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Overview

Brief Summary

Biology

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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Description

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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Distribution

Range Description

The Pallass Cat primarily occurs within regions of montane grassland and shrubland steppe of Central Asia, but is found as far west as Western Iran and previously extended into Armenia and Azerbaijan. The core populations of Pallass Cat occur in Mongolia and China, where it occurs up to an elevation of 5,050 m on the Tibetan Plateau (Fox and Dorji 2007). In Russia, the Pallass Cat occurs along the border with Mongolia and China in the Altai, Tyva, Buryatia Republics, and Zabaikalsky Krai (former Chita province) (Koshkarev 1998, Barashkova et al. 2007). They are found in mountain steppe and semi desert foothills in Kazakhstan but at a much lower density than Russia, Mongolia and China (Barashkova Unpublished report). It is distributed within Eastern Kyrgyzstan but its status in the west of the country is currently unknown (Snow Leopard Trust 2014, K. Zhumabai pers. comm. 2014). Populations in the southwest of its range (the Caspian Sea region, Afghanistan and Pakistan) are diminishing, isolated and sparse (Belousova 1993, Nowell and Jackson 1996, Habibi 2003, Hameed et al. 2014). Recent records from Bhutan and Nepal also suggest its presence in the Himalayas but at a very low density (WWF 2012, Thinley 2013, Shrestha et al. 2014).

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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 )

  • Johnson, W., E. Eizirik, J. Pecon-Slattery, W. Murphy, A. Antunes, E. Teeling, S. O'Brien. 2006. The late miocene radiation of modern Felidae: a genetic assessment. Science, 311: 73-77.
  • Ross, S. 2009. Providing and ecological basis for the conservation of the Pallas's cat (Felis manul). Bristol: University of Bristol.
  • Ross, S., B. Munkhtsog, S. Harris. 2012. Determinants of mesocarnivore range use: relative effects of prey and habitat properties on Pallas's cats home-range size. Journal of Mammology, 93: 000-000.
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Range

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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Physical Description

Morphology

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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Ecology

Habitat

Habitat and Ecology

Habitat and Ecology
Pallas's Cats are distributed in landscapes with extreme continental climates - little rainfall, low humidity, and a wide range of temperatures. They are rarely found in areas where the mean ten-day snow cover depth exceeds 10 cm, and a continuous snow cover of 15-20 cm is thought to mark the ecological limit for this species (Sunquist and Sunquist 2002). They are generally associated with montane grassland steppe and shrub steppe, but show habitat specificity, selecting habitats providing hiding cover such as ravines and rocky areas that allow them to move through the landscape without being detected (Ross 2009, Ross et al. 2012). Their optimal habitat is believed to consist of a mix of grassland and shrub steppe with rocky cover, ravines and hill-slopes. They are largely absent from lowland desert basins or flat plains although may penetrate these areas along seasonal river courses or other disruptive cover (Nowell and Jackson 1996, Ross 2009).

Many aspects of Pallass Cat ecology are influenced by predation pressure from larger sympatric carnivores. Pallass Cat is predated by large raptors, the Wolf and Red Foxes. Domestic dogs are also known to be an important introduced predator (Ross et al. 2012). Due to predation pressure Pallass Cat strongly selects habitats providing hiding cover and avoids open habitats. Selection results in a low percentage (20 30%) of the landscape being utilised by the species (Ross 2009). Analyses has shown that the home range size of Pallass Cat is determined by the distribution and availability of preferred cover habitats (Ross et al. 2012).

Pallass Cat also has a strong dependency on marmot burrows and/or rock cavities as refuges. Refuges are a critical resource used on a daily basis for shelter and extremely important for raising young (Ross et al. 2010a).

