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Lifespan, longevity, and ageing

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Maximum longevity: 15.9 years (captivity) Observations: One captive specimen was at least 15.9 years of age when it died (Richard Weigl 2005).
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Benefits

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There are negligible negative effects to humans caused by Pallas cats.

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Jane Waterman, University of Manitoba
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Associations

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Pallas cats face the possibility of predation from a variety of terrestrial and aerial predators, such as red foxes, wolves, domestic dogs and large birds of prey. Humans are also known to hunt Pallas cats, especially in Mongolia where body parts are said to have a medicinal value and furs may be used in subsistence living or nomadic trading.

Given the threats of predation faced by Pallas cats, they spend most of their time in 'safe' habitats, such as rocky hills or ravines, while avoiding open areas like steppe and grasslands. Pallas cats rely primarily on their preferred rocky habitat to avoid detection by predators and escape if pursued. Threatened Pallas cats will run into a ‘bolt hole’, such as a rock crevice or den, slink down low and freeze next to rocks or vegetation, or run into the cover of nearby rocks or ravines. Furthermore, they move slowly compared to other sympatric and predatory carnivores and rely on crypsis and camouflage with their background, this is accomplished with their complex coat pattern and color. Pallas cats may be poor runners; therefore running is an unlikely means of escape from large terrestrial or aerial predators.

Known Predators:

  • red foxes (Vulpes vulpes)
  • Corsac foxes (Vulpes corsac)
  • wolves (Canis lupus)
  • large birds of prey (Aves)

Anti-predator Adaptations: cryptic

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Morphology

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Pallas cats are small felines, about the size of domestic cats but appear heavier due to their long, fluffy fur coats. Their body length ranges 50 to 62 cm, with tail lengths of 23 to 31 cm and they usually weigh between 2.5 and 5.0 kg. They show no significant sexual dimorphism; however, males may be slightly heavier. They have a small spherical head, a thick build, short stocky legs and a thick furry tail that does not thin or become pointed at the end. They have a short, protruding muzzle and a large forehead with bulging, large eyes, set low and directed forward, which gives the impression of a ‘flat face’. This is intensified by the shape of their broad ears and the large bunches of elongated hairs occurring on their cheeks and under their ears.

Pallas cats have very long, fluffy, silky fur that is characterized by white-tipped guard hairs that project prominently over a darker pelage, although there can be considerable inter-individual and sub-species variation of coloration. Their coat is generally a complex combination of light grey, with pale yellow to ocherous or pale yellow to reddish patches on their flanks and a dirty white posterior abdomen and groin. There are often six or seven narrow black transverse strips dorsally, extending onto the sides, which may vary in length but are always located posterior to the shoulders. Their tail is uniformly grey above and below, with a very small black tip and seven narrow black fields surrounding. Their legs are generally grey, with short brown hairs between the digits that do not cover the pads or form tufts. The darkest coloration occurs on their neck and chest, especially between their forelimbs and transitioning into a dirty white throat and abdomen. Their head is mostly a pure, light grey with scattered black spots. Their eyes, upper and lower lips and their nasal region are surrounded by white patches. There are two narrow black strips under their eyes on their cheeks, one of which terminates at their ear and the other extends around to the back of their neck. The back of their ears are grey, with a pale yellow tinge and black fringe at the tip with tufts of white hairs in front and on the inner surface of their ear pinna. White vibrissae are present on their cheeks and over their eyes.

Their skull is rounded and relatively broad laterally, with a very short rostral region and large orbits that are set vertically and directed forward. Their palate is short and broad. Tympanic bullae are set closely together and are not large, but have a swollen and highly developed antero-outer chamber, with a distinct suture between the ectotympanic and endotympanic chambers. Their angular process is short and thin. Their cheek teeth form an acute angle and are in a plane together with a line joining their molars. Their second premolars are invariably absent in the upper row, with short massive upper carnassials that lack an antero-inner cusp. There are no sex-related structural differences in the skull of Pallas cats, aside from the females being somewhat smaller.

Range mass: 2.5 to 5.0 g.

Range length: 500 to 620 mm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: sexes alike

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Life Expectancy

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Adult Pallas cats have an average lifespan of 27.1 months in the wild, with mortality heavily biased towards winter.

Average lifespan
Status: wild:
27.1 months.

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Habitat

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Pallas cats are found in temperate grassland biomes and in the mountain steppe and semi-arid desert habitats characteristic of central Asia. These areas are comprised of open flatlands, hills, upland steppe, rocky outcrops and ravine habitat. Sightings of Pallas cats have been reported up to 1,500 meters above sea level in Russia, and up to 1,700 meters above sea level in Mongolia. The habitat of Pallas cats in Russia is dominated by vegetation composed of Artemesia, Festuca, Stipa, and a variety of small shrubs. These areas are characterized by temperate continental climates with temperatures ranging from 38 degrees Celsius to -43 degrees Celsius. These areas often have small amounts of precipitation (generally less than 200 mm), most of which falls as rain in the summer. Pallas cats are poorly adapted to moving through deep snow, so their habitat is restricted to areas with less than 10 cm of snow cover during the winter.

