull lengths range from 78 to 99 mm in black-capped mar­mots. The an­gu­lar process of black-capped mar­mots is not greatly elon­gated, and is not much longer than the ar­tic­u­lar process. Other dis­tin­guish­ing skull fea­tures of black-capped mar­mots in­clude: a more no­tice­able mandibu­lar sym­ph­ysis; an elon­gated ven­tral half of the in­cisor socket in the jaw; the upper por­tion of the in­cisor socket shifted slightly back from the front; and each coro­noid process an­gles to­wards the back of the skull. Com­pared to hoary mar­mots, black-capped mar­mot have a longer ros­tra, a longer au­di­tory bulla, zy­go­matic arches that branch quickly from the max­illa as they travel to­wards the back of the skull, zy­go­matic arches with a more rounded ap­pear­ance, a smaller mas­toid width, a larger nasal de­pres­sion, and deeper an­gu­lar processes that lower the oc­ciputs in the skull pro­file. (Car­dini, 2004; Hoff­mann, et al., 1979; Vino­gradov and Ar­girop­ulo, 1968; Zim­ina and Gerasi­mov, 1973)

The shape of nasal cav­i­ties for black-capped mar­mots is mid-way be­tween that of Alaska mar­mots and hoary mar­mots. The mar­gin of the pre­max­illa in black-capped mar­mots is al­most straight, but the nasal bones nar­row slowly until they reach their final length. Black-capped mar­mots have a de­fined supra­or­bital notch on the edge of the frontal. The wing of the or­bital does not rise be­yond the upper edge of the lacrimal bone. There are sub­tle dif­fer­ences in the choanal and in­ci­sive foram­ina be­tween the black-capped mar­mot sub­species. (Boyesko­rov, et al., 1994; Hoff­mann, et al., 1979)

Male mar­mots have more pro­nounced sagit­tal crests, al­most dome-shaped cra­ni­ums, a small fora­men mag­num, dif­fer­ences in labial and lin­gual lower jaw mor­pholo­gies, and nar­rower in­ter- and post-or­bital areas on the skull com­pared to fe­males. The den­tal for­mula for black-capped mar­mots is in­cisors 1/1, ca­nines 0/0, pre­mo­lars 2/2, and mo­lars 3/3 to­tal­ing 24 teeth. (Car­dini, 2004; Hoff­mann, et al., 1979)

Black-capped mar­mots have a diploid kary­otype to­tal­ing 40 chro­mo­somes (2n=40 or 20 pairs of chro­mo­somes), while other Palearc­tic diploid species have 38 total chro­mo­somes (2n=38 or 19 pairs of chro­mo­somes). Olympic mar­mots have the same kary­otype as black-capped mar­mots, and so they are often thought to be more closely re­lated to Nearc­tic species as op­posed to Palearc­tic species. (Boyesko­rov, et al., 1994; Hoff­mann, et al., 1979; Le Berre and Ra­mousse, 1994; Step­pan, et al., 1999; Zim­ina and Gerasi­mov, 1973)

De­spite the nu­mer­ous sim­i­lar­i­ties in skull mor­phol­ogy, feet mor­phol­ogy, and pelage color be­tween cer­tain Nearc­tic species (hoary mar­mots and Alaska mar­mots) and black-capped mar­mots, mol­e­c­u­lar stud­ies, such as those based on Cy­tochrome b, in­di­cate that Palearc­tic mar­mot species form a mono­phyletic group. There­fore, black-capped mar­mots do not form a sis­ter group to hoary mar­mots as has been sug­gested in the past, and sim­i­lar fea­tures be­tween the two species are con­ver­gent evo­lu­tion. Im­muno­genetic dif­fer­ences have been noted be­tween the sub­species. (Boesko­rov, et al., 1999; Bran­dler, et al., 2010; Step­pan, et al., 1999; Zim­ina and Gerasi­mov, 1973)

No in­for­ma­tion is avail­able re­gard­ing spe­cific meta­bolic rates of black-capped mar­mots, but en­ergy ex­pen­di­tures may in­crease by 8 to 15 times be­tween hi­ber­na­tion and ac­tive time pe­ri­ods. It has also been noted that an­i­mals in­hab­it­ing moun­tain­ous re­gions with low tem­per­a­tures (5 to 10°C) have lower meta­bolic rates than species that live with higher tem­per­a­tures (Ward and Ar­mitage, 1981a cited in Barash, 1989). ("Squir­rels", 2006; Barash, 1989)