The annual home ranges of Pallass Cat are unusually large for a small felid. Researchers in Mongolia have measured mean annual home ranges as follows (Ross et al. 2012):

Female = 95% kernals 23.1 8.9 km; 100% MCP = 64.1+/-18.6; n = 13

Male = 95% kernals 98.8 17.2 km, 100% MCP = 159.0 +/- 59.3 n = 9

Radio-tracking studies of Pallass Cat in the Daursky State Nature Reserve (Zabaikalsky Krai, Russia) calculated the following ranges (Barashkova and Kiriliuk 2011):


Female = 95% kernels 6.0 3.4 km; 100% MCP = 10.0 6.5 km Males = 95% kernels 22.9 12.9 km; 100% MCPs = 16.5 9.4 km

In Mongolia Pallass Cat eats a broad range of small mammals (Ochotona, Alticola, Meriones, Lasiopodomys, Cricetulus), insects, birds, reptiles, and carrion. Pallass Cat specializes on pikas, which are selected disproportionately to their availability while other more numerous prey items used less than expected. Pikas optimize energy intake per unit foraging by Pallass Cat because pikas are 24 times larger than other available small mammal prey (Ross et al. 2010b). Pallass Cat also feeds on jerboas (Dipus sagitta and Allactaga spp) and lambs of the Argali Sheep (Ovis ammon) in Mongolia (Murdoch et al. 2006). Similar feeding preferences have been observed in China, where Pallass Cats feed predominately on pikas, small rodents, birds (partridge Pyrrhocorax), hares (Lepus) and marmots (Marmota), and appear to be most numerous where pikas and voles are abundant (Wozencraft 2008).

Like its prey, Pallass Cat is predominantly active in the early morning and evening (crepuscular), but can be active at any time of day (Ross 2009). Although unproven it is likely that Pallass Cat populations fluctuate widely with their small mammal prey base (Purevsuren 2004). In addition there is a strong overlap of Pallass Cat distribution with the distribution of pika, their preferred prey (Ross 2009).


Systems
  • Terrestrial
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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

  • Heptner, V., A. Sludski. 1992. Mammals of the Soviet Union. Volume 2 Part 2: Carnivora (Felidae and Hyenidae). Lieden, Netherlands: E.J. Brill Publishers.
  • Ross, S., B. Kamnitzer, S. Harris. 2010. Den-site selection is critical for Pallas's cats (Otocolubus manul). Canadian Journal of Zoology, 88: 905-913.
  • Ross, S., J. Murdoch, D. Mallon, J. Sanderson, A. Barashkova. 2012. "Felis manul" (On-line). IUCN Red List of Threatened Species. Accessed November 03, 2012 at http://www.iucnredlist.org/details/15640/0.
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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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Trophic Strategy

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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Associations

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

  • Kenny, D., M. Lappin, F. Knightly, J. Baier, M. Brewer, D. Getzy. 2002. Toxoplasmosis in Pallas's cats (Felis manul) at the Denver Zoological Gardens. Journal of Zoo and Wildlife Medicine, 33: 131-138.
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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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Known prey organisms

Otocolobus manul preys on:
Procapra gutturosa

This list may not be complete but is based on published studies.
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Life History and Behavior

Behavior

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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Life Expectancy

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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Lifespan, longevity, and ageing

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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Reproduction

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)

  • Brown, L., J. Brown, B. Munkhtsog, W. Swanson. 2005. Exploring the ecological basis for extreme susceptibility of Pallas's cats (Felis manul) to fatal toxoplasmosis. Journal of Wildlife Diseases, 41: 691-700.
  • Clutton-Brock, T. 1989. Mammalian mating systems. Proceedings of the Royal Society of London, 236: 339-372.
  • Heptner, V., A. Sludski. 1992. Mammals of the Soviet Union. Volume 2 Part 2: Carnivora (Felidae and Hyenidae). Lieden, Netherlands: E.J. Brill Publishers.
  • Ross, S. 2009. Providing and ecological basis for the conservation of the Pallas's cat (Felis manul). Bristol: University of Bristol.
  • Ross, S., J. Murdoch, D. Mallon, J. Sanderson, A. Barashkova. 2012. "Felis manul" (On-line). IUCN Red List of Threatened Species. Accessed November 03, 2012 at http://www.iucnredlist.org/details/15640/0.
  • Swanson, W., J. Brown, D. Wildt. 1996. Influence of seasonality on reproductive traits of male Pallas's cat (Felis manul) and implications for captive management. Journal of Zoo and Wildlife Medicine, 27: 234-240.
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Conservation

Conservation Status

IUCN Red List Assessment


Red List Category
NT
Near Threatened

Red List Criteria

Version
3.1

Year Assessed
2015

Assessor/s
Ross, S., Barashkova, Y., Farhadinia, mf, Appel, A., Riordan, P., Sanderson, J. & Munkhtsog, B.

