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The Pallas's cat (Otocolobus manul), also known as the manul, is a small wild cat with long and dense light grey fur, and rounded ears set low on the sides of the head. Its head-and-body length ranges from Template:Cvt with a Template:Cvt long bushy tail. It is well camouflaged and adapted to the cold continental climate in its native range, which receives little rainfall and experiences a wide range of temperatures.
The Pallas's cat was first described in 1776 by Peter Simon Pallas, who observed it in the vicinity of Lake Baikal. Since then, it has been recorded across a large region in Central Asia, albeit in widely spaced sites from the Caucasus, Iranian Plateau, Hindu Kush, parts of the Himalayas, Tibetan Plateau to the Altai-Sayan region and South Siberian Mountains. It inhabits rocky montane grasslands and shrublands, where the snow cover is below Template:Cvt. It finds shelter in rock crevices and burrows, and preys foremost on lagomorphs and rodents. The female gives birth to between two and six kittens in spring.
Due to its widespread range and assumed large population, the Pallas's cat has been listed as Least Concern on the IUCN Red List since 2020. Some population units are threatened by poaching, prey base decline due to rodent control programs, and habitat fragmentation as a result of mining and infrastructure projects.
The Pallas's cat has been kept in zoos since the early 1950s. Template:As of 60 zoos in Europe, Russia, North America and Japan participate in Pallas's cat captive breeding programs.
EtymologyEdit
'Manul' is the Pallas's cat's name in the Mongolian language. It is called 'manol' in the Kyrgyz language.<ref name=Pallas/> The common name 'Pallas's cat' was coined by William Thomas Blanford in honour of Peter Simon Pallas.<ref>Template:Cite book</ref>Template:Failed verification
TaxonomyEdit
Felis manul was the scientific name used by Peter Simon Pallas in 1776, who first described a Pallas's cat that he had encountered near the Dzhida River southeast of Lake Baikal.<ref name="Pallas1776">Template:Cite book</ref><ref name="Pallas">Template:Cite book</ref><ref name=ems1951/> Several Pallas's cat zoological specimens were subsequently described:
- Felis nigripectus proposed by Brian Houghton Hodgson in 1842 was based on three specimens from Tibet.<ref name=Hodgson1842>Template:Cite journal</ref>
- Otocolobus manul ferrugineus proposed by Sergey Ognev in 1928 was an erythristic specimen from the Kopet Dag mountains.<ref>Template:Cite journal</ref>
Otocolobus was proposed by Johann Friedrich von Brandt in 1842 as a generic name.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref> Reginald Innes Pocock recognized the taxonomic rank of Otocolobus in 1907, described several Pallas's cat skulls in detail and considered the Pallas's cat an aberrant form of Felis.<ref name=Pocock1907>Template:Cite journal</ref>
In 1951, John Ellerman and Terence Morrison-Scott considered
- the nominate subspecies Felis manul manul to be distributed from Russian Turkestan to Transbaikalia;<ref name=ems1951>Template:Cite book</ref>
- F. m. nigripecta to be distributed in Tibet and Kashmir;<ref name=ems1951/>
- F. m. ferruginea occurring from southwestern Turkestan and the Kopet Dag mountains to Afghanistan and Balochistan.<ref name=ems1951/>
Since 2017, the Cat Classification Task Force of the Cat Specialist Group recognises only two subspecies as valid taxa, namely:<ref name=CatSG2017>Template:Cite journal</ref>
- O. m. manul syn. O. m. ferrugineus in the western and northern part of Central Asia from Iran to Mongolia;
- O. m. nigripectus in the Himalayas from Kashmir to Bhutan.
