Theclouded leopard (Neofelis nebulosa), also calledmainland clouded leopard, is a wildcat inhabiting dense forests from the foothills of theHimalayas throughNortheast India andBhutan tomainland Southeast Asia intoSouth China. It was first described in 1821 on the basis of a skin of an individual fromChina. 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 theforest floor.
The clouded leopard is thesister taxon to otherpantherine cats, having genetically diverged 9.32 to 4.47 million years ago. Today, the clouded leopard islocally extinct inSingapore,Taiwan, and possibly also inHainan Island andVietnam. The wild population is believed to be in decline, with fewer than 10,000 adults and no more than 1,000 in any subpopulation. It has therefore been listed asVulnerable on theIUCN Red List since 2008. The population is threatened by large-scaledeforestation and commercialpoaching 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 inzoological 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 byGeorges Cuvier in 1823 was based on a clouded leopard skin from Java.[9]It was considered a clouded leopardsubspecies byReginald Innes Pocock in 1917.[10] In 2006, it was identified as a distinctNeofelis species, theSunda clouded leopard.[11][12] Populations in Taiwan and Hainan Island are considered to belong to the mainland clouded leopard.[8]
Skulls of clouded leopard andPanthera species were analysedmorphologically in the 1960s. Results indicate that the clouded leopard forms anevolutionary link between thePantherinae and theFelinae.[13]Phylogenetic analysis of thenuclear DNA in tissue samples from all Felidae species revealed that theevolutionary radiation of the Felidae began in theMiocene around14.45 to 8.38 million years ago in Asia.[14][15] Analysis ofmitochondrial DNA of all Felidae species indicates a radiation at16.76 to 6.46 million years ago.[16]The clouded leopard is thesister taxon to all other members of the Pantherinae, diverging9.32 to 4.47 million years ago, based on analysis of their nuclear DNA.[14]The clouded leopard from mainland Asia reachedBorneo andSumatra via a now submergedland bridge probably during thePleistocene, when populations became isolated during periods of global cooling and warming.Genetic analysis of hair samples of the clouded leopard and itssister 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]
Itshyoid 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. Itsulnae andradii 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 strongoccipital andsagittal crests. Thecanine 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 abite force at thecanine tip of 544.3Newton and abite force quotient at the canine tip of 122.4.[21]The clouded leopard is often referred to as a "modern-daysabre-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 theTaroko 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]
The clouded leopard is a solitary cat. Early accounts depict it as a rare, secretive,arboreal, andnocturnal inhabitant of denseprimary 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 ofsupination 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 toscent mark byspraying urine andrubbing 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; theyprusten 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 showedcrepuscular 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 wereradio-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 theKhao 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 ofKhlong Saeng Wildlife Sanctuary andKhao Sok National Park during camera trapping surveys; 15 individuals were identified in a core zone of 200 km2 (77 sq mi) withpopulation 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]
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 andterrestrialvertebrates.[19]Pocock presumed that it is adapted for preying uponherbivorousmammals 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 onsouthern pig-tailed macaque (Macaca nemestrina),[50]Indian hog deer (Axis porcinus),Asiatic brush-tailed porcupine (Atherurus macrourus),Malayan pangolin (Manis javanica) andBerdmore's ground squirrel (Menetes berdmorei).[39] In India, a clouded leopard also preyed on aBengal slow loris (Nycticebus bengalensis).[51] Known prey species in China includebarking deer (Muntiacus sp.) andpheasants.[52]In northern Peninsular Malaysia, a male clouded leopard was photographed while carrying abinturong (Arctictis binturong) in its jaws.[53]
Both males and females average 26 months at first reproduction. The female is inestrus for about six days, with herestrous cycle lasting about 30 days.[54] 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 onelitter 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 agestation period of 93 ± 6 days.[54] 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.[55]One individual has lived to be almost 17 years old.[56]
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.[58] They are threatened by habitat loss following large–scaledeforestation and commercial poaching for thewildlife trade.[59] 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.[60]
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.[61]
A clouded leopard resting in a tree trunk at theToronto Zoo
The clouded leopard is listed inCITES Appendix I. Hunting is banned in Bangladesh, China, India,Malaysia, Myanmar, Nepal, Taiwan, Thailand and Vietnam.[54] These bans, however, are poorly enforced in India, Malaysia and Thailand.[59][failed verification]
Clouded leopards have been kept in zoos since the early 20th century. The internationalstudbook was initiated in the 1970s. Coordinated breeding programs were started in the 1980s and encompass theEuropean Endangered Species Programme, theSpecies Survival Plan, and the Indian Conservation Breeding Programme. As of 2014, 64 institutions keep clouded leopards.[63]
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 usingartificial insemination with cubs successfully born in 1992, 2015 and 2017.[64]
A study onmorbidity andmortality rate of 271 captive clouded leopards across 44 zoos in Europe, Asia and Australia showed that 17% of them died because ofrespiratory disease, 12% due to maternal neglect and starvation, 10% from generalized infectious disease, 10% fromdigestive diseases, and 10% from trauma.[65]
In March 2011, two breeding females at theNashville Zoo at Grassmere gave birth to three cubs, which were raised by zookeepers. Each cub weighed 0.23 kg (0.5 lb).[66] In June 2011, two cubs were born at thePoint Defiance Zoo & Aquarium. The breeding pair was brought from theKhao Kheow Open Zoo in Thailand in an ongoing education and research exchange program.[67] Four cubs were born at Nashville Zoo in 2012.[68] In May 2015, four cubs were born in Point Defiance Zoo & Aquarium.[69]
The clouded leopard is the state animal of the Indian state ofMeghalaya.[70] In the 1970s, the print ofRama Samaraweera's paintingClouded leopard was a best-seller in the US.[71]
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