Reviewer/s
Nowell, K., Hunter, L., Mallon, D., Breitenmoser-Wrsten, C., Lanz, T. & Breitenmoser, U.

Contributor/s
Zhumabai uulu, K., Khorozyan, I. & Murdoch, J.D.

Justification

Pallass Cat has a wide but fragmented distribution in the grasslands and montane steppe of Central Asia. Even in optimal steppe habitat in Mongolia Pallass Cats occur at a very low density of two to six Pallass Cats/100 km2 (Ross in prep.). Their low density is believed to be a result of interspecific predation which restricts their use of habitats to those with good cover from predators, whilst maintaining access to prey. A consequence of habitat specialization is that a low percentage of the landscape is suitable for their needs, and their low density means that relatively large areas are required for conservation of viable populations (Ross et al. 2012). Pallass Cats are dependent on marmot burrows or other available cavities to provide dens for daily use and rearing young, which further restricts habitat availability (Ross et al. 2010a). Using the Pallass Cat studbook data (Barclay 2013) generation length has been estimated at 3.61 years. However, most populations now experience disturbances resulting in reduced lifespan and lower reproductive output (e.g. Ross 2009). Predation by sympatric carnivores, domestic dogs and human offtake are the main causes of mortality (Ross 2009). While recent records have resulted in small expansions of the species known range, increases in the number and extent of livestock, herders and herding dogs is resulting in increased habitat degradation, displacement and mortalities (Damdinsuren et al. 2008, Barashkova and Smelansky 2011, Ross et al. 2012). Mineral exploitation and infrastructure developments have also increased substantially in Central Asia increasing fragmentation throughout its core range (Selles 2013). Pallass Cat continues to be at serious risk from a declining prey base due to pika and rodent control programmes and secondary poisoning. A lack of range-wide data means there is little empirical bases on which to estimate population size and status of Pallass Cat. However, based on low detection rates, increases in habitat degradation and the species biological susceptibility to disturbance, populations are suspected to be fragmented and to have declined by 1015% over the past 11 years (three generations). The species is listed as Near Threatened as it may qualify as Vulnerable under criterion C1 in the future when the population size drops below 10,000 mature individuals (it is currently about 15,000).


History
  • 2008
    Near Threatened (NT)
  • 2002
    Near Threatened (NT)
  • 1996
    Lower Risk/least concern (LR/lc)
  • 1994
    Insufficiently Known (K)
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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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Status

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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Population

Population
Pallass Cat has a wide distribution across Central Asia and west into Iran. However it is rarely sighted. Due to recent habitat degradation and habitat specialisation shown by the species the population is likely to be fragmented and much of the supposed distribution is likely to be unoccupied by the species. Undocumented populations of Pallass Cat are still being found resulting in small extensions of its range into Nepal and Bhutan. At the same time a lack of recent records from range countries indicates possible expatriation from its historic range.

Mongolia is probably the stronghold of Pallas's Cat. In the steppe grasslands of Central Mongolia, Ross (2009) radio-collared 29 cats and estimated density at 4-8 cats/100 km. High densities of Pallass Cat have also been recorded in Russia indirectly estimated using snow tracking (Barashkova and Kiriliuk 2011). The largest Russian populations occur in Tyva Republic and Zabaikalsky Krai, with important populations in Altai and Buryatia republics (Barashkova et al. 2007).