PhylogenyEdit
Phylogenetic analysis of the nuclear DNA in tissue samples from all Felidae species revealed that the evolutionary radiation of the Felidae began in Asia during the late Miocene around Template:Mya.<ref name=Johnson2006>Template:Cite journal</ref><ref name=Werdelin2010>Template:Cite book</ref> Analysis of mitochondrial DNA of all Felidae species indicates a radiation at around Template:Mya.<ref name=Li_al2016>Template:Cite journal</ref> The Pallas's cat is estimated to have genetically diverged from a common ancestor with the genus Prionailurus between Template:Mya based on analysis of nuclear DNA.<ref name=Johnson2006/> Based on analysis of mitochondrial DNA, it diverged Template:Mya from a common ancestor with Felis.<ref name=Li_al2016/>
CharacteristicsEdit
Template:Multiple image The Pallas's cat's fur is light grey with pale yellowish-ochre or pale yellowish-reddish hues.<ref name=Hodgson1842/> Some hair tips are white and some blackish. Its fur is greyer and denser with fewer markings visible in winter than in the summer.<ref name=Geptner1972>Template:Cite book</ref>Template:Rp The forehead and top of the head are light grey with small black spots. It has two black zigzag lines on the cheeks running from the corner of the eyes to the jaw joints.<ref name=Hodgson1842/> Its chin, whiskers, lower and upper lips are white.<ref name=Geptner1972/>Template:Rp It has narrow black stripes on the back, consisting of five to seven dark transversal lines across the lower back.<ref name=Hodgson1842/> Its grey tail has seven narrow black rings and a black tip.<ref name=Geptner1972/>Template:Rp The underfur is Template:Cvt long and 19 μm thick, and the guard hairs up to Template:Cvt long and 93 μm thick on the back. Its fur is soft and dense with up to Template:Cvt.<ref name=Geptner1972/>Template:Rp
The Pallas's cat's ears are grey with a yellowish tinge on the back and a darker rim, but with whitish hair in front and in the ear pinnae. Its rounded ears are set low on the side, such that it can peer over an object and show only a relatively small part of the head above the eyes without depressing the ears. This can give its face a look of ferocity and unrest.<ref name=Pocock1907/> Its eyes are encircled by white. The iris is yellowish, and its pupils contract to small circular disks in sunlight.<ref name=Pocock1907/>Template:Rp Among the Felinae, it shares this trait of round pupils with Puma, Herpailurus and Acinonyx species.<ref>Template:Cite book</ref>
The Pallas's cat is about the size of a domestic cat (Felis catus).<ref name=Hodgson1842/> Its stocky posture with the long and dense fur make it appear stout and plush. Its head-to-body is Template:Cvt long with a Template:Cvt long tail. It weighs Template:Cvt.<ref name=WCoW>Template:Cite book</ref> Its body is stout, and its skull is rounded with a short nasal bone, an enlarged cranial part and rounded zygomatic arches. Its orbits are large and directed forward. Its legs are short with short and sharp retractile claws.<ref name=Ognev1930>Template:Cite journal</ref>
The skull of males is Template:Cvt long and Template:Cvt wide at the base. Females have a Template:Cvt long and Template:Cvt wide skull.<ref name=Geptner1972/>Template:Rp The lower carnassial teeth are powerful, and the upper carnassials are short and massive. The first pair of upper premolars is absent. The dental formula is Template:DentalFormula.<ref name=Ognev1930/> It has a bite force at the canine tip of 155.4 newtons and a bite force quotient at the canine tip of 113.8.<ref>Template:Cite journal</ref>
The mitochondrial genome of the Pallas's cat consists of 16,672 base pairs containing 13 protein-coding, 22 transfer RNA and two ribosomal RNA genes and one non-coding RNA control region.<ref>Template:Cite journal</ref>
Distribution and habitatEdit
