Felis manul

Geographic Range

Pallas cats ( Otocolobus manul , Pallas 1776; syn. Felis manul , reclassified by Johnson et al . 2006) are wild monotypic felines of the genus Otocolobus , 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 palearctic
  • native native
• oriental oriental
  • native native

Habitat

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 temperate

• Terrestrial Biomes
• savanna or grassland savanna or grassland
• mountains mountains

Physical Description

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.

• Other Physical Features
• endothermic endothermic
• homoiothermic
• bilateral symmetry bilateral symmetry

• Sexual Dimorphism
• sexes alike

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 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.

• Key Reproductive Features
• iteroparous iteroparous
• seasonal breeding seasonal breeding
• year-round breeding year-round breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual sexual
• viviparous 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 female parental care
• pre-weaning/fledging
  • provisioning
    • female
  • protecting
    • female
• pre-independence
  • provisioning
    • female
  • protecting
    • female

Lifespan/Longevity

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

Behavior

Pallas cats are solitary, crepuscular carnivores. The lack of nocturnal species found in their diet supports the conclusion that Pallas cats are not nocturnal. Pallas cats are habitat specialists, having adjusted their behaviors and feeding habits to spend as much time as possible within the relatively ‘safe’ areas of rocky and ravine habitats. There is no obvious social structure indicated by space use, male home range boundaries and core areas are not exclusive from other males. Female home ranges are likely separated from those of other females, due to a patchy distribution of the desirable rocky habitat, separated by the less desirable open steppe habitats. Male Pallas cats are significantly more likely to use the relatively less safe edge areas of their home ranges, which maximizes their prey intake, to gain energy before the mating season; whereas females, and especially females with kittens, are more likely to stay within the core rocky areas of their home range and minimize time spent in open steppe habitats.

The availability of suitable den sites is a critical factor influencing Pallas cats' habitat use. Dens are used for giving birth, raising young, thermoregulation, feeding, mating and cover or escape from predators. Food is sometimes taken back to summer and winter dens, especially when kittens are present. Females take live prey, or whole uneaten birds , back to the maternal den where kittens play hunt. Wild Pallas cats have never been observed resting outside of their dens and most activities that do not require the cat to be on the surface, are undertaken in the den. Ironically, these cats cannot excavate their own burrows and therefore rely very heavily on the availability of abandoned marmots' burrows for winter dens, in addition to rock crevices and caves in the summer.

Pallas cats use a variety of behaviors when they are hunting including stalking, flushing and ambushing prey. Stalking occurs when the cat creeps slowly and low to the ground, using vegetation or rocks as cover, until it is close enough to pounce. Flushing occurs when cats walk quickly through summer undergrowth, to flush out and capture small mammals and birds . This technique has not been observed in the winter, likely because it requires high prey density and tall grass cover. Finally, ambush hunting occurs when cats wait outside a burrow for prey to emerge, before attacking, this is most commonly seen in winter.

• Key Behaviors
• terricolous
• crepuscular crepuscular
• motile motile
• solitary solitary

Home Range

Pallas cats have very large home ranges for a mesocarnivore of their size. Male home ranges are four to five times the size of females. Male home ranges vary from 20.9 square kilometers to 207.0 square kilometers, with a mean size of 98.8 (±17.2) square kilometers and a ‘core’ use area of 16.8 square kilometers. Female home ranges vary from 7.4 square kilometers to 125.2 square kilometers, with a mean size of 23.1 (±8.9) square kilometers and a ‘core’ use area of 4.2 square kilometers. Neither prey availability, nor season has a significant effect on home range size for either sex. Female home range size appears to be influenced by the connectivity of rocky habitats within the surrounding home range area. Females strongly select for rocky, connected hill-slopes and ravines as ‘core areas’ that offer maximum protection from predators. The home range size for female cats with kittens is reduced; this is likely for mothers to stay close to the maternal den, to guard her litter. In contrast, males maintain large home ranges throughout variable habitat (open steppe, rocky and ravine areas) and spend a significantly greater amount of time in less safe ‘edge areas’. This allows them to maximize their overlap with female home ranges, thereby enhancing their potential to reproduce come mating season and access higher prey density areas to gain energy before mating season begins.

Communication and Perception

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 chemical

• Other Communication Modes
• scent marks scent marks

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

Food Habits

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.

• Primary Diet
• carnivore carnivore
  • eats terrestrial vertebrates

• Animal Foods
• mammals
• carrion carrion
• insects

Predation

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.

• Anti-predator Adaptations
• cryptic cryptic

Ecosystem Roles

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 eversmannii ), 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.

Economic Importance for Humans: Positive

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

Economic Importance for Humans: Negative

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

Conservation Status

Pallas cats ( Otocolobus 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.

Additional Links

Encyclopedia of Life

Contributors

Nicole Caithness (author), University of Manitoba, Jane Waterman (editor), University of Manitoba, Leila Siciliano Martina (editor), Texas State University.

References

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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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