Across the Tibetan plateau, Pallas's Cat is considered widespread but nowhere very common (Nowell and Jackson 1996). The species is considered rare and uncommon in Afghanistan (Habibi 2003) and India (Ladakh, Kashmir and Sikkim; Chanchani 2008). Few records exist from Pakistan, but the Pallass Cat is confirmed to occur in Baluchistan province, the northwest district of Chitral and in Gilgit-Baltistan province (Hameed et al. 2014). Pallass Cat is broadly distributed across Iran with most records coming from the northern mountains, but also being found in arid areas of the south and central regions (Aghili et al. 2008, Farhadinia et al. 2012, Joolaee et al. 2014). In Turkmenistan Pallass Cat is extremely rare and largely unknown. Records have only come from the mountains of the south and west, and in the highlands of the north-west (Lukarevsky 2001). Since 1983 Pallass Cat has only been recorded in the Syunt-Khasardag Reserve (S.I. Zabelin, pers. comm., cited in Lukarevsky 2001). Pallass Cat is found in eastern Kyrgyzstan, with the majority of records coming from mountainous regions in Issyk Kul Province and southern Naryn Province (K. Zhumabai pers. comm. 2014). Its occurrence in western Kyrgyzstan remains unconfirmed. No information is available on the status of the Pallass Cat in Tajikistan (Saidov 2007) and Uzbekistan (Azimov 2009), and as a result populations are assumed to be small or locally extinct. It has disappeared from much of its former range around the Caspian Sea (Belousova 1993). Its status in Azerbaijan is uncertain, and the last observation of the Pallass Cat in Armenia was made in the 1920s (Ognev 1935, Khorozyan 2010).

The area of occupancy (AOO) of Pallass Cat is estimated as 2,269,000 km. This estimate however is based on very few records (<500) from the past 15 years. Large areas between these sightings, where the status of the Pallass Cat is assumed but uncertain, are also included in the AOO. For the purpose of red listing and due to our uncertainty we have taken a precautionary approach to estimating the population size of mature individuals. Our estimate is based upon only 30% of the estimated AOO actually supporting Pallass Cat populations. Within this area we have used a density of three Pallass Cats/100 km. Using these approximations and assuming 25% of the population is immature, the population size is estimated as 15,315 mature individuals. As the population size is close to that classified as Least Concern (LC), it is possible that with more information Pallass Cat could be listed under the LC category.

Population Trend
Decreasing
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Threats

Major Threats
New information regarding Pallass Cats biological susceptibility to threats has recently been made available. Pallass Cat has several known and tested causes of vulnerability, including feeding and habitat specialization, large range sizes, dependency on shelters made by other threatened species (marmots), and vulnerability to pastoralist activities which are ubiquitous throughout its range (high associated mortality and displacement by herding activities).

The most serious threat to Pallass Cat is habitat degradation and fragmentation. Degradation is occurring through increased human population size and expansion of animal husbandry across the species range (Barashkova et al. 2007, Damdinsuren et. al. 2008, Huanguang and van Veen 2014). In Mongolia livestock numbers have increased from 33 million in 1999 to 45 million in 2013, with a forecast annual increase of 10.2% (National Statistical Office of Mongolia 2013). As well as habitat degradation, with livestock come domestic herding dogs which are known to be an important predator of Pallass Cat (Barashkova and Smelansky 2011, Farhadinia et al. 2012, Ross et al. 2012, Joolaee et al. 2014). Across the region other sources of habitat fragmentation include infrastructural developments, development of the mining industry (Awehali 2011, Paltsyn et al. 2012, Selles 2013), and the petrochemical industry in China (Abdukadir 1997). Due to habitat loss it is very likely that Pallass Cat populations are becoming increasingly fragmented and isolated. In Russia, there is believed to be a risk of loss of isolated subpopulations, particularly in Buryatia Republic (A. Barashkova pers. comm. 2014).

Another serious threat is depletion of their prey base through poisoning and over-hunting. Pikas and rodents have been targeted as pests by poisoning campaigns in China and Mongolia because they are believed to compete for forage that could otherwise be utilised by livestock, contribute to grassland degradation, and can serve as vectors for the plague (Clark et al. 2006, Smith and Xie 2008, Delibes-Mateos et al. 2011; Harris et al. 2014). Pika make up the majority of Pallass Cat diet and are extremely important for the species persistence (Ross et al. 2010b). Poisoning continues in China where pika populations have been reduced to less than 5% of pre-control densities (Lai and Smith 2003). In Mongolia poison bait campaigns to control small mammal numbers have occurred in all provinces (Clark et al. 2006, Winters 2006) and are continuing. Control of rodents in Russia is also on-going but only at small localized scales which are currently not expected to threaten the species (Shilova and Tchabovsky 2009, A. Barashkova pers. comm. 2014).

