Marmota camtschatica

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).

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.

• Biogeographic Regions
• palearctic palearctic
  • native native

• Other Geographic Terms
• holarctic 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.

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.

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 ).

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.

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.

• Habitat Regions
• polar polar
• terrestrial terrestrial

• Terrestrial Biomes
• tundra tundra
• taiga taiga
• mountains mountains

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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).

• Other Physical Features
• endothermic endothermic
• heterothermic heterothermic
• bilateral symmetry bilateral symmetry

• Sexual Dimorphism
• male larger

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.

• Mating System
• monogamous monogamous
• cooperative breeder 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.

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.

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

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.

• Parental Investment
• altricial altricial
• female parental care 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.

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.

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.

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.

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.

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.

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.

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.

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

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.

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.

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.

• Communication Channels
• visual visual
• acoustic acoustic
• chemical chemical

• Other Communication Modes
• scent marks scent marks

• Perception Channels
• visual visual
• tactile tactile
• acoustic acoustic
• chemical 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.

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.

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.

• Primary Diet
• herbivore herbivore
  • folivore folivore
  • frugivore frugivore
  • granivore 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.

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.

• Anti-predator Adaptations
• cryptic cryptic

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.

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.

• Ecosystem Impact
• disperses seeds
• soil aeration 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.

• Positive Impacts
• food 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.

• 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.

Additional Links

Encyclopedia of Life

Contributors

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

References

Allainé, D. 2000. Sociality, mating system and reproductive skew in marmots: evidence and hypotheses. Behavioural Processes , 51: 21-34.

Armitage, K. 1992. Social organization and fitness strategies of marmots. Proceedings of 1st international symposium on alpine marmot (Marmota marmots) and genera Marmota. (ed. B. Bassano, P. Durio, U. Gallo Orsi, and E. Macchi), Torino, Italy: Dipartmento di Produzionni Animali, Epidemiologia ed Ecologia: 89-94.

Armitage, K. 2007. Evolution of sociality in marmots: it begins with hibernation. Pp. 356-367 in Rodent societies: an ecological and evolutionary perspective . Chicago, Illinois: University of Chicago Press.

Barash, D. 1989. Marmots: social behavior and ecology . Stanford, California: Stanford University Press.

Bibikov, D. 1994. On the size of the marmots (Marmota), their annual cycle and adaptations to the “short summer”. In: Actual Problems of Marmots Investigation (Collection of Scientific Articles) . Moscow, Russia: ABF Publishing House.

Blumstein, D., K. Armitage. 1999. Cooperative breeding in marmots. Oikos , 84/3: 369-382.

Boeskorov, G., E. Zholnerovskaya, N. Vorontsov, E. Lyapunova. 1999. Intraspecific divergence of the black-capped marmot Marmota camtschatica (Sciuridae, Marmotinae). Zoologichesky Zhurnal , 78/7: 866-877.

Boyeskorov, G., V. Vasiliev, Y. Lukovtsev. 1994. The black-capped marmot (Marmota camtschatica Pall.) in Yakutia. Polish Ecological Studies , 20/3-4: 519-522.

Brandler, O., E. Lyapunova, A. Bannikova, D. Kramerov. 2010. Phylogeny and systematics of marmots (Marmota, Sciuridae, Rodentia) inferred from Inter-SINE PCR data. Russian Journal of Genetics , 46/3: 283-292.

Cardini, A. 2004. Evolution of marmots (Rodentia, Sciuridae): combining information on labial and lingual sides of the mandible. Acta Theriologica , 49/3: 301-318.

Erbajeva, M., N. Alexeeva. 2009. A new look at Pleistocene marmot diversity of Transbaikalia. Ethology Ecology & Evolution , 21: 237-241.

Filonov, K. 1961. Data on the ecology of Marmota camtschatica in the Barguzin state forest. Trudy Barguzinsk Gosudarst Zapovednika , 3: 169-180.

Hayssen, V., A. van Tienhoven, A. van Tienhoven. 1993. Asdell’s pattern of mammalian reproduction: a compendium of species-specific data . Ithaca, New York: Cornell University Press.