• Other Physical Features
• endothermic
• heterothermic
• bilateral symmetry

• Sexual Dimorphism
• male larger

• Range mass

  2.0 to 7.5 kg

  4.41 to 16.52 lb

• Range length

  450 to 510 mm

  17.72 to 20.08 in

Re­pro­duc­tion

Blacked-caped mar­mots live in fam­ily groups con­sist­ing of one dom­i­nant re­pro­duc­tive pair and sev­eral off­spring. Black-capped mar­mots are monog­a­mous and off­spring ex­hibit de­layed ma­tu­rity and de­layed dis­per­sal. As a re­sult, fam­ily groups ex­hibit re­pro­duc­tive sup­pres­sion and co­op­er­a­tive breed­ing. In­breed­ing may occur if re­pro­duc­tive sup­pres­sion is not com­plete. (Al­lainé, 2000; Blum­stein and Ar­mitage, 1999; Kapitonov, 1978; Se­menov, et al., 2001a; Se­menov, et al., 2001b)

• Mating System
• monogamous
• cooperative breeder

Male and fe­male black-capped mar­mots reach sex­ual ma­tu­rity around 3 years of age. How­ever, be­cause of their so­cial sys­tem nei­ther males nor fe­males re­pro­duce for some time after they reach ma­tu­rity. Fe­male black-capped mar­mots bear a lit­ter every two or more years. The se­vere en­vi­ron­men­tal con­di­tions of the habits ex­ploited by black-capped mar­mots do not allow fe­male mar­mots to build up enough en­ergy stores to hi­ber­nate, grow, re­pro­duce, and main­tain daily ac­tiv­i­ties to pro­duce a lit­ter every year. Sub­or­di­nate fe­males do not pro­duce lit­ters even dur­ing the years when the dom­i­nant fe­male has not pro­duced a lit­ter. ("Squir­rels", 2006; Al­lainé, 2000; Ar­mitage, 2007; Blum­stein and Ar­mitage, 1999; Le Berre and Ra­mousse, 1994; Mosolov and Tokarsky, 1994; Se­menov, et al., 2001b)

Black-capped mar­mots mate in the bur­row, usu­ally in April, be­fore they emerge from hi­ber­na­tion in mid-May. Thus, mat­ing be­hav­iors are not known. Par­tu­ri­tion oc­curs in early to mid-June, and may occur be­fore or up to one to two weeks after the mother emerges from the bur­row after hi­ber­na­tion. In­for­ma­tion was not avail­able about ges­ta­tion in black-capped mar­mots, but ges­ta­tion in mar­mots gen­er­ally last about 30 to 32 days. Mar­mot off­spring are weaned and be­come in­de­pen­dent at least 30 to 42 days after birth, but re­main with their par­ents for sev­eral years. Black-capped mar­mots give birth to off­spring that are 33 g and about 107 mm long. The av­er­age lit­ter size of black-capped mar­mots is 5, but lit­ter sizes will vary from 3 to 11. ("Squir­rels", 2006; Ar­mitage, 2007; Barash, 1989; Boyesko­rov, et al., 1994; Hayssen, et al., 1993; Hoff­mann, et al., 1979; Nowak, 1999; Se­menov, et al., 2001b; Vino­gradov and Ar­girop­ulo, 1968)

• Key Reproductive Features
• iteroparous
• seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• viviparous

• Breeding interval

  Female black-capped marmots bear a litter every two or more years.

• Breeding season

  Black-capped marmots mate during April.

