Clouded leopard

This article is about the animal species. For the infantry fighting vehicle, see CM-32 Armoured Vehicle.

Clouded leopard
Clouded leopard in Kaeng Krachan National Park
Conservation status
Vulnerable  (IUCN 3.1) [1]
CITES Appendix I (CITES) [1]
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Family:	Felidae
Subfamily:	Pantherinae
Genus:	Neofelis
Species:	N. nebulosa [2]
Binomial name
Neofelis nebulosa [2] (Griffith, 1821)
Distribution of the clouded leopard [1]

The clouded leopard (Neofelis nebulosa), also called mainland clouded leopard, is a wild cat inhabiting dense forests from the foothills of the Himalayas through Northeast India and Bhutan to mainland Southeast Asia into South China. It was first described in 1821 on the basis of a skin of an individual from China. The clouded leopard has large dusky-grey blotches and irregular spots and stripes reminiscent of clouds. Its head-and-body length ranges from 68.6 to 108 cm (27.0 to 42.5 in) with a 61 to 91 cm (24 to 36 in) long tail. It uses its tail for balancing when moving in trees and is able to climb down vertical tree trunks head first. It rests in trees during the day and hunts by night on the forest floor.

The clouded leopard is the sister taxon to other pantherine cats, having genetically diverged 9.32 to 4.47 million years ago. Today, the clouded leopard is locally extinct in Singapore, Taiwan, and possibly also in Hainan Island and Vietnam. The wild population is believed to be in decline with fewer than 10,000 adults and no more than 1,000 in each subpopulation. It has therefore been listed as Vulnerable on the IUCN Red List since 2008. The population is threatened by large–scale deforestation and commercial poaching for the wildlife trade. Its body parts are offered for decoration and clothing, though it is legally protected in most range countries.

The clouded leopard has been kept in zoological gardens since the early 20th century. Captive breeding programs were initiated in the 1980s. In captivity, the clouded leopard has an average lifespan of 11 years.

Taxonomy and phylogeny

Felis nebulosa was proposed by Edward Griffith in 1821 who first described a skin of a clouded leopard that was brought alive from Guangdong in China to the menagerie at Exeter Exchange in London. ^[3] Felis macrosceloides proposed by Brian Houghton Hodgson in 1841 was a clouded leopard specimen from Nepal. ^{[4] [5]} Felis brachyura proposed by Robert Swinhoe in 1862 was a clouded leopard skin from Taiwan. ^[6] The generic name Neofelis was proposed by John Edward Gray in 1867 who subordinated all three to this genus. ^[7] At present, N. nebulosa is considered a monotypic species due to lack of evidence for subspeciation. ^[8]

Felis diardi proposed by Georges Cuvier in 1823 was based on a clouded leopard skin from Java. ^[9] It was considered a clouded leopard subspecies by Reginald Innes Pocock in 1917. ^[10] In 2006, it was identified as a distinct Neofelis species, the Sunda clouded leopard. ^{[11] [12]} Populations in Taiwan and Hainan Island are considered to belong to the mainland clouded leopard. ^[8]

Phylogeny

Skulls of clouded leopard and Panthera species were analysed morphologically in the 1960s. Results indicate that the clouded leopard forms an evolutionary link between the Pantherinae and the Felinae. ^[13] Phylogenetic analysis of the nuclear DNA in tissue samples from all Felidae species revealed that the evolutionary radiation of the Felidae began in the Miocene around 14.45 to 8.38 million years ago in Asia. ^{[14] [15]} Analysis of mitochondrial DNA of all Felidae species indicates a radiation at 16.76 to 6.46 million years ago. ^[16] The clouded leopard is the sister taxon to all other members of the Pantherinae, diverging 9.32 to 4.47 million years ago, based on analysis of their nuclear DNA. ^[14] The clouded leopard from mainland Asia reached Borneo and Sumatra via a now submerged land bridge probably during the Pleistocene, when populations became isolated during periods of global cooling and warming. Genetic analysis of hair samples of the clouded leopard and its sister species the Sunda clouded leopard (N. diardi) indicates that they diverged 2.0–0.93 million years ago. ^[11]

Phylogenetic relationships of the clouded leopard as derived through analysis of

nuclear DNA: [14]
Felidae Felinae Pantherinae Panthera Clouded leopard

combined mitochondrial and nuclear DNA: [11]
Felidae Felinae Pantherinae Panthera  Neofelis  Sunda clouded leopard Mainland clouded leopard

