Felis manul is distributed throughout Central Asia, from western Iran to western China.
Biogeographic Regions: oriental (Native )
Body Length: 500-600mm
Tail Length: 210-310mm
The skull of Felis manul is relatively convex and rounded with a short rostrum and a strongly enlarged cranium. The large, forward facing orbits are set low on the skull in relation to the forehead. The tympanic bullae are slightly swollen in Felis manul. The coronoid process is broad and angled slightly backwards, and the angular process is short and thin. There seems to be no structural differences in skull morphology between males and females, although females skulls are usually smaller.
Felis manul is similar in size to a large domestic cat. It has long dense fur, which is generally gray or pale reddish in color. Its white guard hairs give it a frosted appearance. The dark colored fur on its underside is nearly twice as long as the fur on its back. It has a series of five to seven narrow black stripes running transversely across its lower back. The long tail is black tipped, with a series of five to seven black rings running down its length. Its short, stumpy legs are generally similar in color to the fur on the rest of the body with indistinct black bands sometimes present. The fur under the surface of the paws is generally short and reddish in appearance. Its head is small and has broad, white rimmed eyes. The eyes are unique in that they contract in small circles instead of slits like most other small wild cats. It has low set rounded ears that are generally buff colored and can have dark tips. There is silvery-gray fur with black spots on the forehead and crown. There are two narrow black stripes running down from the corners of each eye. The lips, chin and neck are white, with a slight reddish tint near the upper lip. Like most felines, it has long white whiskers.
There is huge variation in coat color across the entire range of this species. This has caused some scientists to refer to them with three different subspecies classifications. Felis manul manul has the most common coloration found (as described above), and is found throughout most of the species' range, but most frequently in Mongolia and China. F. m. ferrugineus appears to be more reddish orange in color, with distinct reddish spots and stripes. It is generally found from the Caspian Sea to Pakistan. The third subspecies, F. m. nigripectus appears more grayish in color, and has a particularly distinct silver-gray winter coat. It is found in central Asiatic Russia, Nepal, and Tibet.
Range mass: 2 to 5 kg.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: sexes alike
Habitat and Ecology
In the grass and shrub steppe of central Mongolia, annual home ranges were found to be strikingly large for a small felid, although it is not clear if such large ranges are typical for the species (Brown et al. 2003). Ongoing research there (S. Ross pers. comm. 2008) measured home ranges as follows:
Female = 100% MCP = 50.8 +/- 43.0 kmÂ², 95% MCP = 27.1 +/- 23.6 kmÂ²; n = 10
Male = 100% MCP = 158.5 +/- 179.2 kmÂ², 95% MCP = 100.4 +/- 101.2 kmÂ² n = 8
During the study Pallas cats were found to have a strong association with rocky, steep areas and were rarely found in open grasslands (where they may be more vulnerable to predation by sympatric carnivores: S. Ross pers. comm.. 2008).
Preliminary results based on radio-tracking of Pallas cats in Daursky state nature reserve (Chita region of Russia) shows that annual home range varies from 5 to 30 kmÂ² (n=3) (Kiriliuk et al. 2008 via pers. comm. A Barashkova 2008).
In China, Pallas cats feed predominately on pikas (Ochotona), small rodents (Alticola, Meriones, Cricetulus), birds (partridge-Pyrrhocorax), hares (Lepus) and marmots (Marmota), and appear to be most numerous where pikas and voles are abundant and not living under deep snow cover (Wozencraft et al. in press). In Mongolia, preliminary analysis of scats indicated that gerbils (Meriones spp) and jerboas (Dipus sagitta and Allactaga spp) were the main prey, wih lambs of the Argali sheep (Ovis ammon) taken during the spring (Murdoch et al. 2006). Populations may fluctate widely with their small mamal prey base (Purevsuren 2004). Activity is predominantly crepuscular, although they can be active at any time (S. Ross pers comm. 2008).
Although the manul is found throughout central Asia, its habitat is very specific. It lives on rocky steppes and stony outcroppings only, and has rarely been seen in the lowland areas. It has been found at altitudes up to 4,800m but only in areas where deep snow does not accumulate.
Range elevation: 4800 (high) m.
Habitat Regions: temperate ; terrestrial
Terrestrial Biomes: desert or dune ; mountains
Throughout its habitat range, Felis manul preys primarily on pikas and small rodents. It is adept at stalking and ambushing these animals on the rocky steppes where it lives. It has been known to occasionally eat small birds and insectivores as well.