The Pallas's cat's range extends from the Caucasus eastward to Central Asia, Mongolia and adjacent parts of Dzungaria and the Tibetan Plateau. It inhabits montane shrublands and grasslands, rocky outcrops, scree slopes and ravines in areas, where the continuous snow cover is below Template:Cvt.<ref name=Geptner1972/> In the southwestern part of its range, the habitat of the Pallas's cat is affected by cold and dry winters, and moderate to low rainfall in warm summers.<ref name=Moqanaki_al2019>Template:Cite journal</ref> The typical vegetation in this part consists of small shrubs, sagebrush (Artemisia), Festuca and Stipa grasses.<ref name=Geptner1972/> In the central part of its range, it inhabits hilly landscapes, high plateaus and intermontane valleys that are covered by dry steppe or semi-desert vegetation, such as low shrubs and xerophytic grasses.<ref name=Barashkova_al2019>Template:Cite journal</ref> The continental climate in this region exhibits a range of Template:Cvt between the highest and lowest air temperatures, dropping to Template:Cvt in winter.<ref name=Geptner1972/>Template:Rp
The Greater Caucasus region is considered climatically suitable for the Pallas's cat.<ref name=Moqanaki_al2019/> In Armenia, an individual was killed near Vedi in the mountains of Ararat Province in the late 1920s.<ref name=Ognev1930/> In January 2020, an individual was sighted about Template:Cvt farther north in Tavush Province; the habitat at this location transitions from semi-desert to montane steppe at an elevation of about Template:Cvt.<ref>Template:Cite journal</ref> Records in Azerbaijan are limited to a Pallas's cat skin found in Karabakh and a sighting of an individual in Julfa District, both in the late 20th century.<ref name=Aghili_al2008>Template:Cite journal</ref>
On the Iranian Plateau, two Pallas's cats were encountered near the Aras River in northwestern Iran before the 1970s.<ref name=Geptner1972/> In the area, an individual was captured at an elevation of about Template:Cvt near Azarshahr in East Azerbaijan Province in 2008.<ref name=Aghili_al2008/> In the same year, a camera trap recorded a Pallas's cat on the southern slopes of the central Alborz Mountains in Khojir National Park shortly after heavy snowfall.<ref>Template:Cite journal</ref> Farther east in the Alborz Mountains, an individual was recorded among rocks at an elevation of Template:Cvt in 2016.<ref>Template:Cite journal</ref> In the Aladagh and Kopet Dag Mountains, the Pallas's cat was recorded inside and in the vicinity of protected areas.<ref name=Farhadinia_al2016>Template:Cite journal</ref> In the south of the Zagros Mountains, an individual was caught in a corral used by transhumant pastoralists in Abadeh County in 2012. The surrounding area consists of rocky steppe habitat dominated by mountain almond (Prunus scoparia), Astragalus and Artemisia.<ref>Template:Cite journal</ref> In the Hindu Kush, a Pallas's cat was observed sunbathing at the fringe of a rocky high-elevation plain near Dasht-e Nawar in Afghanistan's Koh-i-Baba range in April 2007.<ref>Template:Cite report</ref> The Pallas's cat was also photographed multiple times in Bamyan Province between 2015 and 2017.<ref name=Moqanaki_al2019/> In Pakistan's Qurumber National Park in Gilgit-Baltistan, an individual was recorded on a ridge in a juniper dominated forest at Template:Cvt in July 2012.<ref>Template:Cite journal</ref>
In the Transcaspian Region, its presence was first reported in the Kopet Dag mountains and in the vicinity of the Tedzhen and Murghab Rivers in the late 19th century.<ref>Template:Cite journal</ref> In Turkmenistan's Sünt-Hasardag Nature Reserve, a camera trap recorded an individual in 2019. The Pallas's cat is allegedly also present in Köpetdag Nature Reserve.<ref>Template:Cite report</ref>
Historical records of the Pallas's cat are known in the Surxondaryo Region and Gissar Range along the border of Tajikistan and Uzbekistan.<ref name=Geptner1972/> In Kyrgyzstan, it is present at high elevations of Sarychat-Ertash State Nature Reserve and in the foothills of the Alay Range.<ref name=Barashkova_al2019/> In 2013, a dead female was found in a valley near Engilchek, Kyrgyzstan.<ref>Template:Cite journal</ref> In Kazakhstan, it inhabits the highlands and steppes of central and east Kazakhstan Region, the periphery of the Betpak-Dala Desert, the northern Balkhash District and the Tarbagatai Mountains.<ref name=Barashkova_al2019/>