Hoffmann, R., J. Koeppl, C. Nadler. 1979. The relationships of the amphiberingian marmots (Mammalia: Sciuridae). Occasional Papers of the Museum of Natural History University of Kansas , 83: 1-56.

Kapitonov, V. 1978. Winter burrow digging of the Kamchaatka marmot in northwestern Verkhoyansk Russian-SFSR USSR. Byulleten’ Moskovskogo Obshchestva Ispyttelei Prirody Otdel Biologichesii , 83/1: 43-51.

Le Berre, M., R. Ramousse. 1994. Can marmots be considered as ecological and biological indicators?. Polish Ecological Studies , 20/2: 489-495.

Lee, T., B. Barnes, C. Buck. 2009. Body temperature patterns during hibernation in a free-living Alaska marmot (Marmota broweri). Ethology Ecology & Evolution , 21: 403-413.

Mosolov, V., V. Tokarsky. 1994. The black-capped marmot (Marmota camtschatica Pall.) in the Kronotsky Reserve. In: Actual Problems of Marmots Investigation (Collection of Scientific Articles) . Moscow, Russia: ABF Publishing House.

Nikolsky, A., N. Formozov, V. Vasiljev, G. Bojeskorov. 1991. Geographic variation of the sound signal of black-capped marmot, Marmota-camtschatica (Rodentia, Sciuridae). Zoologichesky Zhurnal , 70/2: 155-159.

Nowak, R. 1999. Marmots. Pp. 1251-1253 in Walker’s Mammals of the World , Vol. 2, 6th Edition. Baltimore, Maryland: The John Hopkins University Press.

Semenov, Y., R. Ramousse, M. Le Berre, V. Vassiliev, N. Solomov. 2001. Aboveground activity rhythm in Arctic black-capped marmot (Marmota camtschatica bungei Katschenko 1901) under polar day conditions. Acta Oecologica , 22: 99-107.

Semenov, Y., R. Ramousse, M. Le Berre, Y. Tutukarov. 2001. Impact of the black-capped marmot (Marmota camtschatica bungei) on floristic diversity of arctic tundra in northern Siberia. Arctic, Antarctic, and Alpine Research , 33/2: 204-210.

Semenov, Y., R. Ramuss, M. Leberre. 2000. Effects of ecological factors on the diurnal activity rhythms of Yakutian black-capped marmots (Marmota camtschatica bungei) in the Arctic. Russian Journal of Ecology , 31/2: 118-122.

Steppan, S., M. Akhverdyan, E. Lyapunova, D. Fraser, N. Vorontsov, R. Hoffmann, M. Braun. 1999. Molecular phylogeny of the marmots (Rodentia: Sciuridae): tests of evolutionary and biogeographic hypotheses. Systematic Biology , 48/4: 715/734.

Tokarsky, V., A. Valentsev. 1994. Distribution, biology, and breeding in captivity of black-capped marmot, Marmota-camtschatica (Rodentia, Sciuridae). Zoologichesy Zhurnal , 73/7-8: 209-222.

Tsytsulina, K. 2008. "Marmota camtschatica" (On-line). IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. Accessed November 22, 2012 at http://www.iucnredlist.org/details/12831/0 .

Vasil'ev, V. 2000. Specific features of hibernation in the black-capped marmot (Marmota camtschatica) from Yakutia. Zoologichesky Zhurnal , 79/9: 1114-1123.

Vinogradov, B., A. Argiropulo. 1968. Marmota camtschatica Pallas (1811), Kamchata or black-capped marmot. In Fauna of the U.S.S.R: Key to Rodents. [Translated from Russian by Jean Salkind] . Jerusalem: Zoological Institute of the Academy of Sciences of the U.S.S.R.

Zharov, V. 1976. Factors limiting the population density of the Kamchatka marmot Marmota-camtschatica on the Barguzin Mountain Range USSR. Zoologicheskii Zhurnal , 55/10: 1584-1586.

Zimina, R., I. Gerasimov. 1973. The periglacial expansion of marmots (Marmota) in Middle Europe during the Late Pleistocene. Journal of Mammalogy , 54/2: 327-340.

2006. Squirrels. Pp. 150-165 in The Encyclopedia of Mammals , Vol. 1, 2nd Edition. New York, New York: Facts on File.