Clouded leopard

This article is about the Asian feline species. For the Taiwanese infantry fighting vehicle, see CM-32 Armoured Vehicle.
Clouded leopard
A clouded leopard at the San Antonio Zoo and Aquarium.
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Felidae
Subfamily: Pantherinae
Genus: Neofelis
Species: N. nebulosa[2]
Binomial name
Neofelis nebulosa
(Griffith, 1821)
Clouded leopard range
Synonyms

Felis macrocelis
Felis marmota

The clouded leopard (Neofelis nebulosa) is a wild cat found from the Himalayan foothills through mainland Southeast Asia into China, and has been classified as Vulnerable in 2008 by the International Union for Conservation of Nature (IUCN). Its total population size is suspected to be fewer than 10,000 mature individuals, with a decreasing population trend, and no single population numbering more than 1,000 adults.[1]

Taxonomy

Two cladograms proposed for the Panthera lineage

The scientific name of the clouded leopard is Neofelis nebulosa. It is one of the two members of the genus Neofelis, and is classified under the family Felidae.[3] The other member of this genus is the Sunda clouded leopard (N. diardi), that was considered a subspecies of the clouded leopard till as recently as 2006.[4][5] The clouded leopard was first described by British zoologist Edward Griffith in 1821.[3]

The clouded leopard is part of the Panthera lineage, one of the eight lineages of Felidae. This lineage comprises the species of Panthera and Neofelis. The Neofelis species diverged first from the lineage, followed by the snow leopard.[6] Genetic analysis of hair samples of the two Neofelis species indicates that they diverged 1.4 million years ago, after having used a now submerged land bridge to reach Borneo and Sumatra from mainland Asia.[4] Subsequent branching in the lineage is disputed. [6] Broadly, two different cladograms have been proposed for the Panthera lineage.[7][8][9][10] The clouded leopard is considered to form an evolutionary link between the big cats and the small cats.[11] It represents the smallest of the big cats, but despite its name, it is not closely related to the leopard.[12]

Characteristics

Close-up of face

The fur of clouded leopards is of a dark grey or ochreous ground-colour, often largely obliterated by black and dark dusky-grey blotched pattern. There are black spots on the head, and the ears are black. Partly fused or broken-up stripes run from the corner of the eyes over the cheek, from the corner of the mouth to the neck, and along the nape to the shoulders. Elongated blotches continue down the spine and form a single median stripe on the loins. Two large blotches of dark dusky-grey hair on the side of the shoulders are each emphasized posteriorly by a dark stripe, which passes on to the foreleg and breaks up into irregular spots. The flanks are marked by dark dusky-grey irregular blotches bordered behind by long, oblique, irregularly curved or looped stripes. These blotches yielding the clouded pattern suggest the English name of the cat. The underparts and legs are spotted, and the tail is marked by large, irregular, paired spots. Females are slightly smaller than males.[12]

Their irises are usually either greyish-green or brownish-yellow in color. Their legs are short and stout, with broad paws. They have rather short limbs compared to the other big cats, but their hind limbs are longer than their front limbs to allow for increased jumping and leaping capabilities. Their ulnae and radii are not fused, which also contributes to a greater range of motion when climbing trees and stalking prey.[13]

Melanistic clouded leopards are uncommon. Clouded leopards weigh between 11.5 and 23 kg (25 and 51 lb). Females vary in head-to-body length from 68.6 to 94 cm (27.0 to 37.0 in), with a tail 61 to 82 cm (24 to 32 in) long. Males are larger at 81 to 108 cm (32 to 43 in) with a tail 74 to 91 cm (29 to 36 in) long.[14] Their shoulder height varies from 50 to 55 cm (20 to 22 in).[15]

They have exceptionally long, piercing canine teeth, the upper being about three times as long as the basal width of the socket.[12] The upper pair of canines may measure 4 cm (1.6 in) or longer.[14] They are often referred to as a “modern-day saber tooth” because they have the largest canines in proportion to their body size, matching the tiger in canine length. The first premolar is usually absent, and they also have a very distinct long and slim skull with well-developed occipital and sagittal crests to support the enlarged jaw muscles.[13]

Etymology

The scientific name of the genus Neofelis is a composite of the Greek word νεο- meaning "new", and the Latin word feles meaning "cat", so it literally means "new cat."[16][17]

Distribution and habitat

Clouded leopard at Aizawl, Mizoram, India

Clouded leopards occur from the Himalayan foothills in Nepal and India to Myanmar, Bhutan, Thailand, Peninsular Malaysia, Indochina, and in China south of the Yangtze River. They are regionally extinct in Taiwan.[1] Clouded leopards prefer open- or closed-forest habitats to other habitat types.[18] They have been reported from relatively open, dry tropical forest in Myanmar and in Thailand.[19]