In the South Siberian Mountains, it inhabits grasslands on the Ukok Plateau and in the Altai, Kuray and Saylyugem Mountains.<ref name=BS2011>Template:Cite journal</ref> It is also present in Chagan-Uzun and Argut river basins, Mongun-Taiga, Uvs Lake Basin, Sayano-Shushenski Nature Reserve, Tunkinsky National Park, Lake Gusinoye basin and in the interfluves of the Selenga, Chikoy and Khilok rivers.<ref>Template:Cite journal</ref> In the eastern Sayan Mountains, its presence was documented for the first time in 1997.<ref>Template:Cite journal</ref> In Transbaikal, it inhabits montane steppes at elevations of Template:Cvt, where annual rainfall ranges from Template:Cvt.<ref>Template:Cite journal</ref> In 2013, an individual was observed on the Vitim Plateau.<ref>Template:Cite journal</ref>
The Pallas's cat inhabits the semi-desert steppe of Ikh Nartiin Chuluu Nature Reserve in Mongolia.<ref>Template:Cite journal</ref> In Khustain Nuruu National Park and Gobi Gurvansaikhan National Park, it prefers rocky and rugged habitats that provides cover and camouflage.<ref name=Ross_al2012>Template:Cite journal</ref><ref>Template:Cite journal</ref> On the Tibetan plateau, two Pallas's cats were observed in undulating alpine meadow amidst plateau pika (Ochotona curzoniae) colonies at Template:Cvt in western China's Qumarlêb County in 2001. One of them swam across an irrigation channel.<ref>Template:Cite journal</ref> In Gêrzê County, an individual was sighted in desert steppe habitat at an elevation of Template:Cvt in 2005.<ref>Template:Cite journal</ref> In 2011, the Pallas's cat was photographed in an alpine meadow in the core area of Sanjiangyuan National Nature Reserve.<ref>Template:Cite journal</ref> In Ruoergai, it was observed at several places in habitat that was frequented by pastoralists and their livestock herds.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>
The presence of the Pallas's cat in the Himalayas was first reported in Ladakh's Indus valley in 1991.<ref>Template:Cite journal</ref> In Changthang Wildlife Sanctuary, Pallas's cats were sighted close by riverbanks at elevations of Template:Cvt in 2013 and 2015.<ref>Template:Cite journal</ref> In Gangotri National Park, a Pallas's cat was photographed in rocky alpine scrub at Template:Cvt in 2019.<ref>Template:Cite journal</ref> In Sikkim, an individual was observed on a rocky slope at an elevation of Template:Cvt in the vicinity of Tso Lhamo Lake in 2007.<ref>Template:Cite journal</ref> In December 2012, the Pallas's cat was recorded for the first time in the Nepal Himalayas. It was photographed in the upper Marshyangdi river valley in alpine pastures at elevations of Template:Cvt and Template:Cvt in Annapurna Conservation Area.<ref>Template:Cite journal</ref> In Shey-Phoksundo National Park, Pallas's cat scat was detected at Template:Cvt in 2016, the globally highest record to date.<ref name=Werhahn_al2018>Template:Cite journal</ref> In January 2012, it was recorded for the first time in Bhutan, namely in rolling hills dominated by glacial outwash and alpine steppe vegetation in Wangchuck Centennial National Park.<ref>Template:Cite report</ref> In autumn 2012, it was also photographed at an elevation of Template:Cvt in Jigme Dorji National Park.<ref>Template:Cite journal</ref> In 2019, scat samples of two individuals were found in Sagarmatha National Park, providing the first genetic evidence of the cat's presence in the eastern Himalayas.<ref>Template:Cite journal</ref>
Behaviour and ecologyEdit
The Pallas's cat is solitary.<ref name=Geptner1972/> Of nine Pallas's cat kittens observed in captivity, only the two males scent marked by spraying urine.<ref>Template:Cite book</ref>