Pallas cats are habitat specialists, confining their activity largely to, or near rocky and ravine areas, as a predator avoidance strategy. Pallas cats use marmots' (Marmota sibirica) burrows and rock crevices as dens on a daily basis, such den sites are essential habitat for this species. Dens are chosen significantly more in rocky or ravine habitats. There are three main den types used by Pallas cats on an annual basis including summer dens that are either abandoned marmots' burrows or rock dens; maternal dens, which are usually rock dens with many entrances for predator avoidance or escape; and winter dens, which are usually marmots' burrows and provide increased thermoregulatory and heat retention benefits over rock dens. Pallas cats show fidelity to a particular den for consecutive days and often return to the same den after spending time in other parts of their home range.

Habitat Regions: temperate

Terrestrial Biomes: savanna or grassland ; mountains

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Distribution

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Pallas cats (Felis manul, Pallas 1776; syn. Felis manul, reclassified by Johnson et al. 2006) are wild monotypic felines of the genus Felis, which are from the leopard cat lineage, endemic to central Asia. They are most abundant in Mongolia and the Tibetan Plateau; however, they have a broad but patchy distribution across central Asia and are considered rare and uncommon from the Caspian Sea through southern Turkmenistan, Kazakhstan, Kyrgyzstan, Iran, Afghanistan, Baluchistan, Ladakh, western and central China and southern Russia and Siberia.

Biogeographic Regions: palearctic (Native ); oriental (Native )

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Trophic Strategy

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The majority (85.5%) of Pallas cats' diet is comprised of small rodents and pikas (Ochotona dauurica). Pikas are an especially important component of their diet in the summer months. Pallas cats are dietary specialists who depend on pikas for the majority of their energy requirements, as they are two to three times larger than other available prey species. Specializing in capturing pikas reduces their foraging costs, per unit of energy gain. In addition, pikas are relatively slow moving compared to rodents and use distinct trails to move between burrows, making them much easier to locate and capture. In the winter months, there is a distinct diversification of Pallas cats' prey base, to a more generalized foraging strategy, likely due to decreasing availability of all prey. In a study of 146 scat samples, the most frequent small mammal remains were Daurian pikas (Ochotona dauurica), Mongolian gerbils (Meriones unguiculatus) and mountain voles (Alticola stoliczkanus), but there were also small amounts of passerines, carrion and insects present. There are also sexual differences in dietary diversity, with females narrowing their dietary niche to a greater degree than males in summer and generalizing their dietary niche to a greater degree than males in winter.

Animal Foods: mammals; carrion ; insects

Primary Diet: carnivore (Eats terrestrial vertebrates)

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Jane Waterman, University of Manitoba
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Associations

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There has been very little research in regards to the role Pallas cats play in steppe ecosystems. Pallas cats are ‘mesocarnivores,’ which suggests that they are mid-trophic level carnivores whose diet is comprised mostly of meat, but also some invertebrate material. They are successful predators of pikas and small rodents, and are in turn predated upon by larger carnivores, such as red foxes and wolves. They depend quite strongly on abandoned marmots' dens for shelter in the winter. Pallas cats may undergo interspecific competition with the following species: red foxes (Vulpes vulpes), Corsac foxes (Vulpes corsac) Eurasian badgers (Meles meles), steppe polecats (Mustela eversmanii), mountain weasels (Mustela altaica) and a number of raptor birds.

Pallas cats are highly susceptible to the obligate intracellular coccidian, Toxoplasma gondii, in captivity worldwide. Domestic cats, and other members of Family Felidae are likely the definitive hosts, it is transmitted placentally and through lactation, from mother to kittens. In Pallas cats, no treatments are completely effective at clearing the body of the tissue cysts caused by T. gondii and once it is contracted, it is most often fatal. This is likely the limiting factor of successful propagation of this species in captivity, as there is a high newborn mortality in kittens (~60%) within four months of birth. The cause of this extreme susceptibility to this condition in Pallas cats is unknown. The maternal immune response in seropositive pregnant Pallas cats is not protective for the kittens, as it is in domestic cats and other cat species. In a study of 15 wild and 9 captive Pallas cats, 87% of the wild individuals were negative for T. gondii, while 100% of the captive individuals were positive for the parasite. This lack of T. gondii in wild Pallas cats suggests that they have minimal exposure to this parasite in their natural habitat and only become infected after being brought into captivity. Furthermore, T. gondii oocysts have poor survivorship at high altitudes and extreme temperatures, which likely makes the hot summers and bitterly cold winters typical of Pallas cats' habitat, a difficult environment for the parasite to propagate and spread. Should the population of Pallas cats in the wild decrease to a point where extinction is imminent, it is unlikely that current captive reproductive individuals could be re-introduced, due to the severe mortality associated with T. gondii, unless a successful treatment is developed.

Commensal/Parasitic Species:

  • Toxoplasma gondii
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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Benefits

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Pallas cats have little economic importance to humans. Their furs are not valuable in today’s market, and international trade in their pelts has largely ceased since the 1980’s. Mongolia still permits hunting for ‘household purposes’; however, the permitting system is ineffective and furs are likely still illegally exported to China. Their fat and organs are still used as medicines in nomadic Mongolia and Russia and domestic dogs from nomad camps in Mongolia hunt them. Pallas cats have a negligible effect on agriculture, public health and wildlife management. It is possible that Pallas cats may provide a pest-control benefit for agriculture through their hunting of pikas and small rodents, which are the target of wide-spread and common pest control programs throughout central Asia.