Characteristics

Face

Skull

The clouded leopard's fur is of a dark grey or ochreous ground-color, 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. Its legs are short and stout, and paws broad. Females are slightly smaller than males. ^[17]

Its hyoid bone is ossified, making it possible to purr. Its pupils contract into vertical slits. ^[18] Irises are brownish yellow to grayish green. Melanistic clouded leopards are uncommon. It has rather short limbs compared to the other big cats. Its hind limbs are longer than its front limbs to allow for increased jumping and leaping capabilities. Its ulnae and radii are not fused, which also contributes to a greater range of motion when climbing trees and stalking prey. 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. ^[19] Its shoulder height varies from 50 to 55 cm (20 to 22 in). ^[20]

Its skull is long and low with strong occipital and sagittal crests. The canine teeth are exceptionally long, the upper being about three times as long as the basal width of the socket. The first premolar is usually absent. ^[17] The upper pair of canines measure 4 cm (1.6 in) or longer. ^[19] It has a bite force at the canine tip of 544.3 Newton and a bite force quotient at the canine tip of 122.4. ^[21] The clouded leopard is often referred to as a "modern-day sabre-tooth" because it has the largest canines in proportion to its body size. ^[18]

Distribution and habitat

Clouded leopard at Aizawl, Mizoram, India

The clouded leopard occurs from the Himalayan foothills in Nepal, Bhutan and India to Myanmar, southeastern Bangladesh, Thailand, Peninsular Malaysia and to south of the Yangtze River in China. It is locally extinct in Singapore and Taiwan. ^[1]

Clouded leopards were found in Nepal in 1987 and 1988, having previously been presumed to be extinct in the country. ^[22] Since then, the clouded leopard has been recorded in Shivapuri Nagarjun National Park and in Annapurna Conservation Area. ^{[23] [24]} Between 2014 and 2015, it was also recorded in Langtang National Park at an elevation range of 1,823–3,498 m (5,981–11,476 ft). ^[25]

In India, it occurs in the states of Sikkim, northern West Bengal, Tripura, Mizoram, Manipur, Assam, Nagaland and Arunachal Pradesh, as well as in the Meghalaya subtropical forests. ^{[26] [27] [28] [29]} In Pakke Tiger Reserve, a clouded leopard was photographed in semi-evergreen forest at an elevation of 144 m (472 ft). ^[30] In Sikkim, clouded leopards were photographed by camera traps at elevations of 2,500–3,720 m (8,200–12,200 ft) between April 2008 and May 2010 in the Khangchendzonga Biosphere Reserve. ^[31] In Manas National Park, 16 individuals were recorded during a survey in November 2010 to February 2011. ^[32] Between January 2013 and March 2018, clouded leopards were also recorded in Dampa Tiger Reserve, Eaglenest Wildlife Sanctuary and Singchung-Bugun Village Community Reserve, in Meghalaya's Nongkhyllem National Park and Balpakram-Baghmara landscape. ^[33]

In Bhutan, it was recorded in Royal Manas National Park, Jigme Singye Wangchuck National Park, Phibsoo Wildlife Sanctuary, Jigme Dorji National Park, Phrumsengla National Park, Bumdeling Wildlife Sanctuary and several non-protected areas. ^{[34] [35]} In Bangladesh, it was recorded in Sangu Matamuhari in the Chittagong Hill Tracts in 2016. ^[36] In Myanmar, it was recorded by camera traps for the first time in the hill forests of Karen State in 2015. ^[37]

In Thailand, it inhabits relatively open, dry tropical forest in Huai Kha Khaeng Wildlife Sanctuary and closed-forest habitats in Khao Yai National Park. ^{[38] [39] [40]} In Laos, it was recorded in Nam Et-Phou Louey National Protected Area in dry evergreen and semi-evergreen forests. ^[41] In Cambodia, it was recorded in deciduous dipterocarp forest in Phnom Prich Wildlife Sanctuary between 2008 and 2009, ^[42] and in Central Cardamom Mountains National Park, Southern Cardamom National Park, Botum Sakor National Park and Phnom Samkos Wildlife Sanctuary between 2012 and 2016. ^[43] In Peninsular Malaysia, it was recorded in Taman Negara National Park, Ulu Muda Forest, Pasoh Forest Reserve, Belum-Temengor, Temengor Forest Reserve and in a few linkages between 2009 and 2015. ^[44]