In Bangladesh its presence is uncertain[20] However, there have been some recent reports of clouded leopard in Chittagong Hill Tracts of south-eastern Bangladesh. In 2009 a few clouded leopards were sighted. Among these one cub was caught by the local indigenous people.[21] Also there are some uncertain reports of clouded leopard sightings in the south-eastern and north-eastern parts of the country.[22]

In India, they occur in Assam, northern West Bengal, Sikkim, Arunachal Pradesh, Manipur, Meghalaya, Mizoram, Nagaland, and Tripura.[23][24][25][26] In the Himalayas, they were camera-trapped at altitudes of 2,500–3,720 m (8,200–12,200 ft) between April 2008 and May 2010 in the Khangchendzonga Biosphere Reserve, Sikkim.[27]

Clouded leopards were thought to be extinct in Nepal since the late 1860s. But in 1987 and 1988, four individuals were found in the central part of the country, close to Chitwan National Park and in the Pokhara Valley. These findings extended their known range westward, suggesting they are able to survive and breed in degraded woodlands that previously harboured moist subtropical semideciduous forest.[28] Since then, individuals have been recorded in the Shivapuri Nagarjun National Park and in the Annapurna Conservation Area.[29][30]

Distribution of subspecies

At present, these three subspecies of Neofelis nebulosa are recognized:[31]

Ecology and behavior

Clouded leopards are the most talented climbers among the cats. In captivity, they have been observed to climb down vertical tree trunks head first, and hang on to branches with their hind paws bent around branchings of tree limbs. They are capable of supination and can even hang down from branches only by bending their hind paws and their tail around them. When jumping down, they keep hanging on to a branch this way until the very last moment. They can climb on horizontal branches with their back to the ground, and in this position make short jumps forward. When balancing on thin branches, they use their long tails to steer. They can easily jump up to 1.2 m (3.9 ft) high.[11]

Clouded leopards have been observed to scent mark in captivity by urine-spraying and head-rubbing on prominent objects. Presumably such habits are used to mark their territory in the wild, although the size of their home ranges is unknown. Like other big cats, they do not appear able to purr, but they otherwise have a wide range of vocalisations, including mewing, hissing, growling, moaning, and snorting. When communicating, two individuals will emit low snorting sounds that are called prusten when approaching each other in a friendly manner. They also use long-call communication used over large distances, which could either be a type of mating call between different territories or a warning call to other cats encroaching on other territories.[13] Apart from information stemming from observations of captive clouded leopards, little is known of their natural history and behavior in the wild. Early accounts depict them as rare, secretive, arboreal, and nocturnal denizens of dense primary forest. More recent observations suggest they may not be as arboreal and nocturnal as previously thought. They may use trees as daytime rest sites, but also spend a significant proportion of time on the ground. Some daytime movement has been observed, suggesting they are not strictly nocturnal but crepuscular. However, the time of day when they are active depends on their prey and the level of human disturbance.[14]

They live a solitary lifestyle, resting in trees during the day and hunting at night. When hunting, clouded leopards either come down from their perches in the trees and stalk their prey or lie and wait for the prey to come to them. After making a kill and eating, they usually retreat to the trees to digest and rest.[34]

Their partly nocturnal and far-ranging behaviour, their low densities, and because they inhabit densely vegetated habitats and remote areas makes the counting and monitoring of clouded leopards extremely difficult. Consequently, little is known about their behaviour and status. Available information on their ecology is anecdotal, based on local interviews and a few sighting reports.[35]

Home ranges have only been estimated in Thailand:

Little is known of the diet of clouded leopards. Their prey includes both arboreal and terrestrial vertebrates.[14] Pocock presumed they are adapted for preying upon herbivorous mammals of considerable bulk because of their powerful build and the deep penetration of their bites, attested by their long canines.[12] Confirmed prey species include hog deer, slow loris, brush-tailed porcupine, Malayan pangolin and Indochinese ground squirrel.[18] Known prey species in China include barking deer and pheasants.[37]

Reproduction

Both males and females average 26 months at first reproduction. Mating usually occurs during December and March. The males tend to be very aggressive during sexual encounters and have been known to bite the female on the neck during courtship, severing her vertebrae. With this in mind, male and female compatibility has been deemed extremely important when attempting breeding in captivity. The pair will meet and mate multiple times over the course of several days. The male grasps the female by the neck and the female responds with vocalization that encourages the male to continue. The male then leaves and is not involved in raising the kittens.[13] Estrus last six days on average, estrous cycle averages 30 days. After a gestation period of 93 ± 6 days, females give birth to a litter of one to five, most often three cubs.[38]