Animal Foods: birds; mammals
Primary Diet: carnivore (Eats terrestrial vertebrates)
Known prey organisms
This list may not be complete but is based on published studies.
Life History and Behavior
Lifespan, longevity, and ageing
Very little is known about the reproductive habits of these solitary felines. What is known comes from studies of Felis manul in the former Soviet Union.
Litters generally range from 3 to 6 kittens in size, but some have been found to have as many as eight. Like many other felines, the kittens are blind and helpless when borne. They are typically about 12cm long and weigh around 300g at birth. The kittens molt around the age of two months, and have been observed hunting by the age of three to four months. The average life span is around 11-12 years.
Range number of offspring: 4 to 6.
Range gestation period: 74 to 75 days.
Parental Investment: altricial
IUCN Red List Assessment
Red List Category
Red List Criteria
- 2002Near Threatened
- 1996Lower Risk/least concern(Baillie and Groombridge 1996)
- 1994Insufficiently Known(Groombridge 1994)
The conservation status of Felis manul is insufficiently known due to lack of information about its range and relative numbers. It was widely hunted at one time, but availability of the animal steadily decreased until the 1980s, when hunting was finally prohibited throughout most of its range.
Although hunting no longer seems to be a problem for Felis manul, in some parts of the Russian Federation, the small rodents and pikas that the manul feeds on are being poisoned because they are considered to be carriers of disease. These prey animals are also being poisoned in some parts of China where they are believed to compete with livestock for graze. It is not clear which is a bigger threat to Felis manul, the exposure to these poisons or the decreasing food supply.
CITES: appendix ii
IUCN Red List of Threatened Species: near threatened
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, Pakistan, India (Ladakh) and Iran (Nowell and Jackson 1996, Habibi 2004), and has disappeared from much of its former range around the Caspian Sea (Belousova 1993) and Pakistan's Balochistan province (Husain 2001). Populations are small and threatened in Armenia, Azerbaijan, Russia's Krasnoyarsk region and Turkmenistan (Anon. 2008).
In Russia, the Tyva and Chita regions may have the largest populations, estimated at 2,000-2,200 and 2,100-3,000, respectively. Populations in Altai and Buryatia republics were estimated at 450-550 and 250-350 (Barashkova et al. 2007).
The manul has long been hunted for its fur in relatively large numbers in Mongolia, Russia and China, although international trade in manul pelts has largely ceased since the late 1980s (Nowell and Jackson 1996). Mongolia is the only range state which permits hunting of Pallas cats for "household purposes." The permitting system is ineffective, and Pallas cat furs are illegally exported to China (Murdoch et al. 2006). Wingard and Zahler (2006) estimate that there are approximately 1,000 hunters of Pallas's cats in Mongolia, with a mean estimated harvest of two cats per year. Pallas 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 marmot and hares (S. Ross pers. comm. 2008) Their fat and organs are used as medicine in Mongolia and Russia, and they are killed by domestic dogs (IUCN Cats Red List workshop, 2007 and A. Barashkova pers. comm. 2008).
Approximately 12% of the species? range in Mongolia occurs within protected areas (Mongolia Mammal Assessment, 2006), although Murdoch et al. (2007) found that the manul's preferred steppe-shrub 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.
In Russia, about 6% of the species' range is protected, as follows: about 1% of it is situated the state nature reserves: "Altaisky" (Altai Republic), "Sayano-Shushensky" (Krasnoyarsk region), "Ubsunurskaya Kotlovina" (Tyva Republic), "Daursky" and "Sokhodinsky" (Chita RegionHowever, only the Tyvan and Daursky state reserv protect significant habitat. About 4% of Pallas' Cat area is situated in the national parks: "Tunkinsky" (Buryatia Republic) and "Alkhanai" (Aginsky Buryatsky autonomus region, which will belong to Chita Region soon). About 0.5% of area is situated in federal wildlife refuges: "Altacheisky" (Buryatia Republic) and partly "Tsasucheisky Bor" (Chita Region). Buryatia Republic also has regional protected areas where llas cats are found (Barashkova et al. 2007).
In China it is reported from the following nature reserves: Xuelingyunshan, Tuomuerfeng, Luoshan, Baijitan, Qinghaihuniaodao, Wanglang. Wolong, Zhumulangmafeng, Kalamailishan, Qitaihuangmobanhuangmo, Aerjinshan, Ganjiahu (Xinjiang), Luobupoyeluotuo (China Species Information Service 2008).
Since 2009 this species is now a legally protected species in Afghanistan, banning all hunting and trade in its parts within the country.