• Range number of offspring

  3 to 11

• Average number of offspring

  5

• Average age at sexual or reproductive maturity (female)

  3 years

• Average age at sexual or reproductive maturity (male)

  3 years

Parental in­vest­ment has not been well-doc­u­mented for black-capped mar­mots. Nests are con­structed in the bur­row from dried veg­e­ta­tion. It is not known whether or not the fa­ther or sub­or­di­nate adults pro­vide ad­di­tional care for the off­spring once they are born, or whether the mother pro­vides sole care. Both par­ents and sub­or­di­nates pro­vide ther­moreg­u­la­tory ben­e­fits to the ju­ve­niles dur­ing hi­ber­na­tion by help­ing main­tain an op­ti­mal hi­ber­nac­ula tem­per­a­ture. The off­spring have the po­ten­tial to in­herit the home range if ei­ther of their par­ents dies. (Al­lainé, 2000; Ar­mitage, 2007; Blum­stein and Ar­mitage, 1999; Nowak, 1999; Se­menov, et al., 2001b)

• Parental Investment
• altricial
• female parental care
• post-independence association with parents
• inherits maternal/paternal territory

Lifes­pan/Longevity

The lifes­pan of black-capped mar­mots is cur­rently un­known, al­though most mar­mot spe­ices are long-lived. ("Squir­rels", 2006; Ar­mitage, 1992; Ar­mitage, 2007)

Be­hav­ior

Black-capped mar­mots are a very so­cial species that live in iso­lated fam­ily groups. Fam­ily groups in­clude a re­pro­duc­tive pair and its off­spring (Kapitonov, 1978 cited in Se­menov et al., 2001b). Off­spring may in­clude year­lings, ju­ve­niles from the cur­rent breed­ing sea­son, and male and fe­male sub­or­di­nate adults (sex­u­ally ma­ture off­spring that have not dis­persed). Male off­spring may dis­perse into un­in­hab­ited ter­ri­to­ries when they are 2-3 years old. Kam­chatka mar­mot fam­i­lies av­er­age 1.8 adult males, 1.7 adult fe­males, and 4.3 young. The size of the fam­ily de­pends on the amount of re­sources avail­able. Stud­ies have shown that black-capped mar­mots may move be­tween fam­ily groups, in­di­cat­ing that groups may not con­sist of only close rel­a­tives (Mashkin, 2003 cited in Ar­mitage, 2007). A black-capped mar­mot bur­row ex­ca­vated dur­ing the win­ter re­vealed two adults, four sub-adults, and four ju­ve­niles. This find­ing sup­ports the the­ory that the fam­ily group not only shares the same sum­mer ter­ri­tory but also ex­ploits joint hi­ber­na­tion. ("Squir­rels", 2006; Al­lainé, 2000; Ar­mitage, 2007; Barash, 1989; Filonov, 1961; Kapitonov, 1978; Mosolov and Tokarsky, 1994; Se­menov, et al., 2001a; Se­menov, et al., 2001b)

Black-capped mar­mots are ac­tive dur­ing the day (di­ur­nal) and seden­tary. They for­age ex­ten­sively dur­ing the sum­mer to put on weight in prepa­ra­tion for hi­ber­na­tion. De­spite al­most 24 hours of day­light in sum­mer months, stud­ies have found that black-capped mar­mots fol­low a typ­i­cal di­ur­nal cir­ca­dian rhythm. At night, black-capped mar­mots rest and seek shel­ter in their bur­rows, but dur­ing the day, or when the sun reaches an angle of 17°, above­ground ac­tiv­ity be­gins. Daily ac­tiv­i­ties have not been found to end at a par­tic­u­lar angle of the sun as oc­curs when ac­tiv­i­ties begin. Feed­ing pat­terns seem to fol­low a bi-modal cycle with mar­mots most ac­tive first thing in the day and then again later in the day. When tem­per­a­tures reach 15-25°C, black-capped mar­mots seek shel­ter in their bur­rows, only re-sur­fac­ing to feed once tem­per­a­tures de­crease. Dis­pers­ing adult males that have yet to find a ter­ri­tory are no­madic, spend­ing 1 to 3 nights in any par­tic­u­lar bur­row be­fore mov­ing on to the next. (Barash, 1989; Se­menov, et al., 2001a; Se­menov, et al., 2001b; Se­menov, et al., 2000)