Initially, the young are blind and helpless, much like the young of many other cats, and weigh from 140 to 280 g (4.9 to 9.9 oz). Unlike adults, the kittens' spots are "solid" — completely dark rather than dark rings. The young can see within about 10 days of birth, are active within five weeks, and are fully weaned at around three months of age. They attain the adult coat pattern at around six months, and probably become independent after around 10 months. Females are able to bear one litter each year.[14] The mother is believed to hide her kittens in dense vegetation while she goes to hunt, though little concrete evidence supports this theory, since their lifestyle is so secretive.[13]

In captivity, they have an average lifespan of 11 years. One individual has lived to be almost 17 years old.[39][40]

Threats

Many of the remaining forest areas are too small to ensure the long-term persistence of clouded leopard populations.[35] They are threatened by habitat loss following large–scale deforestation and commercial poaching for the wildlife trade. Skins, claws, and teeth are offered for decoration and clothing, bones and meat as substitute for tiger in traditional Asian medicines and tonics, and live animals for the pet trade. Few poaching incidents have been documented, but all range states are believed to have some degree of commercial poaching. In recent years, substantial domestic markets existed in Indonesia, Myanmar, and Vietnam.[41][42]

In Myanmar, 301 body parts of at least 279 clouded leopards, mostly skins and skeletons, were observed in four markets surveyed between 1991 and 2006. Three of the surveyed markets are situated on international borders with China and Thailand, and cater to international buyers, although clouded leopards are completely protected under Myanmar's national legislation. Effective implementation and enforcement of CITES is considered inadequate.[43]

Conservation

A clouded leopard resting atop a tree trunk at the Toronto Zoo

Neofelis nebulosa is listed in CITES Appendix I and protected over most of its range. Hunting is banned in Bangladesh, China, India, Malaysia, Myanmar, Nepal, Taiwan, Thailand, and Vietnam. It is not legally protected outside Bhutan's protected areas. Hunting is regulated in Laos. No information about its protection status is available from Cambodia.[38] These bans, however, are poorly enforced in India, Malaysia, and Thailand.[41]

In the United States, the clouded leopard is listed as endangered under the Endangered Species Act, further prohibiting trade in the animals or any parts or products made from them.[2]

In captivity

A clouded leopard at the Feline Conservation Center, Rosamond, California

Early captive-breeding programs involving clouded leopards were not very successful, largely due to ignorance of courtship activity among them in the wild. Experience has taught keepers that introducing pairs of clouded leopards at a young age gives opportunities for the pair to bond and breed successfully. Males have the reputation of being aggressive towards females. Facilities breeding clouded leopards need to provide the female a secluded, off-exhibit area.[15] Modern breeding programs involve carefully regulated introductions between prospective mating pairs, and take into account the requirements for enriched enclosures. Stimulating natural behavior by providing adequate space to permit climbing minimizes stress. This, combined with a feeding program that fulfills the proper dietary requirements, has promoted more successful breeding in recent years.

In Tripura, a national park was set up in the Sipahijola Wildlife Sanctuary where clouded leopards are kept in enclosures in a zoological park.[44]

In March 2011, two breeding females at the Nashville Zoo at Grassmere in Nashville, Tennessee, gave birth to three cubs, which are being raised by zookeepers. Each cub weighed 0.5 lb (0.23 kg).[45] In June 2011, two cubs were born at the Point Defiance Zoo & Aquarium in Tacoma, Washington. The breeding pair was brought from the Khao Kheow Open Zoo in Thailand in an ongoing education and research exchange program.[46] Four cubs were born at the Nashville Zoo in 2012.[47] On May 22, 2015, four more cubs were born at Tacoma's Point Defiance Zoo & Aquarium. The cubs were the fourth litter born to Chai Li and her mate Nah Fun.[48]

As of December 2011, 222 clouded leopards are believed to exist in zoos.[49]

In culture

The Rukai people of Taiwan considered the hunting of clouded leopards a taboo.[50]

In the media

A clouded leopard is the antagonist of the fantasy animated feature film Rugrats Go Wild. Chrissie Hynde provides the voice of the animal.

