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.
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
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]
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
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
terrestrialvertebrates.[19]
Pocock presumed that it is adapted for preying upon
herbivorousmammals 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
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]
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
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]
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
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]
^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.
^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.
^Hodgson, B. H. (1853).
"Felis macrosceloides". Proceedings of the Zoological Society of London. I. Mammalia: Plate XXXVIII.
^Gray, J. E. (1867). "Notes on the skulls of cats (Felidae)". Proceedings of the Zoological Society of London. 35: 258–277.
^
abKitchener, 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.
^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.
^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.
ISBN978-0-19-923445-5.
Archived from the original on 25 September 2018. Retrieved 16 September 2019.
^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.
S2CID84998140.
^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.
^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.
^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.
^Fazio, J. M. (2010). Behavioral Assessment of the Clouded Leopard (Neofelis nebulosa); A Comparative Study of Reproductive Success (MSc). Fairfax, VA: George Mason University.
^Austin, S. C. & Tewes, M. E. (1999). "Ecology of the clouded leopard in Khao Yai National Park, Thailand". Cat News (31): 17–18.
^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.
^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.
^Lam, W.Y.; Hedges, L. & Clements, G.R. (2014). "First record of a clouded leopard predating on a binturong". Cat News (60): 16.
^
abcNowell, 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.
^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.
^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.
S2CID230543392.
^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.
PMID32212558.
S2CID213184824.