The Pallas's cat uses caves, rock crevices and marmot burrows as shelter.<ref name=Geptner1972/>Template:Rp In central Mongolia, 29 Pallas's cats were fitted with radio collars between June 2005 and October 2007. They used 101 dens during this time, including 39 winter dens, 42 summer dens and 20 dens for raising kittens. The summer and winter dens usually had one entrance with a diameter of Template:Cvt. They resided in the summer dens for 2–21 days, and in the winter dens for 2–28 days. Summer and maternal dens were close to rocky habitats with little direct sunlight, whereas winter dens were closer to ravines.<ref name=Ross_al2010>Template:Cite journal</ref> The home ranges of 16 females varied from Template:Cvt. The home ranges of nine males varied from Template:Cvt and overlapped those of one to four females and partly also those of other males. The sizes of their home ranges decreased in winter.<ref name=Ross_al2012/>
In an unprotected area in central Mongolia, Pallas's cats were mainly crepuscular between May and August, but active by day from September to November.<ref>Template:Cite journal</ref> Pallas's cats recorded in four study areas in the western Mongolian Altai mountains were also active during the day, but with a lower frequency at sites where livestock was present.<ref>Template:Cite journal</ref>
Hunting and dietEdit
The Pallas's cat is a highly specialised predator of small mammals, which it catches by stalking or ambushing near exits of burrows. It also pulls out rodents with its paws from shallow burrows. In the Altai Mountains, remains of long-tailed ground squirrel (Urocitellus undulatus), flat-skulled shrew (Sorex roboratus), Pallas's pika (Ochotona pallasi) and bird feathers were found near breeding burrows of Pallas's cats. In Transbaikal, it preys on Daurian pika (Ochotona dauurica), steppe pika (O. pusilla), Daurian ground squirrel (Spermophilus dauricus) and young of red-billed chough (Pyrrhocorax pyrrhocorax).<ref name=Geptner1972/> Scat samples of the Pallas's cat collected in the bufferzone of Khustain Nuruu National Park in central Mongolia contained foremost remains of Daurian pika, Mongolian gerbil (Meriones unguiculatus), Mongolian silver vole (Alticola semicanus) and remains of passerine birds, beetles and grasshoppers.<ref>Template:Cite journal</ref> Brandt's vole (Lasiopodomys brandtii) dominated in the diet of Pallas's cats in Mongolia's Sükhbaatar Province after the irruptive growth of this vole population during 2017 to 2020.<ref>Template:Cite journal</ref> Scat found in Shey-Phoksundo National Park contained remains of pika species and of woolly hare (Lepus oiostolus).<ref name=Werhahn_al2018/> Remains of a cypriniform fish were found in Pallas's cat scat in Gongga Mountain Nature Reserve.<ref>Template:Cite journal</ref>
Reproduction and life cycleEdit
The female is sexually mature at the age of about one year.<ref name=Mellen1989>Template:Cite thesis</ref> She is in estrus for 26 to 42 hours.<ref name=Schauenberg1978>Template:Cite journal</ref> Gestation lasts 66 to 75 days.<ref name=Mellen1989/><ref name=Schauenberg1978/>
A captive male Pallas's cat housed under natural lighting conditions showed increased aggressive and territorial behaviour at the onset of the breeding season, lasting from September to December. Its blood contained three times more testosterone than in the non-breeding season, and its ejaculate was more concentrated with more normal sperm forms and a higher motility of sperm.<ref name=Swanson1996>Template:Cite journal</ref>
In the wild, the female gives birth to a litter of two to six kittens between the end of April and late May. The newborn kittens' fur is fuzzy, and their eyes are closed until the age of about two weeks.<ref name=Geptner1972/>Template:Rp A newborn male kitten born in a zoo weighed Template:Cvt, measured Template:Cvt and had a Template:Cvt long tail.<ref name=Mellen1989/>
In central Mongolia, seven females with kittens were observed using 20 dens for 4–60 days. Their maternal dens were either among rocks, or in former burrows of the Tarbagan marmot (Marmota sibirica), and had at least two entrances.<ref name=Ross_al2010/> In Iran, a Pallas's cat was observed using cavities of aged Greek juniper (Juniperus excelsa) as breeding dens for a litter of four kittens.<ref>Template:Cite journal</ref>