Due to sympatric predation the Pallass Cat has a dependency on marmot burrows and rock cavities, particularly for raising young (Ross et al. 2010a). Most marmot species remain non-threatened but the Siberian Marmot which overlaps Pallass Cats range in Russia and Mongolia has declined due to overharvesting and is now classified as Endangered (Batbold et al. 2008). The decline of this species may result in the loss of keystone resources for the Pallass Cat and other steppe species (Ross et al. 2010a).
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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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Management

Conservation Actions

Conservation Actions
Pallass Cat is listed under CITES Appendix II (as Felis manul). Hunting of this species is prohibited in all range countries except Mongolia (Nowell and Jackson 1996), where it has no legal protection despite being classified as Near Threatened in the country (Wingard and Zahler 2006). Trophy hunters can purchase hunting licenses to export trophies, from which US$70 is allocated to the government (Clark et al. 2006).

Approximately 12% of the species range in Mongolia occurs within protected areas (Clark et al. 2006), although Murdoch et al. (2007) found that the Pallass Cat's preferred shrub-steppe habitat was under-represented in an important protected area in Central Mongolia (the Ikh Nartiin Chuluun Nature Reserve), and that illegal hunting inside the protected area was frequent.

Due to new initiatives within Russia, protection of Pallass Cat within reserves has increased since 2008. Approximately 13% of Pallass Cat's range in Russia is now situated within protected areas. Pallass Cat habitats are included in Zabaikalsky Krai: federal refuge "Dzeren's Valley", regional wildlife refuges "Aginskaya Steppe", "Gornaya Steppe" and "Semenovsky", protective zone of Sokhondo Nature Reserve near Mongolian border. Pallass cat is also found in Sailughemsky National Park (Altai Republic) and in Shuisky Nature Park (Tyva Republic). Tyvan and Daurian State Nature Reserves remain the most important protected areas for Pallass Cat conservation. The newly created federal refuge Dzeren's Valley (subordinated to Daursky reserve) is fully situated within Pallass Cat's range (Barashkova, pers. comm. 2014, UNDP/GEF 2014).

In China Pallass Cat is reported in the following nature reserves: Xuelingyunshan, Tuomuerfeng, Luoshan, Baijitan, Qinghaihuniaodao, Wanglang, Wolong, Zhumulangmafeng, Kalamailishan, Qitaihuangmobanhuangmo, Aerjinshan, Ganjiahu (Xinjiang), Luobupoyeluotuo (China Species Information Service 2008). It is reportedly present in 29 Chinese Nature Reserves (Jutzeler et al. 2010).

The cryptic behaviour and rarity of Pallass Cat has resulted in extremely low detection rates of the species across its range. However, fundamental to conserving Pallass Cat and recognizing it within conservation action plans is understanding its occurrence. Dedicated surveys to understand the species occupancy, habitats and resources associated with its presence are needed across its range. Following this, more specific conservation action should be implemented to protect resources important for the cat's conservation.

Pallass Cat has long been hunted for its fur in relatively large numbers in Mongolia, Russia and China, but international trade in Pallass Cat pelts has largely ceased since the late 1980s (Nowell and Jackson 1996). Mongolia is the only range state which permits hunting of Pallass Cat for "household purposes". The permitting system is said to be ineffective, and Pallass Cat furs are illegally exported to China (Murdoch et al. 2006). Wingard and Zahler (2006) estimated that there are approximately 1,000 hunters of Pallas's Cats in Mongolia, with a mean estimated harvest of two cats per year. Pallass Cats are also shot due to them being mistaken for marmots, which are commonly hunted. They are also trapped incidentally in leghold traps set for wolves and foxes, and in snares set for marmot and hares (Ross 2009). Their fat and organs are used as medicine in Mongolia and Russia (IUCN Cats Red List workshop 2007 and A. Barashkova, pers. comm. 2014).
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Conservation