Bur­row­ing ac­tiv­i­ties occur dur­ing the sum­mer. When the ground thaws black-capped mar­mots are able to add on to their ex­ten­sive un­der­ground tun­nels and some tun­nels have been found to reach 113 m in length (Kapitonov, 1960 cited in Zim­ina and Gerasi­mov, 1973). Bur­rows are the cu­mu­la­tive re­sult of many gen­er­a­tions of mar­mots adding on to ex­ist­ing sys­tems. Each bur­row has mul­ti­ple open­ings and mul­ti­ple cham­bers. Bur­row open­ings are under or near large rocks and 17 to 18 cm in di­am­e­ter. Three types of bur­rows, sum­mer, win­ter, and tem­po­rary, can be found through­out the mar­mot home range, and are main­tained dur­ing the sum­mer. (Barash, 1989; Mosolov and Tokarsky, 1994; Zim­ina and Gerasi­mov, 1973)

Adult mar­mots have an alert pos­ture, where they sit up­right on their haunches and scan their sur­round­ings. Mar­mots di­rect an­tag­o­nist be­hav­iors to­wards in­trud­ers while more am­i­ca­ble be­hav­iors are di­rected to­wards rel­a­tives. Black-capped mar­mots use large rocks as look-out posts duri­ing vig­i­lant be­hav­iors. (Ar­mitage, 2007; Barash, 1989; Mosolov and Tokarsky, 1994)

Black-capped mar­mots have phys­i­o­log­i­cal and be­hav­ioral adap­ta­tions that en­able them to sur­vive in cold en­vi­ron­ments with per­mafrost. Com­pared to other mar­mots, black-capped mar­mots have a higher per­cent­age of their body mass con­sist­ing of sub­cu­ta­neous and periv­is­ceral fat. This fat also does not so­lid­ify until tem­per­a­tures reach -3°C to -7°C. Black-capped mar­mots have the abil­ity to forego moult­ing on all or part of their body to re­duce en­ergy ex­pended grow­ing new fur. Win­ter bur­rows, or hi­ber­nac­ula, are con­structed in areas that re­ceive deep snow cover dur­ing the win­ter. The hi­ber­nac­u­lum is pre­pared for the win­ter by in­su­lat­ing the roof and walls. The roof of the bur­row is cov­ered with rocks and the walls of the bur­row are lined with as much as 9 to 12 kg of grass and other veg­e­ta­tive ma­te­r­ial to as­sist with heat re­ten­tion dur­ing win­ter. (Bibikov, 1994; Filonov, 1961; Se­menov, et al., 2001a; Vasil'ev, 2000; Zim­ina and Gerasi­mov, 1973)

The last mar­mot to enter the hi­ber­nac­ula plugs the en­trance in such a way that air spaces be­come trapped in the plug ma­te­r­ial. This plug con­sists of plant ma­te­r­ial, soil, feces, and rocks, and not only pro­vides in­creased in­su­la­tion but also pro­vides pro­tec­tion from preda­tors. In ad­di­tion to sav­ing en­ergy by group hi­ber­nat­ing, black-capped mar­mots have the abil­ity to lower body tem­per­a­tures to around 0°C dur­ing hi­ber­na­tion and hi­ber­nate when am­bi­ent tem­per­a­tures are below 0°C (Vasil’ev, 2000 cited in Lee, Barnes, and Buck, 2009). Even dur­ing arousal pe­ri­ods, body tem­per­a­tures do not rise to their usual ac­tive body tem­per­a­ture. Mar­mots also roll into balls and press to­gether while in hi­ber­na­tion. Arousals occur every two to four weeks, in which case the mar­mots are be­lieved to defe­cate and uri­nate. ("Squir­rels", 2006; Barash, 1989; Kapitonov, 1978; Lee, et al., 2009; Vasil'ev, 2000)

Hi­ber­na­tion in black-capped mar­mots usu­ally lasts for 8-9 months of the year from the end of Au­gust or mid-Sep­tem­ber, de­pend­ing on the en­vi­ron­men­tal con­di­tions and how long food sources are avail­able above­ground, to about the mid­dle of May. Black-capped mar­mots in cap­tiv­ity did not enter hi­ber­na­tion until No­vem­ber, or 1.5-2 months later then in the wild. In ad­di­tion to cold tem­per­a­tures, hi­ber­na­tion also be­gins as a re­sult of food de­pri­va­tion and body fat con­tent. From Feb­ru­ary to March, black-capped mar­mots were found to have the deep­est bouts of tor­por. Adult tor­por bout length typ­i­cally de­creased in du­ra­tion in mid-April, or when the mar­mots were near­ing the time to mate. Ju­ve­niles have long tor­por bouts into May with some bouts stretch­ing for 20 days. ("Squir­rels", 2006; Barash, 1989; Bibikov, 1994; Se­menov, et al., 2001b; Vino­gradov and Ar­girop­ulo, 1968)