The last confirmed record of a Formosan clouded leopard dates to 1989, when the skin of a young individual was found in the Taroko National Park. ^[45] It was not recorded during an extensive camera trapping survey conducted from 1997 to 2012 in more than 1,450 sites inside and outside Taiwanese protected areas. ^[46]

Behaviour and ecology

In the San Antonio Zoo and Aquarium, Texas, US

The clouded leopard is a solitary cat. Early accounts depict it as a rare, secretive, arboreal, and nocturnal inhabitant of dense primary forest. ^[18]

It is one of the most talented climbers among the cats. Captive clouded leopards 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. ^[13] They are capable of supination and can hang down from branches only by bending their hind paws and their tail around them. They can jump up to 1.2 m (3 ft 11 in) high. ^[13] They use trees as daytime rest sites, but also spend time on the ground when hunting at night. Captive clouded leopards have been observed to scent mark by spraying urine and rubbing their heads on prominent objects. ^[19] Their vocalisations include a short high-pitched meow call, a loud crying call, both emitted when a cat is trying to locate another one over a long or short distance; they prusten and raise their muzzle when meeting each other in a friendly manner; when aggressive, they growl with a low-pitched sound and hiss with exposed teeth and wrinkled nose. ^[47]

Radio-collared clouded leopards were foremost active by night but also showed crepuscular activity peaks. ^[39] Clouded leopards recorded in northeast India were most active in the late evening after sunset. ^[33]

Home ranges have only been estimated in Thailand:

• Four individuals were radio-collared in Phu Khieo Wildlife Sanctuary from April 2000 to February 2003. Home ranges of two females were 25.7 km² (9.9 sq mi) and 22.9 km² (8.8 sq mi), and of two males 29.7 km² (11.5 sq mi) and 49.1 km² (19.0 sq mi). ^[39]
• Two individuals were radio-collared during a study from 1997 to 1999 in the Khao Yai National Park. The home range of one female was 39.4 km² (15.2 sq mi), of the one male 42 km² (16 sq mi). Both individuals had a core area of 2.9 km² (1.1 sq mi). ^[48]

In 2016, clouded leopards were detected in the forest complex of Khlong Saeng Wildlife Sanctuary and Khao Sok National Park during camera trapping surveys; 15 individuals were identified in a core zone of 200 km² (77 sq mi) with population density estimated at 5.06 individuals per 100 km² (39 sq mi); but only 12 individuals were identified in an edge zone of 297 km² (115 sq mi), which is more disturbed by humans, with density estimated at 3.13 individuals per 100 km² (39 sq mi). ^[49]

Hunting and diet

When hunting, the clouded leopard stalks its prey or waits for the prey to approach. After making and feeding on a kill, it usually retreats into trees to digest and rest. Its prey includes both arboreal and terrestrial vertebrates. ^[19] Pocock presumed that it is adapted for preying upon herbivorous mammals of considerable bulk because of its powerful build, long canines and the deep penetration of its bites. ^[17] In Thailand, clouded leopards have been observed preying on southern pig-tailed macaque (Macaca nemestrina), ^[50] Indian hog deer (Axis porcinus), Bengal slow loris (Nycticebus bengalensis), Asiatic brush-tailed porcupine (Atherurus macrourus), Malayan pangolin (Manis javanica) and Berdmore's ground squirrel (Menetes berdmorei). ^[39] Known prey species in China include barking deer (Muntiacus sp.) and pheasants. ^[51] In northern Peninsular Malaysia, a male clouded leopard was photographed while carrying a binturong (Arctictis binturong) in its jaws. ^[52]

Reproduction and life cycle

Clouded leopard cub

Both males and females average 26 months at first reproduction. The female is in estrus for about six days, with her estrous cycle lasting about 30 days. ^[53] In the wild, mating usually occurs between December and March. The pair mates multiple times over the course of several days. The male grasps the female by the neck who responds with vocalization. Occasionally, he also bites her during courtship and is very aggressive during sexual encounters. Females can bear one litter each year. The male is not involved in raising the cubs. ^[19]

The female gives birth to a litter of one to five, mostly three cubs, after a gestation period of 93 ± 6 days. ^[53] Cubs are born with closed eyes and weigh from 140 to 280 g (4.9 to 9.9 oz). Their spots are solid dark, rather than dark rings. Their eyes open after about 10 days. They are active within five weeks and fully weaned at around three months of age. They attain the adult coat pattern at around six months and become independent after around 10 months. ^[19]

Captive clouded leopards have an average lifespan of 11 years. ^[54] One individual has lived to be almost 17 years old. ^[55]

The generation length of the clouded leopard is about seven years. ^[56]

Threats

A coat made of clouded leopard skin. Poaching for illegal trade of skin is one of the main threats to the clouded leopard.