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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Relevance to Humans and Ecosystems

Benefits

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

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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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Wikipedia

Pallas's cat

Not to be confused with the Pampas cat

The Pallas's cat (Otocolobus manul), also called manul, is a small wild cat with a broad but fragmented distribution in the grasslands and montane steppes of Central Asia. It is negatively affected by habitat degradation, prey base decline, and hunting, and has therefore been classified as Near Threatened by IUCN since 2002.[1]

The Pallas’s cat was named after the German naturalist Peter Simon Pallas, who first described the cat in 1776 under the binomial Felis manul.[2]

Characteristics[edit]

The ears are set low and wide apart

The Pallas's cat is about the size of a domestic cat, its body is 46 to 65 cm (18 to 26 in) long and its tail is 21 to 31 cm (8.3 to 12.2 in) long. It weighs 2.5 to 4.5 kg (5.5 to 9.9 lb). The combination of its stocky posture and long, dense fur makes it appear stout and plush. Its fur is ochre with dark vertical bars on the torso and forelegs. The winter coat is greyer and less patterned than the summer coat. There are clear black rings on the tail and dark spots on the forehead. The cheeks are white with narrow black stripes running from the corners of the eyes. The chin and throat are also white, merging into the greyish, silky fur of the underparts. Concentric white and black rims around the eyes accentuate their rounded shape. The legs are proportionately shorter than those of other cats, the ears are set very low and wide apart, and it has unusually short claws. The face is shortened compared with other cats, giving it a flattened face. The pupils are circular. The shorter jaw has fewer teeth than is typical among cats, with the first pair of upper premolars missing, but the canine teeth are large.[3]

Distribution and habitat[edit]

Pallas's cats are native to the steppe regions of Central Asia, where they inhabit elevations of up to 5,050 m (16,570 ft) in the Tibetan Plateau.[4] They inhabit Mongolia, Tajikistan, Kyrgyzstan, Pakistan, Kazakhstan, Kashmir, and occur across much of western China. They also are found in the Transbaikal regions of Russia, and less frequently, in the Altai, Tyva, and Buryatia Republics.[1] In 1997, they were reported for the first time as being present in the eastern Sayan Mountains.[5]

Until the early 1970s, only two Pallas's cats were recorded in the Transcaucasus, both encountered near the Araks River in northwestern Iran, but no records existed from Azerbaijan.[6] Populations in the Caspian Sea region, in Afghanistan and Pakistan, are thought to be declining and becoming increasingly isolated.[7][8]

In recent years, several Pallas' cats were photographed for the first time during camera trapping surveys:

Distribution of subspecies[edit]

Pallas's cat at the Edinburgh Zoo

Three subspecies are recognized:[16]

Ecology and behaviour[edit]

Pallas's cat at the Zurich Zoo

Pallas's cats are solitary. Both males and females scent mark their territory. They spend the day in caves, rock crevices, or marmot burrows, and emerge in the late afternoon to begin hunting. They are not fast runners, and hunt primarily by ambush or stalking, using low vegetation and rocky terrain for cover. They feed largely on diurnally active prey species such as gerbils, pikas, voles and chukar partridges, and sometimes catch young marmots.[3]

Reproduction[edit]

The breeding season is relatively short due to the extreme climate in the cat's native range. Oestrus lasts between 26 and 42 hours, which is also shorter than in many other felids. Pallas's cats give birth to a litter of around two to six kittens after a gestation period of 66 to 75 days, typically in April or May. Such large litters may compensate for a high rate of infant mortality in the harsh environment. The young are born in sheltered dens, lined with dried vegetation, feathers, and fur. The kittens weigh around 90 g (3.2 oz) at birth, and have a thick coat of fuzzy fur, which is replaced by the adult coat after around two months. They are able to begin hunting at four months, and reach adult size at six months. Pallas's cats have been reported to live up to 11 years in captivity.[3]

Threats[edit]