References

  1. 1 2 3 4 Sanderson, J., Khan, J. A., Grassman, L. & Mallon, D. P. (2008). "Neofelis nebulosa". IUCN Red List of Threatened Species. Version 2015.2. International Union for Conservation of Nature.
  2. 1 2 Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 545. ISBN 978-0-8018-8221-0. OCLC 62265494.
  3. 1 2 Wilson, D.E.; Reeder, D.M., eds. (2005). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 546. ISBN 978-0-8018-8221-0. OCLC 62265494.
  4. 1 2 Buckley-Beason, V.A.; Johnson, W.E.; Nash, W.G.; Stanyon, R.; Menninger, J.C.; Driscoll, C.A.; Howard, J.; Bush, M.; Page, J.E.; Roelke, M.E.; Stone, G.; Martelli, P.P.; Wen, C.; Ling, L.; Duraisingam, R.K.; Lam, P.V.; O'Brien, S.J. (2006). "Molecular evidence for species-level distinctions in clouded leopards". Current Biology 16 (23): 2371–2376. doi:10.1016/j.cub.2006.08.066.
  5. ↑ Kitchener, A.C.; Beaumont, M.A.; Richardson, D. (2006). "Geographical variation in the clouded leopard, Neofelis nebulosa, reveals two species". Current Biology 16 (23): 2377–2383. doi:10.1016/j.cub.2006.10.066. PMID 17141621.
  6. 1 2 Turner, A. (1987). "New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora)". Annals of the Transvaal Museum 34: 319–347.
  7. ↑ Johnson, W.E.; 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 (New York) 311 (5757): 73–77. PMID 16400146.
  8. ↑ Werdelin, L.; Yamaguchi, N.; Johnson, W.E.; O'Brien, S.J. (2010). "Phylogeny and evolution of cats (Felidae)" (PDF). Biology and Conservation of Wild Felids: 59–82.
  9. ↑ Davis, B.W.; Li, G.; Murphy, W.J. (2010). "Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, Panthera (Carnivora: Felidae)" (PDF). Molecular Phylogenetics and Evolution 56 (1): 64–76. doi:10.1016/j.ympev.2010.01.036.
  10. ↑ Mazák, J.H.; Christiansen, P.; Kitchener, A.C.; Goswami, A. (2011). "Oldest known pantherine skull and evolution of the tiger". PLoS ONE 6 (10): e25483. doi:10.1371/journal.pone.0025483.
  11. 1 2 Hemmer, H. (1968). Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) II: Studien zur Ethologie des Nebelparders Neofelis nebulosa (Griffith 1821) und des Irbis Uncia uncia (Schreber 1775). [Researching the phylogenetic history of the Pantherinae II: Studies into the ethology of the clouded leopard Neofelis nebulosa and snow leopard Uncia uncia.] Veröffentlichungen der Zoologischen Staatssammlung München 12:155–247.
  12. 1 2 3 4 Pocock, R.I. (1939). The Fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1. Taylor and Francis, Ltd., London. Pp 247–253
  13. 1 2 3 4 5 McGinley, M.; Turnage, C. (2012). "Clouded Leopard". Encyclopedia of Earth Eds. Cutler J. Cleveland (Washington, D.C.: Environmental Information Coalition, National Council for Science and the Environment). Archived from the original on 2013-12-29.
  14. 1 2 3 4 5 Sunquist, M.; Sunquist, F. (2002). Wild cats of the World. Chicago: University of Chicago Press. pp. 278–284. ISBN 0-226-77999-8.
  15. 1 2 Clouded leopard SSP (2000). Clouded leopard (Neofelis nebulosa) Husbandry Guidelines. American Zoo and Aquarium Association.
  16. ↑ Perseus Digital Library. Greek Dictionary νεο Headword Search Result
  17. ↑ Perseus Digital Library. Latin Dictionary feles Headword Search Result
  18. 1 2 3 Grassman Jr., L. I.,Tewes, M. E., Silvy, N. J., Kreetiyutanont, K. (2005). "Ecology of three sympatric felids in a mixed evergreen forest in North-central Thailand". Journal of Mammalogy 86: 29–38.
  19. ↑ Rabinowitz, A. R., Walker, S. R. (1991). "The carnivore community in a dry tropical forest mosaic in Huai Kha Khaeng Wildlife Sanctuary, Thailand". Journal of Tropical Ecology 7: 37−47.
  20. ↑ "Clouded Leopard - IUCNRedlist.org".
  21. ↑ "Rare Clouded Leopard caught in Bangladesh BBC News.".
  22. ↑ "Introduction to wildlife of Bangladesh".
  23. ↑ Choudhury, A.U. (1996). The clouded leopard. Cheetal 35 (1-2): 13–18.
  24. ↑ Choudhury, A. (1997). "The clouded leopard in Manipur and Nagaland". Journal of the Bombay Natural History Society 94(2): 389–391.