Clouded leopard require larger areas of intact forest than are present in many parts of their range. ^[57] They are threatened by habitat loss following large–scale deforestation and commercial poaching for the wildlife trade. ^[58] 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, despite the protected status of clouded leopards in Myanmar. Some markets are located near Myanmar's borders with China and Thailand and are used to facilitate cross-border smuggling. ^[59]

In Nepal, 27 cases of clouded leopard body parts were discovered between November 1988 and March 2020 in nine districts of the country, comprising at least 51 individual clouded leopards. In 17 of these cases, the poachers and traders were arrested. ^[60]

Conservation

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

The clouded leopard is listed in CITES Appendix I. Hunting is banned in Bangladesh, China, India, Malaysia, Myanmar, Nepal, Taiwan, Thailand and Vietnam. ^[53] These bans, however, are poorly enforced in India, Malaysia and Thailand. ^{[58] [ failed verification ]}

In the United States, the clouded leopard is listed as endangered under the Endangered Species Act, prohibiting trade in live animals or body parts. ^[2]

International Clouded Leopard Day is celebrated each year on 4 August since 2018 in zoos and conservation organizations all over the world. ^[61]

In captivity

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

Clouded leopards have been kept in zoos since the early 20th century. The international studbook was initiated in the 1970s. Coordinated breeding programs were started in the 1980s and encompass the European Endangered Species Programme, the Species Survival Plan, and the Indian Conservation Breeding Programme. As of 2014, 64 institutions keep clouded leopards. ^[62]

Early captive-breeding programs involving clouded leopards were not successful, largely due to ignorance of their courtship behaviour. Males have the reputation of being aggressive towards females. For breeding success, it has been deemed extremely important that male and female clouded leopards are compatible. ^[19] Introducing pairs at a young age gives them opportunities to bond and breed successfully. Facilities breeding clouded leopards need to provide the female with a secluded, off-exhibit area. ^[20] There has been some recent captive breeding success using artificial insemination with cubs successfully born in 1992, 2015 and 2017. ^[63]

A study on morbidity and mortality rate of 271 captive clouded leopards across 44 zoos in Europe, Asia and Australia showed that 17% of them died because of respiratory disease, 12% due to maternal neglect and starvation, 10% from generalized infectious disease, 10% from digestive diseases, and 10% from trauma. ^[64]

In March 2011, two breeding females at the Nashville Zoo at Grassmere gave birth to three cubs, which were raised by zookeepers. Each cub weighed 0.23 kg (0.5 lb). ^[65] In June 2011, two cubs were born at the Point Defiance Zoo & Aquarium. The breeding pair was brought from the Khao Kheow Open Zoo in Thailand in an ongoing education and research exchange program. ^[66] Four cubs were born at Nashville Zoo in 2012. ^[67] In May 2015, four cubs were born in Point Defiance Zoo & Aquarium. ^[68]

In culture

The clouded leopard is the state animal of the Indian state of Meghalaya. ^[69] In the 1970s, the print of Rama Samaraweera's painting Clouded leopard was a best-seller in the US. ^[70]