Positive Impacts: body parts are source of valuable material; controls pest population

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Conservation Status

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Pallas cats (Felis manul, syn. Felis manul) are listed as “Near Threatened” by the IUCN Red List of Threatened Species, and under Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). Their population is decreasing across their range, the population in the western part of their range around the Caspian Sea is likely extirpated. They are widespread but uncommon across the Tibetan Plateau and are rare and uncommon in Afghanistan, Pakistan, India, Iran, Armenia, Azerbaijan, Russia’s Krasnoyarsk region and Turkmenistan. The Russian regions of Tyva and Chita hold the largest populations in Russia, while Mongolia appears to be the species ‘stronghold’, as they are widely distributed, but still declining across most of the country.

The major threats facing this species include declining prey bases, habitat degradation from economic expansion and direct threats posed by humans and their dogs. There have been long-term and effective vermin control programs for pikas and marmots, implemented on a large scale across central Asia. This poses a direct threat to the survival of Pallas cats. Not only do pikas represent their primary food source, but marmots' dens are an essential habitat requirement and are used extensively by Pallas cats, especially throughout the cold winter months. A decrease in either of these prey or den building species will likely have a serious impact on the populations of Pallas cats in the wild. Furthermore, habitat degradation from agriculture and mining exploration is occurring on a wide scale in Mongolia and Russia, which may lead to a further reduction in suitable rocky habitat for these cats and contribute to further fragmentation of their suitable habitat. Finally, although their furs are of relatively little economic value, even in Mongolia, Pallas cats are still occasionally shot by nomadic hunters or, more often, trapped accidentally in leg traps set for marmots, foxes and wolves.

The lack of knowledge of the ecology, reproductive strategies and population dynamics of this species makes long-term conservation efforts extremely difficult. While Pallas cats are known to occur within protected areas and wildlife and nature reserves in Mongolia, China and Russia, it is estimated that the steppe-grassland biome is the least protected of all major biomes in the world, when this is coupled with their large home ranges and patchy distributions, the effectiveness of these reserves in preserving a viable population, remains to be seen.

CITES: appendix ii

IUCN Red List of Threatened Species: near threatened

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Behavior

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Captive Pallas cats spray and cheek rub like many Felids, but this behavior is unconfirmed in wild populations. These behaviors likely provide temporal information between individuals and may reduce the probability of hostile encounters.

Communication Channels: chemical

Other Communication Modes: scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Reproduction

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Pallas cats display a polygamous mating system, typical of most felids, males mate with several females in a given mating season. There is little, to no dimorphism between sexes; however, scratches were seen on male’s faces, suggesting an ad-hoc mating strategy, where several males compete for a female. Females were not observed with fighting injuries. Mating may occur in dens, with the male staying in close proximity to the den for three to four days after copulation, likely to guard his mates during receptivity. Breeding is highly seasonal and is photo-dependent; females produce one litter of kittens per year. Mating occurs between December and March, with litters born between late March and May.

Mating System: polygynous

Gestation in female Pallas cats lasts for approximately 75 days; their altricial kittens are born blind and helpless, with dense fuzzy pelage. Litter sizes in captivity average 3.57 (±0.53) individuals per litter, but have ranged up to six or eight per litter; however, the average litter size in the wild is not known. In Mongolia, 31.9% of kittens survived to adulthood, with no significant difference between males and females.

Breeding interval: Pallas cats breed once a year.

Breeding season: The breeding season for Pallas cats occurs from December to March.

Average number of offspring: 3.57.

Range gestation period: 74 to 75 days.

Range time to independence: 4 to 5 months.

Average age at sexual or reproductive maturity (female): 1 years.

Average age at sexual or reproductive maturity (male): 1 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous

Male Pallas cats display no parental care. Kittens generally remain within the den for two months after birth, at which time they ‘molt’ into an adult coat and usually weigh between 500 to 600 g. When the kittens are three to four months old, they follow their mother for foraging in social mother-offspring groups. Kittens disperse between four to five months of age, by which time they have usually reached adult size and weight. Young may have large dispersal movements away from their maternal dens and mature quickly, they become reproductively viable within their first year.

Parental Investment: female parental care ; pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female)

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Caithness, N. 2013. "Felis manul" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Felis_manul.html
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Nicole Caithness, University of Manitoba
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Biology

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Solitary and secretive, Pallas's cat is slow but purposeful in its movements, using its environment to conceal itself and blend into its background. Pallas's cat has crepuscular activity peaks but can be active at any time during the day and night. It shelters in small caves or rock crevices and most commonly in the abandoned burrows of marmots and other animals (7). This species occurs at very low densities with as few as eight to eleven cats within 100 square kilometres. Home ranges can be from 8 to 100 square kilometres with males occupying larger areas than females and overlapping those of several females (7). This adept predator hunts by stalking and ambushing its prey, walking at speed and opportunistically pouncing on prey. It will also wait at entrances to burrows and pounce when a small mammal exits. Pallas's cat feeds primarily on rodents and small mammals such as mouse hares, pikas, murines, voles and ground squirrels, but also small birds, lizards and grasshoppers (2) (3) (4) (7). Pallas's cat is a seasonal breeder, with most litters born in April and May (6). Males follow a female for three to four days while mating, perhaps guarding her from other males while she is sexually receptive. Injuries found on males during this period suggest that fights break out between males wanting the same female (7). Litters of three to six kittens are born after a gestation period of nine to ten weeks (6). Those born in April or May will disperse by the end of August and are the size of small adults by October. Both male and female Pallas's cats breed at an age of 10 to 11 months (7).
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Conservation