Two-month-old kittens weigh Template:Cvt, and their fur gradually grows longer. They start hunting at the age of about five months and reach adult size by the age of six to seven months.<ref name=Geptner1972/>Template:Rp
ThreatsEdit
In China, Mongolia and Russia, the Pallas's cat was once hunted for its fur in large numbers of more than 10,000 skins annually. In China and the former Soviet Union, hunting of the Pallas's cat decreased in the 1970s when it became legally protected. Mongolia exported 9,185 skins in 1987, but international trade has ceased since 1988.<ref>Template:Cite book</ref> However, domestic trade of its skins and body parts for medicinal purposes continues in the country, and it may be hunted throughout the year.<ref name=Wingard2006>Template:Cite report</ref>
Cases of herding dogs killing Pallas's cats were reported in Iran, Kazakhstan and the Altai Republic.<ref name=Farhadinia_al2016/><ref name=BS2011/><ref name=Barashkova_al2019/> Pallas's cats have also fallen victim in traps set for small mammals in Kazakhstan and in the Altai Republic. In Transbaikal, the Pallas's cat is threatened by poaching. In Mongolia, the use of the rodenticide bromadiolone in the frame of rodent control measures in the early 21st century poisoned the prey base of carnivores and raptors.<ref name=Barashkova_al2019/> In the Sanjiangyuan region of the Tibetan Plateau, Template:Convert of grassland was poisoned between 2005 and 2009, leading to an estimated loss of Template:Convert of pika biomass.<ref>Template:Cite journal</ref> The Pallas's cat may be negatively affected by habitat fragmentation due to mining and infrastructure projects.<ref name=iucn />
ConservationEdit
On the IUCN Red List, the Pallas's cat is classified as Least Concern since 2020 because of its wide-spread range and assumed large global population. It is listed in CITES Appendix II. Hunting it is prohibited in all range countries except Mongolia. Since 2009, it is legally protected in Afghanistan, where all hunting and trade with its body parts is banned.<ref name=iucn /> On the Mongolian Red List of Mammals, it is listed as Near Threatened since 2006.<ref name=Wingard2006/> In China, it is listed as Endangered.<ref>Template:Cite book</ref> In Turkmenistan, it is proposed to be listed as Critically Endangered due to the scarcity of contemporary records.<ref>Template:Cite book</ref>
In captivityEdit
Between 1951 and 1979, the Beijing Zoo kept 16 Pallas's cats, but they lived for less than three years.<ref>Template:Cite journal</ref> In 1984, the Pallas's cat was designated as a priority species for captive breeding of the American Association of Zoos and Aquariums's Species Survival Plan.<ref>Template:Cite book</ref> Almost half of the kittens born in member zoos died within the first 30 days, reaching the highest mortality rate in captivity of any small wild cat.<ref>Template:Cite journal</ref>
Zoos in the former Soviet Union received most of the wild-caught Pallas's cats from the Transbaikal region and a few from Mongolia. Moscow Zoo initiated a studbook for the Pallas's cat in 1997. Since 2004, the Pallas's cat international studbook has been managed by the Royal Zoological Society of Scotland, which also coordinates the captive breeding program for the Pallas's cat within the European Endangered Species Programme. As of 2018, 177 Pallas's cats were kept in 60 zoos in Europe, Russia, North America and Japan.<ref>Template:Cite journal</ref>
In 2011, a female Pallas's cat was artificially inseminated for the first time with semen from the male at the Cincinnati Zoo. After 69 days, she gave birth to four kittens, of which one was stillborn.<ref>Template:Cite book</ref>
In popular cultureEdit
The Pallas's cat is featured in a Russian Internet meme known as "Pet the cat" introduced in 2008; the meme is a picture of a Pallas's cat that invites the reader to pet it in the image's caption.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> In 2012, the Pallas's cat became the mascot of Moscow Zoo.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>
ReferencesEdit
External linksEdit
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