• Key Behaviors
• terricolous
• fossorial
• diurnal
• motile
• sedentary
• hibernation
• social

• Range territory size

  0.01 to 0.21 km^2

• Average territory size

  0.13 km^2

Home Range

Home ranges oc­cu­pied by black-capped mar­mots vary in size de­pend­ing on the qual­ity of the site. One study noted that Yakut­ian mar­mots oc­cu­pied ranges that were 10 to 15 ha (0.1 to 0.15 sq km), while an­other study noted that black-capped mar­mots oc­cu­pied ranges that were 1.0 to 2.5 ha (0.01 to 0.025 sq km) in size. Ter­ri­to­ries in­clude sum­mer and win­ter bur­rows as well as for­ag­ing areas and other re­quired re­sources. Kam­chatka mar­mots have an av­er­age home range of 13 ha, or 0.13 sq km, (range 1.5 to 21 ha; 0.015 to 0.21 sq km). Mar­mot fam­ily home ranges do not over­lap, and small mar­mot colonies are often formed. (Filonov, 1961; Mosolov and Tokarsky, 1994; Se­menov, et al., 2001a)

Com­mu­ni­ca­tion and Per­cep­tion

Black-capped mar­mots use vo­cal­iza­tions such as alarm calls to no­tify group mem­bers of nearby preda­tors. The du­ra­tion of the main call is 0.2 sec­onds (Nikol’skii, 1976 cited in Hoff­mann, Koeppl, and Nadler, 1979) and has an av­er­age fre­quency of 3000 Hz. Kam­chatka mar­mots have a unique alarm call com­pared to the other two black-capped mar­mot sub­sep­cies, Bar­guzin and Yakut­ian mar­mots. Ter­ri­to­ries may be marked using dif­fer­ent ol­fac­tory cues, and vo­cal­iza­tions are used to in­di­cate mar­mot pres­ence on those ter­ri­to­ries. All mar­mots have cheek and anal glands which they use to scent mark rocks and veg­e­ta­tion. ("Squir­rels", 2006; Barash, 1989; Blum­stein and Ar­mitage, 1999; Hoff­mann, et al., 1979; Nikol­sky, et al., 1991; Nowak, 1999)

Mar­mots have promi­nent tails that can be quite bushy. Sim­i­lar to other mar­mot species that in­habit moun­tain­ous re­gions, black-capped mar­mots use their tails in vi­sual com­mu­ni­ca­tion. (Barash, 1989; Bibikov, 1994)

• Communication Channels
• visual
• acoustic
• chemical

• Other Communication Modes
• scent marks

• Perception Channels
• visual
• tactile
• acoustic
• chemical

Food Habits

The short grow­ing sea­son (3 to 4 months) makes it dif­fi­cult for black-capped mar­mots to ob­tain the fat re­serves nec­es­sary to sur­vive hi­ber­na­tion. There is often lit­tle or no food avail­able when the mar­mots start to emerge from hi­ber­na­tion, which is often why the preg­nant fe­male re­mains in the bur­row until she gives birth in June. Black-capped mar­mots are typ­i­cally her­bi­vores, con­sum­ing grasses, forbs, fruits (berries), seeds (in­clud­ing conifer cones), and shrubs. Early plant growth is pre­ferred over later growth. ("Squir­rels", 2006; Barash, 1989; Bibikov, 1994; Nowak, 1999; Se­menov, et al., 2001b; Vino­gradov and Ar­girop­ulo, 1968; Zim­ina and Gerasi­mov, 1973)