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Hunting of this species is prohibited in Armenia, Azerbaijan, China, India, Iran, Kazakhstan, Kyrgyzstan, Mongolia, Pakistan, Russia, Turkmenistan and Uzbekistan (1). Its listing on Appendix II of the Convention on International Trade in Endangered Species (CITES) strictly regulates international trade of its pelt (4), although many of the countries in which Pallas's cat is found are short of funds for adequate law enforcement and protection (3).
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Description

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Pallas's cat is about the size of a domestic cat but looks much larger due to its stocky build and long, dense coat, which helps shield it from the cold in its frosty habitat (2) (6). The fur, which is nearly twice as long on the underparts and tail as it is on the top and sides (6), seasonally changes colour, from a frosted grey in winter to a grey/fox-red in the spring. Hair length and density also varies seasonally, being longer and heavier in the cold seasons (7). The short, stocky legs are marked with indistinct black bands and the bushy black-tipped tail is encircled with dark rings towards the end (6). Contrasting patches of pale white-cream fur exists on the chin, throat, inner ears and just beneath the eye, while two dark stripes run diagonally across each cheek and the crown is patterned with little black spots (6). The colour and patterning of the hair provides Pallas's cat with a high level of camouflage and amongst rocks it can remain perfectly concealed (7). The small, rounded ears are set low on the sides of the short, broad head, an adaptation to stalking prey in open country where there is little cover (3) (6). Unlike other small cats, the pupils in the large eyes of Pallas's cat contract to small circles rather than slits (3).
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Habitat

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Pallas's cat is adapted to cold, arid habitats in upland hilly areas, such as stony alpine desert and steppe grassland with rocky outcrops, but is generally absent from lowland sandy desert basins (1) (2) (6) (5). Pallas's cat seems to prefer rocky areas and ravines to the open steppe, providing protection from predators in what is otherwise a very open and exposed habitat (7). Although the species has been found up to 4,800 metres above sea level, most records come from much lower elevations, and areas with deep snow cover are generally avoided (1).
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Range

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Widely distributed but rare, Pallas's cat is found from the Caspian Sea in the west through Kazakhstan, Pakistan and northern India to parts of China and Mongolia (4). The species is now thought to be most abundant on the cold grasslands of Mongolia, Inner Mongolia and the Tibetan Plateau. Elsewhere it is considered vulnerable to rare and uncommon (6).
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Status

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Classified as Near Threatened (NT) on the IUCN Red List 2007 (1), and listed on Appendix II of CITES (4). There is currently one recognised subspecies: Otocolobus manul ferrugineus (red manul), which is classified as Lower Risk / Near Threatened (LR/nt) on the IUCN Red List 2007 (1). Two other subspecies have been suggested: O. m. manul and O. m. nigripectus (5).
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Threats

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Pallas's cat has long been hunted for its luxurious fur, with large annual harvests in China and Mongolia (6). With declining numbers in the wild and increasing restrictions, international trade in the animal's pelt has thankfully declined in recent years (1). Significant threats also exist in the form of large scale poisoning of pika and vole populations, an important prey item. This occurs in parts of the Russian Federation, where the pika is considered vermin that carries and transmits the plague, and in parts of China, where it is thought to compete with domestic stock for food (6). Pallas's cats, and particularly its young kittens, are also highly susceptible to toxomoplasmosis, a disease caught from the rodents on which it feeds (8). While there is currently low incidence of disease in its habitat, this could increase with climate change and global warming and give cause for concern. Lastly, habitat fragmentation and development is an increasing issue within the range of Pallas's cats. This may result in local extinction of this already rare species through habitat destruction and increased incidence of domestic dogs which prey heavily on Pallas's cats (7). Human alteration of the steppe ecosystem is also likely to result in adverse changes to the ecological community in which it lives (3).
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Pallas's cat

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The Pallas's cat (Otocolobus manul), also known as the manul, is a small wild cat with long and dense light grey fur, and rounded ears set low on the sides of the head. Its head-and-body length ranges from 46 to 65 cm (18 to 26 in) with a 21 to 31 cm (8.3 to 12.2 in) long bushy tail. It is well camouflaged and adapted to the cold continental climate in its native range, which receives little rainfall and experiences a wide range of temperatures.

The Pallas's cat was first described in 1776 by Peter Simon Pallas, who observed it in the vicinity of Lake Baikal. Since then, it has been recorded across a large region in Central Asia, albeit in widely spaced sites from the Caucasus, Iranian Plateau, Hindu Kush, parts of the Himalayas, Tibetan Plateau to the Altai-Sayan region and South Siberian Mountains. It inhabits rocky montane grasslands and shrublands, where the snow cover is below 15–20 cm (6–8 in). It finds shelter in rock crevices and burrows, and preys foremost on lagomorphs and rodents. The female gives birth to between two and six kittens in spring.

Due to its widespread range and assumed large population, the Pallas's cat has been listed as Least Concern on the IUCN Red List since 2020. Some population units are threatened by poaching, prey base decline due to rodent control programs, and habitat fragmentation as a result of mining and infrastructure projects.

The Pallas's cat has been kept in zoos since the early 1950s. As of 2018, 60 zoos in Europe, Russia, North America and Japan participate in Pallas's cat captive breeding programs.