  25. ↑ Choudhury, A. U. (2003). "The cats in North East India". Cat News 39: 15–19.
  26. ↑ Choudhury, A. U. (1992). The Clouded Leopard in Assam. Oryx 27 (1): 51–53.
  27. ↑ Sathykumar, S., Bashir, T., Bhattacharya, T., Poudyal, K. (2011). Assessing mammal distribution and abundance in intricate Eastern Himalayan habitats of Khangchendzonga, Sikkim, India. Mammalia 75: 257–268.
  28. ↑ Dinerstein, E. and J. N. Mehta (1989). The clouded leopard in Nepal. Oryx 23(4): 199–201.
  29. ↑ Pandey, B. P. (2012). "Clouded leopard in Shivapuri Nagarjun National Park, Nepal". Cat News 57: 24–25.
  30. ↑ Ghimirey, Y., Acharya, R., Adhikary, B., Werhahn, G. and A. Appel (2013). "Clouded leopard camera-trapped in the Annapurna Conservation Area, Nepal". Cat News 58: 25.
  31. ↑ Sunquist, M. E., Sunquist, F. (2009). Family Felidae (Cats) In: Wilson, D. E., Mittermeier, R. A. (eds.) Handbook of the Mammals of the World - Volume 1 Carnivores. Lynx Edicions in association with Conservation International and IUCN. ISBN 978-84-96553-49-1
  32. ↑ Anonymous. (1996). "The mystery of the Formosan clouded leopard". Cat News 24: 16.
  33. ↑ Chiang, P.-J. (2007). Ecology and conservation of Formosan clouded leopard, its prey, and other sympatric carnivores in southern Taiwan. PhD thesis submitted to the Virginia Polytechnic Institute and State University, Virginia.
  34. ↑ "Clouded leopard". A-Z Animals. Archived from the original on 2015-01-16.
  35. 1 2 Wilting, A., Fischer, F., Bakar, S.A., Linsenmair, K.E. (2006) Clouded leopards, the secretive top-carnivore of South-East Asian rainforests: their distribution, status and conservation needs in Sabah, Malaysia. BMC Ecology 2006, 6 (16): 1–13.
  36. ↑ Austin, S. C., Tewes, M. E. (1999). "Ecology of the clouded leopard in Khao Yai National Park, Thailand". Cat News 31: 17–18.
  37. ↑ Feng, L., Lin, L., Zhang, L., Wang, L., Wang, B., Luo A., Yang, S., Smith, J. L. D., Luo S. J. and Zhang, L. (2008). "Evidence of wild tigers in southwest China – a preliminary survey of the Xishuangbanna National Nature Reserve". Cat News 48: 4–6.
  38. 1 2 Nowell, K.; Jackson, P. (1996). "Clouded Leopard". Wild Cats: Status Survey and Conservation Action Plan. IUCN/SSC Cat Specialist Group. Archived from the original on 2007-06-03.
  39. ↑ Achariyo, L. N., Mishra, Ch. G. (1981). "Some notes on the longevity of two species of Indian wild cats in captivity". Journal of the Bombay Natural History Society 78: 155.
  40. ↑ "Clouded leopard". The Aspinall Foundation. n.d. Archived from the original on 2014-04-18.
  41. 1 2 Nowell, K. (2007). Asian big cat conservation and trade control in selected range States: evaluating implementation and effectiveness of CITES Recommendations. A TRAFFIC Report, June 2007
  42. ↑ "Basic Facts About Clouded Leopards". Defenders of Wildlife. Archived from the original on 2014-08-09.
  43. ↑ Shepherd, C. R., Nijman, V. (2008) The wild cat trade in Myanmar. TRAFFIC Southeast Asia, Petaling Jaya, Selangor, Malaysia.
  44. ↑ Anonymous (2007). Tripura to set up National Park for Clouded Leopard. OneIndia.com
  45. ↑ "Cute! Clouded Leopard Cubs Born at Nashville Zoo". LiveScience. 2011-04-06. Archived from the original on 2014-10-03.
  46. ↑ "Welcome Our New Clouded Leopard Cubs". Point Defiance Zoo & Aquarium. 2011-06-24. Archived from the original on 2011-06-27.
  47. ↑ "The Nashville Zoo celebrates two sets of clouded leopard cubs". Associated Press. March 29, 2012. Archived from the original on 2013-06-03.
  48. ↑ "Four extremely rare leopards born in US zoo". RTÉ. May 22, 2015.
  49. ↑ Musi, V. J. (2011). "Cats in Crisis". National Geographic 202 (6): foldout (between 90–91).
  50. ↑ Pei, K. (1999). Hunting System of the Rukai Tribe in Taiwan, Republic of China. Proceedings of the International Union of Game Biologists XXIV Congress, Thessaloniki, Greece.

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