Relevance to Humans and Ecosystems
Economic Importance for Humans: Negative
Felis manul is a rare animal, and has little or no significant negative impact on humans.
Economic Importance for Humans: Positive
At one time Felis manul was widely hunted in Mongolia and China for its fur, but hunting it is now prohibited. Perhaps the best thing Felis manul contributes to human society is its well-developed hunting skills. It hunts and kills small pikas and rodents, some of which are agricultural pests, and others which are "considered to be vectors for the plague" (Wikne, 1999).
Positive Impacts: body parts are source of valuable material
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.
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.
Distribution and habitat
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. 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. In 1997, they were reported for the first time as being present in the eastern Sayan Mountains.
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. Populations in the Caspian Sea region, in Afghanistan and Pakistan, are thought to be declining and becoming increasingly isolated.
In recent years, several Pallas' cats were photographed for the first time during camera trapping surveys:
- in Iran's Khojir National Park in 2008;
- in the Eastern Himalayas: in Bhutan's Wangchuck Centennial Park in April 2012; and above 4,100 m (13,500 ft) in the Jigme Dorji National Park in autumn 2012;
- in Pakistan's Qurumber National Park above 3,400 m (11,200 ft) in July 2012;
- in Nepal's Annapurna Conservation Area above 4,200 m (13,800 ft) in December 2012 and December 2013.
Distribution of subspecies
Three subspecies are recognized:
- O. m. manul (Pallas, 1776) — inhabits the northern part of the range: from Jida River south of Lake Baikal to eastern Siberia;
- O. m. nigripecta (Hodgson, 1842) — inhabits Tibet and Indian Kashmir;
- O. m. ferruginea (Ognev, 1928) — inhabits the south-western part of the range: the mountain ridge of Missanev, Kopet-Dag Mountains, Transcaspia, south-western Turkestan, northern Iran, Baluchistan and Afghanistan.
Ecology and behaviour
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.
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.
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. 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.
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 and is listed as an endangered species in Pakistan-administered Kashmir.
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. 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.
In June 2010, five kittens were born in the Red River Zoo in Fargo. A female was artificially inseminated for the first time at Cincinnati Zoo, and gave birth to three kittens in June 2011. In May 2013, three kittens were born in the Nordens Ark in Sweden.
The Pallas's cat initially was placed in the genus Felis. In 1858, the Russian explorer and naturalist Nikolai Severtzov proposed the name Otocolobus for the species. 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.
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. Otocolobus manul is estimated to have diverged from a leopard cat ancestor about 5.19 million years ago.
- 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.
- 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.
- Sunquist, Mel; Sunquist, Fiona (2002). Wild cats of the World. Chicago: University of Chicago Press. pp. 219–224. ISBN 0-226-77999-8.
- 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.
- Koshkarev, E. (1998). Discovery of manul in eastern Sayan. Cat News 29: 12–13.
- 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.
- Belousova, A. V. (1993). Small Felidae of Eastern Europe, Central Asia and Far East. Survey of the state of populations. Lutreola 2: 16.
- Habibi, K. (2003). Mammals of Afghanistan. Zoo Outreach Organisation, USFWS, Coimbatore, India.
- Chalani, M., Ghoddousi, A., Ghadirian, T., Goljani, R. (2008). First Pallas’s Cat Photo-trapped in Khojir National Park, Iran. Cat News 49: 7.
- WWF Bhutan (2012). Near threatened Pallas’ Cat found in WCP. Wangchuck Centennial Park and WWF, 16 October 2012.
- Thinley, P. (2013). First photographic evidence of a Pallas’s cat in Jigme Dorji National Park, Bhutan. Cat News 58: 27–28.
- 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.
- 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.
- Pokharel, S. (2014). New wild cat species found in ACAP area. República, 12 February 2014.
- Himalayan News Service (2014). Rare wild cat found in Annapurna region. The Himalayan Times, 12 February 2014.
- 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]
- 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.
- 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
- Bray, S. (2010). Pallas’ Cat PMP . Felid TAG Times: 3.
- Smith, S. (2008). "Himalayan kitten at park" (Video). BBC News.
- Associated Press (August 2010). "ND zoo officials boast litter of 5 Pallas kitten". WDAY News.
- CREW (June 2011). "Pallas' cats born from artificial insemination". Cincinnati Zoo’s Center for Conservation and Research of Endangered Wildlife.
- Bohusläningen (June 2013). "pallaskatterna har fått ungar". Bohusläningen.
- 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.
- 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.
- 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.
- O'Brien, S. J., Johnson, W. E. (2007). The evolution of cats. Scientific American July: 68–75.