Taxonomy

Felis manul was the scientific name used by Peter Simon Pallas in 1776, who first described a Pallas's cat that he had encountered near the Dzhida River southeast of Lake Baikal.[3][4][5] Several Pallas's cat zoological specimens were subsequently described:

Otocolobus was proposed by Johann Friedrich von Brandt in 1842 as a generic name.[8][9] Reginald Innes Pocock recognized the taxonomic rank of Otocolobus in 1907, described several Pallas's cat skulls in detail and considered the Pallas's cat an aberrant form of Felis.[10]

In 1951, John Ellerman and Terence Morrison-Scott considered

Since 2017, the Cat Classification Task Force of the Cat Specialist Group recognises only two subspecies as valid taxa, namely:[1]

Phylogeny

Phylogenetic analysis of the nuclear DNA in tissue samples from all Felidae species revealed that the evolutionary radiation of the Felidae began in Asia during the late Miocene around 14.45 to 8.38 million years ago.[11][12] Analysis of mitochondrial DNA of all Felidae species indicates a radiation at around 16.76 to 6.46 million years ago.[13] The Pallas's cat is estimated to have genetically diverged from a common ancestor with the genus Prionailurus between 8.55 to 4.8 million years ago based on analysis of nuclear DNA.[11] Based on analysis of mitochondrial DNA, it diverged 9.4 to 1.46 million years ago from a common ancestor with Felis.[13]

Phylogenetic relationships of the Pallas's cat as derived through analysis of
nuclear DNA:[11] Felidae Felinae Prionailurus

Leopard cat (P. bengalensis)

Fishing cat (P. viverrinus)

Flat-headed cat (P. planiceps)

Rusty-spotted cat (P. rubiginosus)

6.54–3.42 mya Otocolobus

Pallas's cat

8.55–4.8 mya

other Felinae lineages

Pantherinae

mitochondrial DNA:[13] Felidae Felinae Felis

other Felis species

Jungle cat (F. chaus)

6.52–1.03 mya

Pallas's cat

9.40–1.46 mya Prionailurus

Leopard cat

Fishing cat

Flat-headed cat

Rusty-spotted cat

8.76–0.73 mya 13.10–4.21 mya

other Felinae lineages

Pantherinae

Characteristics

Pallas's cat in Edinburgh Zoo
Pallas's cat in Howletts Wild Animal Park

The Pallas's cat's fur is light grey with pale yellowish-ochre or pale yellowish-reddish hues.[6] Some hair tips are white and some blackish. Its fur is greyer and denser with fewer markings visible in winter than in the summer.[14]: 668–669  The forehead and top of the head are light grey with small black spots. It has two black zigzag lines on the cheeks running from the corner of the eyes to the jaw joints.[6] Its chin, whiskers, lower and upper lips are white.[14]: 669  It has narrow black stripes on the back, consisting of five to seven dark transversal lines across the lower back.[6] Its grey tail has seven narrow black rings and a black tip.[14]: 669  The underfur is 40 mm (1.6 in) long and 19 μm thick, and the guard hairs up to 69 mm (2.7 in) long and 93 μm thick on the back. Its fur is soft and dense with up to 9,000 hairs/cm2 (58,000 hairs/in2).[14]: 666 

The Pallas's cat's ears are grey with a yellowish tinge on the back and a darker rim, but with whitish hair in front and in the ear pinnae. Its rounded ears are set low on the side, such that it can peer over an object and show only a relatively small part of the head above the eyes without depressing the ears. This can give its face a look of ferocity and unrest.[10] Its eyes are encircled by white. The iris is yellowish, and its pupils contract to small circular disks in sunlight.[10]: 301  Among the Felinae, it shares this trait of round pupils with Puma, Herpailurus and Acinonyx species.[15]

The Pallas's cat is about the size of a domestic cat (Felis catus).[6] Its stocky posture with the long and dense fur make it appear stout and plush. Its head-to-body is 46 to 65 cm (18 to 26 in) long with a 21 to 31 cm (8.3 to 12.2 in) long tail. It weighs 2.5 to 4.5 kg (5 lb 8 oz to 9 lb 15 oz).[16] Its body is stout, and its skull is rounded with a short nasal bone, an enlarged cranial part and rounded zygomatic arches. Its orbits are large and directed forward. Its legs are short with short and sharp retractile claws.[17]

The skull of males is 87.2–95.1 mm (3.43–3.74 in) long and 66–74 mm (2.6–2.9 in) wide at the base. Females have a 84.1–96.3 mm (3.31–3.79 in) long and 64.7–68.1 mm (2.55–2.68 in) wide skull.[14]: 671–674  The lower carnassial teeth are powerful, and the upper carnassials are short and massive. The first pair of upper premolars is absent. The dental formula is 3.1.2.13.1.2.1 × 2 = 28.[17] It has a bite force at the canine tip of 155.4 newtons and a bite force quotient at the canine tip of 113.8.[18]

The mitochondrial genome of the Pallas's cat consists of 16,672 base pairs containing 13 protein-coding, 22 transfer RNA and two ribosomal RNA genes and one non-coding RNA control region.[19]