See also

• List of largest cats

References

1. Gray, T.; Borah, J.; Coudrat, C.N.Z.; Ghimirey, Y.; Giordano, A.; Greenspan, E.; Petersen, W.; Rostro-García, S.; Shariff, M. & Wai-Ming, W. (2021). "Neofelis nebulosa". IUCN Red List of Threatened Species . 2021: e.T14519A198843258. doi: 10.2305/IUCN.UK.2021-2.RLTS.T14519A198843258.en . Retrieved 19 November 2021.
2. Wozencraft, W. C. (2005). "Species Neofelis nebulosa". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 545–546. ISBN   978-0-8018-8221-0. OCLC   62265494.
3. Griffith, E. (1821). "The Chinese, or tortoiseshell Tiger. Felis nebulosa". General and particular descriptions of the vertebrated animals arranged comfortably to the modern discoveries and improvements in zoology. London: Baldwin, Cradock & Joy. p. 37. Archived from the original on 10 February 2023. Retrieved 19 November 2020.
4. Hodgson, B. H. (1841). "Classified Catalogue of Mammals of Nepal". Calcutta Journal of Natural History and Miscellany of the Arts and Sciences in India. IV: 284–294. Archived from the original on 10 February 2023. Retrieved 10 May 2018.
5. Hodgson, B. H. (1853). "Felis macrosceloides". Proceedings of the Zoological Society of London. I. Mammalia: Plate XXXVIII.
6. Swinhoe R. (1862). "On the mammals of the Island of Formosa (China)". Proceedings of the Zoological Society of London. 30: 347–365. doi:10.1111/j.1469-7998.1862.tb06539.x.
7. Gray, J. E. (1867). "Notes on the skulls of cats (Felidae)". Proceedings of the Zoological Society of London. 35: 258–277.
8. Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; Werdelin, L.; Wilting, A.; Yamaguchi, N.; Abramov, A. V.; Christiansen, P.; Driscoll, C.; Duckworth, J. W.; Johnson, W.; Luo, S.-J.; Meijaard, E.; O’Donoghue, P.; Sanderson, J.; Seymour, K.; Bruford, M.; Groves, C.; Hoffmann, M.; Nowell, K.; Timmons, Z. & Tobe, S. (2017). "A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group" (PDF). Cat News (Special Issue 11): 64–66. Archived (PDF) from the original on 17 January 2020. Retrieved 20 December 2020.
9. Cuvier, G. (1823). "Des ossemens des grands Felis". Recherches sur les ossemens fossiles : où l'on rétablit les caractères de plusieurs animaux dont les révolutions du globe ont détruit les espèces. Volume IV: Les ruminans et les carnassiers fossiles. Paris: G. Dufour & E. d'Ocagne. pp. 407−456. Archived from the original on 10 February 2023. Retrieved 17 November 2020.
10. Pocock, R. I. (1917). "The classification of existing Felidae". The Annals and Magazine of Natural History: Including Zoology, Botany, and Geology. 8th. 20 (119): 329–350. doi:10.1080/00222931709487018.
11. 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. Bibcode:2006CBio...16.2371B. doi:10.1016/j.cub.2006.08.066. PMC   5618441 . PMID   17141620.
12. 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. Bibcode:2006CBio...16.2377K. doi: 10.1016/j.cub.2006.10.066 . PMID   17141621. S2CID   6838593.
13. 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.
14. 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 . 311 (5757): 73–77. Bibcode:2006Sci...311...73J. doi:10.1126/science.1122277. PMID   16400146. S2CID   41672825. Archived from the original on 4 October 2020. Retrieved 11 September 2019.
15. Werdelin, L.; Yamaguchi, N.; Johnson, W. E. & O'Brien, S. J. (2010). "Phylogeny and evolution of cats (Felidae)". In Macdonald, D. W. & Loveridge, A. J. (eds.). Biology and Conservation of Wild Felids. Oxford, UK: Oxford University Press. pp. 59–82. ISBN   978-0-19-923445-5. Archived from the original on 25 September 2018. Retrieved 16 September 2019.
16. Li, G.; Davis, B. W.; Eizirik, E. & Murphy, W. J. (2016). "Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae)". Genome Research. 26 (1): 1–11. doi:10.1101/gr.186668.114. PMC   4691742 . PMID   26518481.