Female manul: Note the "pinhole" shape of its contracted pupil

The manul has long been hunted for its fur in relatively large numbers in China, Mongolia, and Russia, although international trade in manul pelts has largely ceased since the late 1980s.[18] About 1,000 hunters of Pallas's cats are in Mongolia, with a mean estimated harvest of two cats per year. Cats are also shot because they can be mistaken for marmots, which are commonly hunted, and trapped incidentally in leghold traps set for wolves and foxes and snares set for marmots and hares. They are also killed by domestic dogs. The fat and organs of the cats are used as medicine in Mongolia and Russia. While Mongolia has not recorded any trophy exports, skin exports have grown since 2000, with 143 reported exported in 2007.[1]

Conservation[edit]

Otocolobus manul is listed in CITES Appendix II. Hunting of this felid is prohibited in all range countries except Mongolia, where it has no legal protection despite being classified as Near Threatened in the country. Since 2009, the felid is legally protected in Afghanistan, banning all hunting and trade in its parts within the country[1] and is listed as an endangered species in Pakistan-administered Kashmir.[citation needed]

Manul kitten in Parken Zoo, Sweden

In captivity[edit]

In 2010, there were 47 Pallas’s cats in 19 institutions that are members of the Association of Zoos and Aquariums, and four litters were expected. No births and three deaths occurred in 2009. Pallas’s cats have the highest percentage of 30-day mortality of any small cat at 44.9%. The seasonality of their reproduction makes it difficult to control reproductive cycles.[19] Keeping Pallas's cats healthy in captivity is difficult. They breed well, but survival rates are low owing to infections, which are attributed to an underdeveloped immune system. In their natural high-altitude habitat, they would normally not be exposed to viruses causing infection.[20]

In June 2010, five kittens were born in the Red River Zoo in Fargo.[21] A female was artificially inseminated for the first time at Cincinnati Zoo, and gave birth to three kittens in June 2011.[22] In May 2013, three kittens were born in the Nordens Ark in Sweden.[23]

Taxonomic history[edit]

The Pallas's cat initially was placed in the genus Felis.[2] In 1858, the Russian explorer and naturalist Nikolai Severtzov proposed the name Otocolobus for the species.[24] The zoologist Reginald Innes Pocock recognized the taxonomic classification of Otocolobus in 1907, described several skulls in detail, and considered the manul being an aberrant form of Felis.[25]

Following genetic studies, the monotypic genus Otocolobus has been proposed to be placed with the genera Felis and Prionailurus in the tribe Felini, because of a close phylogenetic relationship.[26] Otocolobus manul is estimated to have diverged from a leopard cat ancestor about 5.19 million years ago.[27]

References[edit]