Distribution and habitat

The Pallas's cat's range extends from the Caucasus eastward to Central Asia, Mongolia and adjacent parts of Dzungaria and the Tibetan Plateau. It inhabits montane shrublands and grasslands, rocky outcrops, scree slopes and ravines in areas, where the continuous snow cover is below 15–20 cm (6–8 in).[14] In the southwestern part of its range, the habitat of the Pallas's cat is affected by cold and dry winters, and moderate to low rainfall in warm summers.[20] The typical vegetation in this part consists of small shrubs, sagebrush (Artemisia), Festuca and Stipa grasses.[14] In the central part of its range, it inhabits hilly landscapes, high plateaus and intermontane valleys that are covered by dry steppe or semi-desert vegetation, such as low shrubs and xerophytic grasses.[21] The continental climate in this region exhibits a range of 80 °C (140 °F) between the highest and lowest air temperatures, dropping to −50 °C (−58 °F) in winter.[14]: 684–688 

The Greater Caucasus region is considered climatically suitable for the Pallas's cat.[20] In Armenia, an individual was killed near Vedi in the mountains of Ararat Province in the late 1920s.[17] In January 2020, an individual was sighted about 140 km (90 mi) farther north in Tavush Province; the habitat at this location transitions from semi-desert to montane steppe at an elevation of about 570 m (1,900 ft).[22] Records in Azerbaijan are limited to a Pallas's cat skin found in Karabakh and a sighting of an individual in Julfa District, both in the late 20th century.[23]

The Pallas's cat inhabits rocky slopes in the Koh-i-Baba Range of the Hindu Kush.

On the Iranian Plateau, two Pallas's cats were encountered near the Aras River in northwestern Iran before the 1970s.[14] In the area, an individual was captured at an elevation of about 1,500 m (5,000 ft) near Azarshahr in East Azerbaijan Province in 2008.[23] In the same year, a camera trap recorded a Pallas's cat on the southern slopes of the central Alborz Mountains in Khojir National Park shortly after heavy snowfall.[24] Farther east in the Alborz Mountains, an individual was recorded among rocks at an elevation of 2,441 m (8,009 ft) in 2016.[25] In the Aladagh and Kopet Dag Mountains, the Pallas's cat was recorded inside and in the vicinity of protected areas.[26] In the south of the Zagros Mountains, an individual was caught in a corral used by transhumant pastoralists in Abadeh County in 2012. The surrounding area consists of rocky steppe habitat dominated by mountain almond (Prunus scoparia), Astragalus and Artemisia.[27] In the Hindu Kush, a Pallas's cat was observed sunbathing at the fringe of a rocky high-elevation plain near Dasht-e Nawar in Afghanistan's Koh-i-Baba range in April 2007.[28] The Pallas's cat was also photographed multiple times in Bamyan Province between 2015 and 2017.[20] In Gilgit-Baltistan, an individual was recorded on a ridge in a juniper dominated forest at 3,445 m (11,302 ft) in Pakistan's Qurumber National Park in July 2012.[29]

In the Transcaspian Region, its presence was first reported in the Kopet Dag mountains and in the vicinity of the Tedzhen and Murghab Rivers in the late 19th century.[30] In Turkmenistan's Sünt-Hasardag Nature Reserve, a camera trap recorded an individual in 2019. The Pallas's cat is allegedly also present in Köpetdag Nature Reserve.[31]

Historical records of the Pallas's cat are known in the Surxondaryo Region and Gissar Range along the border of Tajikistan and Uzbekistan.[14] In Kyrgyzstan, it is present at high elevations of Sarychat-Ertash State Nature Reserve and in the foothills of the Alay Range.[21] In 2013, a dead female was found in a valley near Engilchek, Kyrgyzstan.[32] In Kazakhstan, it inhabits the highlands and steppes of central and east Kazakhstan Region, the periphery of the Betpak-Dala Desert, the northern Balkhash District and the Tarbagatai Mountains.[21]

The species inhabits grassland on the Ukuk Plateau.

In the South Siberian Mountains, it inhabits grasslands on the Ukok Plateau and in the Altai, Kuray and Saylyugem Mountains.[33] It is also present in Chagan-Uzun and Argut river basins, Mongun-Taiga, Uvs Lake Basin, Sayano-Shushenski Nature Reserve, Tunkinsky National Park, Lake Gusinoye basin and in the interfluves of the Selenga, Chikoy and Khilok rivers.[34] In the eastern Sayan Mountains, its presence was documented for the first time in 1997.[35] In Transbaikal, it inhabits montane steppes at elevations of 600–800 m (2,000–2,600 ft), where annual rainfall ranges from 150 to 400 mm (5.9 to 15.7 in).[36] In 2013, an individual was observed on the Vitim Plateau.[37]

The Pallas's cat inhabits the semi-desert steppe of Ikh Nartiin Chuluu Nature Reserve in Mongolia.[38] In Khustain Nuruu National Park and Gobi Gurvansaikhan National Park, it prefers rocky and rugged habitats that provides cover and camouflage.[39][40] On the Tibetan plateau, two Pallas's cats were observed in undulating alpine meadow amidst plateau pika (Ochotona curzoniae) colonies at 4,087 m (13,409 ft) in western China's Qumarlêb County in 2001. One of them swam across an irrigation channel.[41] In Gêrzê County, an individual was sighted in desert steppe habitat at an elevation of 5,050 m (16,570 ft) in 2005.[42] In 2011, the Pallas's cat was photographed in an alpine meadow in the core area of Sanjiangyuan National Nature Reserve.[43] In Ruoergai, it was observed at several places in habitat that was frequented by pastoralists and their livestock herds.[44][45]

The Pallas's cat occurs in alpine pastures of the upper Marshyangdi river valley in the central Himalayas.