17. Pocock, R.I. (1939). "Genus Neofelis Gray. The Clouded Leopard". The Fauna of British India, including Ceylon and Burma. Vol. Mammalia. – Volume 1. London: Taylor and Francis, Ltd. pp. 247–253. This article incorporates text from this source, which is in the public domain .
18. Guggisberg, C.A.W. (1975). "Clouded leopard Neofelis nebulosa (Griffiths, 1821)". Wild cats of the World. New York: Taplinger Publishing. pp. 125–130. ISBN   978-0-8008-8324-9.
19. Sunquist, M. & Sunquist, F. (2002). "Clouded leopard Neofelis nebulosa (Griffiths, 1821)". Wild cats of the World. Chicago: University of Chicago Press. pp. 278–284. ISBN   978-0-226-77999-7. Archived from the original on 10 February 2023. Retrieved 26 November 2020.
20. Clouded Leopard SSP (2000). Clouded leopard (Neofelis nebulosa) Husbandry Guidelines (PDF). Silver Spring: Association of Zoos and Aquariums. Archived (PDF) from the original on 17 October 2013. Retrieved 27 January 2013.
21. Christiansen, P. & Wroe, S. (2007). "Bite forces and evolutionary adaptations to feeding ecology in carnivores". Ecology. 88 (2): 347–358. doi:10.1890/0012-9658(2007)88[347:bfaeat]2.0.co;2. PMID   17479753.
22. Dinerstein, E. & Mehta, J. N. (1989). "The clouded leopard in Nepal". Oryx. 23 (4): 199–201. doi: 10.1017/s0030605300023024 . S2CID   86485712.
23. Pandey, B. P. (2012). "Clouded leopard in Shivapuri Nagarjun National Park, Nepal". Cat News (57): 24–25.
24. Ghimirey, Y.; Acharya, R.; Adhikary, B.; Werhahn, G. & Appel, A. (2013). "Clouded leopard camera-trapped in the Annapurna Conservation Area, Nepal". Cat News (58): 25.
25. Can, Ö. E.; Yadav, B. P.; Johnson, P. J.; Ross, J.; D’Cruze, N. & Macdonald, D. W. (2020). "Factors affecting the occurrence and activity of clouded leopards, common leopards and leopard cats in the Himalayas". Biodiversity and Conservation. 29 (3): 839–851. Bibcode:2020BiCon..29..839C. doi: 10.1007/s10531-019-01912-7 .
26. Choudhury, A. U. (1992). "The Clouded Leopard in Assam". Oryx. 27 (1): 51–53. doi: 10.1017/s0030605300023966 .
27. Choudhury, A. U. (1996). "The clouded leopard". Cheetah. 35 (1–2): 13–18.
28. Choudhury, A. (1997). "The clouded leopard in Manipur and Nagaland". Journal of the Bombay Natural History Society. 94 (2): 389–391.
29. Choudhury, A. U. (2003). "The cats in North East India". Cat News. 39: 15–19.
30. Borah, J.; Sharma, T.; Lyngdoh, S. & Tapi, T. (2010). "First photograph of a clouded leopard at Pakke Tiger Reserve, India". Cat News (52): 24–25.
31. 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 (3): 257–268. doi:10.1515/mamm.2011.023. S2CID   52205824.
32. Borah, J.; Sharma, T.; Das, D.; Rabha, N.; Kakati, N.; Basumatary, A.; Ahmed, M. F. & Vattakaven, J. (2014). "Abundance and density estimates for common leopard Panthera pardus and clouded leopard Neofelis nebulosa in Manas National Park, Assam, India". Oryx. 48 (1): 149–155. doi: 10.1017/s0030605312000373 .
33. Mukherjee, S.; Singh, P.; Silva, A.; Ri, C.; Kakati, K.; Borah, B.; Tapi, T.; Kadur, S.; Choudhary, P.; Srikant, S.; Nadig, S.; Navya, R.; Björklund, R. & Ramakrishnan, U. (2019). "Activity patterns of the small and medium felid (Mammalia: Carnivora: Felidae) guild in northeastern India". Journal of Threatened Taxa. 11 (4): 13432–13447. doi: 10.11609/jott.4662.11.4.13432-13447 .
34. Tempa, T.; Hebblewhite, M.; Mills, L. S.; Wangchuk, T. R.; Norbu, N.; Wangchuk, T.; Nidup, T.; Dendup, P.; Wangchuk, D.; Wangdi, Y. & Dorji, T. (2013). "Royal Manas National Park, Bhutan: a hot spot for wild felids". Oryx. 47 (2): 207–210. doi: 10.1017/s0030605312001317 .
35. Penjor, U.; Macdonald, D. W.; Wangchuk, S.; Tandin, T. & Tan, C. K. W. (2018). "Identifying important conservation areas for the clouded leopard Neofelis nebulosa in a mountainous landscape: Inference from spatial modeling techniques". Ecology and Evolution. 8 (8): 4278–4291. Bibcode:2018EcoEv...8.4278P. doi:10.1002/ece3.3970. PMC   5916301 . PMID   29721297.