Roots and bulbs are con­sumed most to­wards the end of the sum­mer when the mar­mots are try­ing to in­crease fat stores. As many as 12 dif­fer­ent plant species are con­sumed, and seeds from Siber­ian dwarf pines (Pinus pumila) are con­sumed just be­fore the mar­mots enter hi­ber­na­tion. Some of the spe­cific species con­sumed by Kam­chatka mar­mots in­clude arc­tic herbs (Anemone sibir­ica), granny's bon­nets (Aqui­le­gia glan­du­losa), Doron­icum flow­ers (Doron­icum bar­gusi­nense), Asian globe­flow­ers (Trol­lius asi­ati­cus), and cranes­bills (Gera­nium alb­i­flo­rum). All parts of a plant may be con­sumed, in­clud­ing the flow­ers, leaves, stems, and roots. (Barash, 1989; Bibikov, 1994; Boyesko­rov, et al., 1994; Filonov, 1961; Zharov, 1976)

In ad­di­tion to plant ma­te­ri­als, most mar­mot species have been found to eat in­sects and their lar­vae, car­rion, small ro­dents lo­cated while bur­row­ing, and bird eggs. Black-capped mar­mots ob­tain water from moun­tain streams, their food, and from melt­ing snow or glac­i­ers. ("Squir­rels", 2006; Barash, 1989; Mosolov and Tokarsky, 1994)

• Primary Diet
• herbivore
  • folivore
  • frugivore
  • granivore
  • lignivore
  • eats sap or other plant foods

• Animal Foods
• insects

• Plant Foods
• leaves
• roots and tubers
• wood, bark, or stems
• seeds, grains, and nuts
• flowers

Pre­da­tion

Be­sides hu­mans, black-capped mar­mots have sev­eral preda­tors in­clud­ing gray wolves, brown bears, golden ea­gles, wolver­ines, and red foxes. Other rap­tors (hawks, owls) and mid- to large-sized car­ni­vores are also po­ten­tial preda­tors of black-capped mar­mots. Since lynxes feed on mar­mots in North Amer­ica, Eurasian lynxes may con­sume these mar­mots in­hab­it­ing Rus­sia. (Barash, 1989; Hoff­mann, et al., 1979; Mosolov and Tokarsky, 1994; Step­pan, et al., 1999)

Bur­rows pro­vide a safe refuge from preda­tors in both sum­mer and win­ter for black-capped mar­mots. The plugs placed in the en­trances to the bur­row dur­ing hi­ber­na­tion pre­vent preda­tors from ac­cess­ing the mar­mots in the win­ter. Alarm calls can be used to alert mem­bers of preda­tors or to no­tify the preda­tor that it has been spot­ted. ("Squir­rels", 2006; Barash, 1989; Blum­stein and Ar­mitage, 1999; Hoff­mann, et al., 1979; Nowak, 1999; Vasil'ev, 2000)

• Anti-predator Adaptations
• cryptic

• Known Predators

  • gray wolves (Canis lupus)
  • brown bears (Ursus arctos)
  • golden eagles (Aquila chrysaetos)
  • wolverines (Gulo gulo)
  • red foxes (Vulpes vulpes)

Ecosys­tem Roles

Black-capped mar­mots sig­nif­i­cantly alter their en­vi­ron­ments through for­ag­ing and bur­row­ing ac­tiv­i­ties, which in turn al­ters the veg­e­ta­tion com­mu­nity. Mea­sures of di­ver­sity (Shan­non-Weaver), eq­ui­tabil­ity, and species rich­ness of vas­cu­lar plants, bryophytes, and lichens dif­fer sig­nif­i­cantly be­tween areas with and with­out mar­mots. Areas im­me­di­ately sur­round­ing black-capped mar­mot bur­rows (core areas) are pre­dom­i­nantly cov­ered by grasses with few bryophytes and cryp­tograms (algae, lichens, fungi) pre­sent. The open tun­dra and the re­gion sur­round­ing the core area typ­i­cally have a greater pro­por­tion of bryophytes, forbs, and cryp­tograms com­pared to the core area. While her­bivory it­self will alter plant com­po­si­tion, for­ag­ing and bur­row­ing may in­di­rectly af­fect species com­po­si­tion through dam­ag­ing plant ma­te­ri­als. Bur­row­ing dis­turbs the soil en­vi­ron­ment, al­ter­ing de­com­po­si­tion, mois­ture and nu­tri­ent cy­cles, as well as pro­vides soil aer­a­tion. Nu­tri­ents, seeds, and soil brought to the sur­face may pro­mote fur­ther plant growth. Mar­mots will also dis­trib­ute nu­tri­ents through­out their range through uri­na­tion and defe­ca­tion. Habi­tat het­ero­gene­ity in­creases as a re­sult of black-capped mar­mot ac­tiv­i­ties. ("Squir­rels", 2006; Se­menov, et al., 2001a)