The presence of the Pallas's cat in the Himalayas was first reported in Ladakh's Indus valley in 1991.[46] In Changthang Wildlife Sanctuary, Pallas's cats were sighted close by riverbanks at elevations of 4,202 and 4,160 m (13,786 and 13,648 ft) in 2013 and 2015.[47] In Gangotri National Park, a Pallas's cat was photographed in rocky alpine scrub at 4,800 m (15,700 ft) in 2019.[48] In Sikkim, an individual was observed on a rocky slope at an elevation of 5,073 m (16,644 ft) in the vicinity of Tso Lhamo Lake in 2007.[49] In December 2012, the Pallas's cat was recorded for the first time in the Nepal Himalayas. It was photographed in the upper Marshyangdi river valley in alpine pastures at elevations of 4,200 m (13,800 ft) and 4,650 m (15,260 ft) in Annapurna Conservation Area.[50] In Shey-Phoksundo National Park, Pallas's cat scat was detected at 5,593 m (18,350 ft) in 2016, the globally highest record to date.[51] In January 2012, it was recorded for the first time in Bhutan, namely in rolling hills dominated by glacial outwash and alpine steppe vegetation in Wangchuck Centennial National Park.[52] In autumn 2012, it was also photographed at an elevation of 4,122 m (13,524 ft) in Jigme Dorji National Park.[53] In 2019, scat samples of two individuals were found in Sagarmatha National Park, providing the first genetic evidence of the cat's presence in the eastern Himalayas.[54]

Behaviour and ecology

Pallas's cat

The Pallas's cat is solitary.[14] Of nine Pallas's cat kittens observed in captivity, only the two males scent marked by spraying urine.[55]

The Pallas's cat uses caves, rock crevices and marmot burrows as shelter.[14]: 690–691  In central Mongolia, 29 Pallas's cat were fitted with radio collars between June 2005 and October 2007. They used 101 dens during this time, including 39 winter dens, 42 summer dens and 20 dens for raising kittens. The summer and winter dens usually had one entrance with a diameter of 15.6 to 23.4 cm (6.1 to 9.2 in). They resided in the summer dens for 2–21 days, and in the winter dens for 2–28 days. Summer and maternal dens were close to rocky habitats with little direct sunlight, whereas winter dens were closer to ravines.[56] The home ranges of 16 females varied from 7.4 to 125.2 km2 (2.9 to 48.3 sq mi). The home ranges of nine males varied from 20.9 to 207.0 km2 (8.1 to 79.9 sq mi) and overlapped those of one to four females and partly also those of other males. The sizes of their home ranges decreased in winter.[39]

In an unprotected area in central Mongolia, Pallas's cats were mainly crepuscular between May and August, but active by day from September to November.[57] Pallas's cats recorded in four study areas in the western Mongolian Altai mountains were also active during the day, but with a lower frequency at sites where livestock was present.[58]

Hunting and diet

The Pallas's cat is a highly specialised predator of small mammals, which it catches by stalking or ambushing near exits of burrows. It also pulls out rodents with its paws from shallow burrows. In the Altai Mountains, remains of long-tailed ground squirrel (Urocitellus undulatus), flat-skulled shrew (Sorex roboratus), Pallas's pika (Ochotona pallasi) and bird feathers were found near breeding burrows of Pallas's cats. In Transbaikal, it preys on Daurian pika (Ochotona dauurica), steppe pika (O. pusilla), Daurian ground squirrel (Spermophilus dauricus) and young of red-billed chough (Pyrrhocorax pyrrhocorax).[14] Scat samples of the Pallas's cat collected in the bufferzone of Khustain Nuruu National Park in central Mongolia contained foremost remains of Daurian pika, Mongolian gerbil (Meriones unguiculatus), Mongolian silver vole (Alticola semicanus) and remains of passerine birds, beetles and grasshoppers.[59] Brandt's vole (Lasiopodomys brandtii) dominated in the diet of Pallas's cats in Mongolia's Sükhbaatar Province after the irruptive growth of this vole population during 2017 to 2020.[60] Scat found in Shey-Phoksundo National Park contained remains of pika species and of woolly hare (Lepus oiostolus).[51] Remains of a cypriniform fish were found in Pallas's cat scat in Gongga Mountain Nature Reserve.[61]

Reproduction and life cycle

Pallas's cat kittens in Parken Zoo

The female is sexually mature at the age of about one year.[62] She is in estrus for 26 to 42 hours.[63] Gestation lasts 66 to 75 days.[62][63]

A captive male Pallas's cat housed under natural lighting conditions showed increased aggressive and territorial behaviour at the onset of the breeding season, lasting from September to December. Its blood contained three times more testosterone than in the non-breeding season, and its ejaculate was more concentrated with more normal sperm forms and a higher motility of sperm.[64]

In the wild, the female gives birth to a litter of two to six kittens between the end of April and late May. The newborn kittens' fur is fuzzy, and their eyes are closed until the age of about two weeks.[14]: 693  A newborn male kitten born in a zoo weighed 89 g (3.1 oz), measured 12.3 cm (4.8 in) and had a 5.5 cm (2.2 in) long tail.[62]

In central Mongolia, seven females with kittens were observed using 20 dens for 4–60 days. Their maternal dens were either among rocks, or in former burrows of the Tarbagan marmot (Marmota sibirica), and had at least two entrances.[56] In Iran, a Pallas's cat was observed using cavities of aged Greek juniper (Juniperus excelsa) as breeding dens for a litter of four kittens.[65]