  1. ^ a b c d e f Ross, S., Murdoch, J., Mallon, D., Sanderson, J., Barashkova, A. (2008). "Otocolobus manul". IUCN Red List of Threatened Species. Version 2014.2. International Union for Conservation of Nature. 
  2. ^ a b Pallas, P. S. (1811). Felis Manul. In: Zoographia Rosso-Asiatica, sistens omnium Animalium in extenso Imperio Rossico et adjacentibus maribus observatorum recensionem, domicillia, mores et descriptiones, anatomen atque icones plurimorum. Petropoli, in officina Caes. Acadamiae scientiarum. Vol. 1 : 20–23.
  3. ^ a b c Sunquist, Mel; Sunquist, Fiona (2002). Wild cats of the World. Chicago: University of Chicago Press. pp. 219–224. ISBN 0-226-77999-8. 
  4. ^ Fox, J. L. and Dorji, T. (2007). High elevation record for occurrence of the manul or Pallas cat on the northwestern Tibetan plateau, China. Cat News 46: 35.
  5. ^ Koshkarev, E. (1998). Discovery of manul in eastern Sayan. Cat News 29: 12–13.
  6. ^ Geptner, V. G., Sludskii, A. A. (1972). Mlekopitaiuščie Sovetskogo Soiuza. Vysšaia Škola, Moskva (in Russian); English translation: Heptner, V. G., Sludskii, A. A., Komarov, A., Komorov, N.; Hoffmann, R. S. (1992). Mammals of the Soviet Union. Volume II, Part 2: Carnivora (Hyaenas and Cats). Smithsonian Institution and the National Science Foundation, Washington, D.C. pp. 665–696.
  7. ^ Belousova, A. V. (1993). Small Felidae of Eastern Europe, Central Asia and Far East. Survey of the state of populations. Lutreola 2: 16.
  8. ^ Habibi, K. (2003). Mammals of Afghanistan. Zoo Outreach Organisation, USFWS, Coimbatore, India.
  9. ^ Chalani, M., Ghoddousi, A., Ghadirian, T., Goljani, R. (2008). First Pallas’s Cat Photo-trapped in Khojir National Park, Iran. Cat News 49: 7.
  10. ^ WWF Bhutan (2012). Near threatened Pallas’ Cat found in WCP. Wangchuck Centennial Park and WWF, 16 October 2012.
  11. ^ Thinley, P. (2013). First photographic evidence of a Pallas’s cat in Jigme Dorji National Park, Bhutan. Cat News 58: 27–28.
  12. ^ Hameed, S., Ud Din, J., Shah, K. A., Kabir, M., Ayub, M., Khan, S., Bischof, R., Nawaz, D. A. and Nawaz, M. A. (2014). Pallas's cat photographed in Qurumber National Park, Gilgit-Baltistan, Pakistan. Cat News 60: 21–22.
  13. ^ Shrestha, B., Ale, S., R. Jackson, Thapa, N., Gurung, L. P., Adhikari, S., Dangol, L., Basnet, B., Subedi, N. and M. Dhakal (2014). Nepal's first Pallas's cat. Cat News 60: 23–24.
  14. ^ Pokharel, S. (2014). New wild cat species found in ACAP area. República, 12 February 2014.
  15. ^ Himalayan News Service (2014). Rare wild cat found in Annapurna region. The Himalayan Times, 12 February 2014.
  16. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. p. 535. ISBN 978-0-8018-8221-0. OCLC 62265494. [not in citation given]
  17. ^ a b c Ellerman, J. R., Morrison-Scott, T. C. S. (1966). Checklist of Palaearctic and Indian mammals 1758 to 1946. Second edition. British Museum of Natural History, London.
  18. ^ Nowell, K., Jackson, P. (1996) Manul Octobulus manul (Pallas, 1776) In: Wild Cats: status survey and conservation action plan. IUCN/SSC Cat Specialist Group, Gland, Switzerland
  19. ^ Bray, S. (2010). Pallas’ Cat PMP . Felid TAG Times: 3.
  20. ^ Smith, S. (2008). "Himalayan kitten at park" (Video). BBC News. 
  21. ^ Associated Press (August 2010). "ND zoo officials boast litter of 5 Pallas kitten". WDAY News. 
  22. ^ CREW (June 2011). "Pallas' cats born from artificial insemination". Cincinnati Zoo’s Center for Conservation and Research of Endangered Wildlife. 
  23. ^ Bohusläningen (June 2013). "pallaskatterna har fått ungar". Bohusläningen. 
  24. ^ Severtzow, M. N. (1858). Notice sur la classification multisériale des Carnivores, spécialement des Félidés, et les études de zoologie générale qui s'y rattachent. Revue et Magasin de Zoologie Pure et Appliquée 2e Série, T. X Séptembre 1858: 386.
  25. ^ Pocock, R. I. (1907). Exhibition of a photograph and the skull of a specimen of the Manul or Pallas’ cat (Felis manul) that had recently died in the Society's Menagerie with some remarks on the species. Proceedings of the Zoological Society of London, 1907: 299–306.
  26. ^ Johnson, W., Eizirik, E., Pecon-Slattery, J., Murphy, W. J., Antunes, A., Teeling, E., O'Brien, S. J. (2006). "The Late Miocene Radiation of Modern Felidae: A Genetic Assessment". Science 311 (5757): 73–77. doi:10.1126/science.1122277. PMID 16400146. 
  27. ^ O'Brien, S. J., Johnson, W. E. (2007). The evolution of cats. Scientific American July: 68–75.

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