The black-capped mar­mot is a host to a par­tic­u­lar flea (Orop­sylla silantiewi) (Kapitonov, 1960b cited in Hoff­mann. Koeppl, and Nadler, 1979). Black-capped mar­mots are also po­ten­tial car­ri­ers of the plague, which poses a po­ten­tial risk to peo­ple. There are no known ces­todes as­so­ci­ated with the black-capped mar­mot. As noted, black-capped mar­mots pro­vide a source of food for many ver­te­brate species. ("Squir­rels", 2006; Ar­mitage, 2007; Hoff­mann, et al., 1979; Mosolov and Tokarsky, 1994)

• Ecosystem Impact
• disperses seeds
• soil aeration

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Be­cause of their fine, soft fur, black-capped mar­mots are often hunted for their pelts. In ad­di­tion to sup­port­ing the fur in­dus­try, it is be­lieved that mar­mots are hunted for food, though po­ten­tially to less of an ex­tent today than in the past. Fur­ther­more, black-capped mar­mots may pro­vide an im­por­tant study or­gan­ism for sci­en­tists who are in­ves­ti­gat­ing hi­ber­na­tion mech­a­nisms for med­ical pur­poses. (Car­dini, 2004; Hoff­mann, et al., 1979; Step­pan, et al., 1999; Tokarsky and Va­lent­sev, 1994; Zim­ina and Gerasi­mov, 1973)

• Positive Impacts
• food
• body parts are source of valuable material

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

The en­vi­ron­ments in­hab­ited by black-capped mar­mots are not often close to human pop­u­la­tions. There­fore black-capped mar­mots do not have a sig­nif­i­cant in­flu­ence on hu­mans. Black-capped mar­mots are po­ten­tial vec­tors of the plague, so they do pose a small health risk; how­ever, this is min­i­mal to non-ex­is­tent given their prox­im­ity to human pop­u­la­tions. Fur­ther­more, mar­mot groups are often erad­i­cated if their home ranges over­lap with human set­tle­ments, specif­i­cally if the mar­mots are be­lieved to com­pete with rein­deer for for­age. Black-capped mar­mot colonies may also be de­stroyed through human ac­tiv­i­ties such as re­source ex­trac­tion. ("Squir­rels", 2006; Mosolov and Tokarsky, 1994; Tokarsky and Va­lent­sev, 1994; Zim­ina and Gerasi­mov, 1973)

• Negative Impacts
• injures humans
  • carries human disease

Con­ser­va­tion Sta­tus

The abun­dance of this species is not well known since the species is wide­spread and not found at high den­si­ties. Mar­mot den­si­ties may fluc­tu­ate from 2 or 3 to 32 mar­mots per 10 sq km. At least two pop­u­la­tions of Yakut­ian mar­mots are en­dan­gered and have been listed in the Red Book of the Sakha Re­pub­lic (Revin et al., 1987 cited in Se­menov et al., 2001a). Within the Mar­mot genus, Yakut­ian mar­mots are con­sid­ered one of the most sus­cep­ti­ble sub­species to ex­tinc­tion. Bar­guzin mar­mots are quite rare and are pro­tected by law. Laws also reg­u­late the hunt­ing of all black-capped mar­mots. Re­searchers have noted that black-capped mar­mot pop­u­la­tions are de­clin­ing and black-capped mar­mots are no longer found in some of their pre­vi­ous ranges. (Filonov, 1961; Mosolov and Tokarsky, 1994; Se­menov, et al., 2001a; Tsyt­sulina, 2008)

• IUCN Red List

  Least Concern
  More information

• IUCN Red List

  Least Concern
  More information

• US Federal List

  No special status

• CITES

  No special status

• State of Michigan List

  No special status

Con­trib­u­tors

Lind­sey Bylo (au­thor), Uni­ver­sity of Man­i­toba, Jane Wa­ter­man (ed­i­tor), Uni­ver­sity of Man­i­toba, Laura Podzikowski (ed­i­tor), Spe­cial Pro­jects.

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