Two-month-old kittens weigh 500–600 g (17.6–21.2 oz), and their fur gradually grows longer. They start hunting at the age of about five months and reach adult size by the age of six to seven months.[14]: 694 

Threats

In China, Mongolia and Russia, the Pallas's cat was once hunted for its fur in large numbers of more than 10,000 skins annually. In China and the former Soviet Union, hunting of the Pallas's cat decreased in the 1970s when it became legally protected. Mongolia exported 9,185 skins in 1987, but international trade has ceased since 1988.[66] However, domestic trade of its skins and body parts for medicinal purposes continues in the country, and it may be hunted throughout the year.[67]

Cases of herding dogs killing Pallas's cats were reported in Iran, Kazakhstan and the Altai Republic.[26][33][21] Pallas's cats have also fallen victim in traps set for small mammals in Kazakhstan and in the Altai Republic. In Transbaikal, the Pallas's cat is threatened by poaching. In Mongolia, the use of the rodenticide bromadiolone in the frame of rodent control measures in the early 21st century poisoned the prey base of carnivores and raptors.[21] In the Sanjiangyuan region of the Tibetan Plateau, 54,147 km2 (20,906+14 sq mi) of grassland was poisoned between 2005 and 2009, leading to an estimated loss of 50,000–80,000 tonnes (55,000–88,000 short tons) of pika biomass.[68] The Pallas's cat may be negatively affected by habitat fragmentation due to mining and infrastructure projects.[2]

Conservation

Pallas's cat on the Russian Federation two-ruble coin, silver, reverse.

On the IUCN Red List, the Pallas's cat is classified as Least Concern since 2020 because of its wide-spread range and assumed large global population. It is listed in CITES Appendix II. Hunting it is prohibited in all range countries except Mongolia. Since 2009, it is legally protected in Afghanistan, where all hunting and trade with its body parts is banned.[2] On the Mongolian Red List of Mammals, it is listed as Near Threatened since 2006.[67] In China, it is listed as Endangered.[69] In Turkmenistan, it is proposed to be listed as Critically Endangered due to the scarcity of contemporary records.[70]

In captivity

Between 1951 and 1979, the Beijing Zoo kept 16 Pallas's cats, but they lived for less than three years.[71] In 1984, the Pallas's cat was designated as a priority species for captive breeding of the American Association of Zoos and Aquariums's Species Survival Plan.[72] Almost half of the kittens born in member zoos died within the first 30 days, reaching the highest mortality rate in captivity of any small wild cat.[73]

Zoos in the former Soviet Union received most of the wild-caught Pallas's cats from the Transbaikal region and a few from Mongolia. Moscow Zoo initiated a studbook for the Pallas's cat in 1997. Since 2004, the Pallas's cat international studbook has been managed by the Royal Zoological Society of Scotland, which also coordinates the captive breeding program for the Pallas's cat within the European Endangered Species Programme. As of 2018, 177 Pallas's cats were kept in 60 zoos in Europe, Russia, North America and Japan.[74]

In 2011, a female Pallas's cat was artificially inseminated for the first time with semen from the male at the Cincinnati Zoo. After 69 days, she gave birth to four kittens, of which one was stillborn.[75]

Etymology

'Manul' is the Pallas's cat's name in the Kyrgyz language. It is called 'manol' in the Mongolian language.[4] The common name 'Pallas's cat' was coined by William Thomas Blanford in honour of Peter Simon Pallas.[76]

In popular culture

The Pallas's cat is featured in a Russian Internet meme known as "Pet the cat" introduced in 2008. It is typically an image macro with a picture of an unfriendly and stern-looking Pallas's cat accompanied by a caption in which the cat invites the reader to pet it.[77][78][79][80] In 2012, the Pallas's cat overwhelmingly won an online vote to become the Moscow Zoo mascot.[81]

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Pallas's cat: Brief Summary

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The Pallas's cat (Otocolobus manul), also known as the manul, is a small wild cat with long and dense light grey fur, and rounded ears set low on the sides of the head. Its head-and-body length ranges from 46 to 65 cm (18 to 26 in) with a 21 to 31 cm (8.3 to 12.2 in) long bushy tail. It is well camouflaged and adapted to the cold continental climate in its native range, which receives little rainfall and experiences a wide range of temperatures.

The Pallas's cat was first described in 1776 by Peter Simon Pallas, who observed it in the vicinity of Lake Baikal. Since then, it has been recorded across a large region in Central Asia, albeit in widely spaced sites from the Caucasus, Iranian Plateau, Hindu Kush, parts of the Himalayas, Tibetan Plateau to the Altai-Sayan region and South Siberian Mountains. It inhabits rocky montane grasslands and shrublands, where the snow cover is below 15–20 cm (6–8 in). It finds shelter in rock crevices and burrows, and preys foremost on lagomorphs and rodents. The female gives birth to between two and six kittens in spring.

Due to its widespread range and assumed large population, the Pallas's cat has been listed as Least Concern on the IUCN Red List since 2020. Some population units are threatened by poaching, prey base decline due to rodent control programs, and habitat fragmentation as a result of mining and infrastructure projects.

The Pallas's cat has been kept in zoos since the early 1950s. As of 2018, 60 zoos in Europe, Russia, North America and Japan participate in Pallas's cat captive breeding programs.

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