36. Petersen, W. J.; Savini, T. & Ngoprasert, D. (2020). "Strongholds under siege: Range-wide deforestation and poaching threaten mainland clouded leopards (Neofelis nebulosa)". Global Ecology and Conservation. 24: e01354. Bibcode:2020GEcoC..2401354P. doi: 10.1016/j.gecco.2020.e01354 .
37. Saw Sha Bwe Moo; Froese, G. Z. L. & Gray, T. N. E. (2017). "First structured camera-trap surveys in Karen State, Myanmar, reveal high diversity of globally threatened mammals". Oryx. 52 (3): 537. doi: 10.1017/S0030605316001113 .
38. 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. doi:10.1017/s0266467400005034. S2CID   84998140.
39. Grassman, L. I. Jr.; 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. doi:10.1644/1545-1542(2005)086<0029:eotsfi>2.0.co;2. S2CID   86640520.
40. Austin, S. C.; Tewes, M. E.; Grassman, L. I. Jr. & Silvy, N. J. (2007). "Ecology and conservation of the leopard cat Prionailurus bengalensis and clouded leopard Neofelis nebulosa in Khao Yai National Park, Thailand". Acta Zoologica Sinica. 53: 1–14.
41. Rasphone, A.; Kamler, J.F. & Macdonald, D.W. (2020). "Temporal partitioning by felids, dholes and their potential prey in northern Laos". Mammal Research. 65 (4): 679–689. doi: 10.1007/s13364-020-00524-9 .
42. Gray, T.N. & Phan, C. (2011). "Habitat preferences and activity patterns of the larger mammal community in Phnom Prich Wildlife Sanctuary, Cambodia". The Raffles Bulletin of Zoology. 59 (2): 311–318.
43. Gray, T.N.; Billingsley, A.; Crudge, B.; Frechette, J.L.; Grosu, R.; Herranz-Muñoz, V.; Holden, J.; Keo, O.; Kong, K.; Macdonald, D. & Neang, T. (2017). "Status and conservation significance of ground-dwelling mammals in the Cardamom Rainforest Landscape, southwestern Cambodia". Cambodian Journal of Natural History. 2017 (1): 38–48.
44. Tan, C. K. W.; Rocha, D. G.; Clements, G. R.; Brenes-Mora, E.; Hedges, L.; Kawanishi, K.; Mohamad, S. W.; Rayan, D. M.; Bolongon, G.; Moore, J. & Wadey, J. (2017). "Habitat use and predicted range for the mainland clouded leopard Neofelis nebulosa in Peninsular Malaysia". Biological Conservation. 206: 65–74. Bibcode:2017BCons.206...65T. doi:10.1016/j.biocon.2016.12.012.
45. Anonymous (1996). "The mystery of the Formosan clouded leopard". Cat News. 24: 16.
46. Chiang, P. J.; Pei, K. J. C.; Vaughan, M. R.; Li, C. F.; Chen, M. T.; Liu, J. N.; Lin, C. Y.; Lin, L. K. & Lai, Y. C. (2015). "Is the clouded leopard Neofelis nebulosa extinct in Taiwan, and could it be reintroduced? An assessment of prey and habitat". Oryx. 49 (2): 261–269. doi: 10.1017/S003060531300063X .
47. Fazio, J. M. (2010). Behavioral Assessment of the Clouded Leopard (Neofelis nebulosa); A Comparative Study of Reproductive Success (MSc). Fairfax, VA: George Mason University.
48. Austin, S. C. & Tewes, M. E. (1999). "Ecology of the clouded leopard in Khao Yai National Park, Thailand". Cat News (31): 17–18.
49. Petersen, W. J.; Steinmetz, R.; Sribuarod, K. & Ngoprasert, D. (2020). "Density and movements of mainland clouded leopards (Neofelis nebulosa) under conditions of high and low poaching pressure". Global Ecology and Conservation. 23: e01117. Bibcode:2020GEcoC..2301117P. doi: 10.1016/j.gecco.2020.e01117 .
50. Davies, R.G. (1990). "Sighting of a clouded leopard (Neofelis nebulosa) in a troop of pigtail macaques (Macaca nemestrina) in Khao Yai National Park, Thailand". Siam Society Natural History Bulletin. 38: 95–96.
51. Feng, L.; Lin, L.; Zhang, L.; Wang, L.; Wang, B.; Luo, A.; Yang, S.; Smith, J. L. D.; Luo, S. J. & Zhang, L. (2008). "Evidence of wild tigers in southwest China – a preliminary survey of the Xishuangbanna National Nature Reserve". Cat News (48): 4–6.
52. Lam, W.Y.; Hedges, L. & Clements, G.R. (2014). "First record of a clouded leopard predating on a binturong". Cat News (60): 16.
53. Nowell, K. & Jackson, P. (1996). "Clouded Leopard". Wild Cats: Status Survey and Conservation Action Plan. Gland: IUCN/SSC Cat Specialist Group. Archived from the original on 3 June 2007.
54. 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. Archived from the original on 2 August 2020. Retrieved 9 April 2020.
55. "Clouded leopard". The Aspinall Foundation. n.d. Archived from the original on 18 April 2014.
56. Pacifici, M.; Santini, L.; Di Marco, M.; Baisero, D.; Francucci, L.; Grottolo Marasini, G.; Visconti, P. & Rondinini, C. (2013). "Generation length for mammals". Nature Conservation. 5 (5): 87–94. doi: 10.3897/natureconservation.5.5734 .
57. 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. 6 (16): 1–13. doi: 10.1186/1472-6785-6-16 . PMC   1654139 . PMID   17092347.
58. Nowell, K. (2007). Asian big cat conservation and trade control in selected range States: evaluating implementation and effectiveness of CITES Recommendations (PDF) (Report). Kuala Lumpur: TRAFFIC. Archived (PDF) from the original on 3 October 2020. Retrieved 9 October 2010.
59. Shepherd, C. R. & Nijman, V. (2008). The wild cat trade in Myanmar (PDF) (Report). Petaling Jaya, Selangor, Malaysia: TRAFFIC Southeast Asia. Archived (PDF) from the original on 23 January 2021. Retrieved 20 December 2020.
60. Ghimirey, Y. & Acharya, R. (2020). "Clouded Leopard Neofelis nebulosa (Grifth, 1821) (Mammalia: Carnivora: Felidae) in illegal wildlife trade in Nepal". Journal of Threatened Taxa. 12 (16): 17229–17234. doi:10.11609/jott.6504.12.16.17229-17234. S2CID   230543392.
61. "International Clouded Leopard Day". The Aspinall Foundation. Archived from the original on 5 August 2019. Retrieved 5 August 2019.
62. Wildlife Institute of India (2014). "Status in captivity". National Studbook of Clouded leopard (Neofelis nebulosa). TR. No. -2014/ 010 (PDF). New Delhi: Central Zoo Authority. p. 5. Archived (PDF) from the original on 11 May 2019. Retrieved 11 May 2019.
63. "First Clouded Leopard Cub Produced with Cryopreserved Semen". Smithsonian's National Zoo. 2017. Archived from the original on 22 October 2020. Retrieved 1 November 2020.
64. Thorel, M.; Pignon, C.; Pascal A.; Donnelly, T.M. & Rivière, J. (2020). "Clouded Leopard (Neofelis nebulosa) morbidity and mortality in captive-bred populations: A comprehensive retrospective study of medical data from 271 individuals in European, Asian, and Australian zoos". Journal of Zoo and Wildlife Medicine. 51 (1): 150–158. doi:10.1638/2019-0048. PMID   32212558. S2CID   213184824.
65. "Cute! Clouded Leopard Cubs Born at Nashville Zoo". LiveScience . 2011. Archived from the original on 3 October 2014.
66. "Welcome Our New Clouded Leopard Cubs". Point Defiance Zoo & Aquarium . 2011. Archived from the original on 27 June 2011.
67. "The Nashville Zoo celebrates two sets of clouded leopard cubs". Associated Press. 2012. Archived from the original on 3 June 2013.
68. "Four extremely rare leopards born in US zoo". RTÉ. 2015. Archived from the original on 24 May 2015. Retrieved 23 May 2015.
69. "State Animals, Birds, Trees and Flowers of India". ENVIS RP on Forestry and Forest Related Livelihoods, Ministry of Environment & Forests, Government of India. 2011. Archived from the original on 8 March 2016. Retrieved 7 June 2017.
70. "DZG inspired acclaimed artist". Archived from the original on 5 March 2018. Retrieved 6 September 2016.

External links

• "Mainland clouded leopard Neofelis nebulosa". IUCN Species Survival Commission Cat Specialist Group.
• "Clouded Leopard Neofelis nebulosa". National Geographic . Archived from the original on 24 February 2012.
• "Taiwan's clouded leopard extinct: zoologists". Focus Taiwan. 2013.
• "Clouded Leopard: Facing Dark Clouds of Extinction". About Animals. 20 June 2016.