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Lifespan, longevity, and ageing

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Maximum longevity: 19.8 years (captivity) Observations: One captive specimen lived 19.8 years (Richard Weigl 2005).
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Conservation Status

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Little is known about the population status of clouded leopards because actual population estimates are difficult to obtain. The chief threat for clouded leopard populations is habitat loss due to deforestation for agricultural purposes. Humans hunt clouded leopards for their pelts and teeth, as well as for use in traditional medicine and culinary trades. In a survey conducted by the IUCN in 1991 in southeastern China, clouded leopard pelts were common on the black market. The Taiwanese purchase most clouded leopard products and the Taiwanese subspecies of clouded leopards (Neofelis nebulosa brachyura) is thought to be extinct as a result. Trade of clouded leopard products has been prohibited by CITES since 1975. Laws now protect clouded leopards over the majority of their range. Hunting is strictly prohibited in Bangladesh, Brunei, China, India, Indonesia, Malaysia, Myanmar, Nepal, Taiwan, Thailand, and Vietnam; hunting is regulated in Laos. The IUCN lists clouded leopards as vulnerable, and they are also listed under the U.S. Endangered Species Act, which prohibits the trade of any part of the animal in the United States. Still, prohibition of hunting of clouded leopards does not necessarily decrease demand and pelts have been reported on sale in urban markets in Myanmar, Laos, Vietnam, Cambodia, Nepal, and Thailand. Clouded leopards face persecution by farmers who feel that their livestock is at risk. Populations have been fragmented by deforestation, increasing the susceptibility of the entire species to infectious disease and natural catastrophic events. Efforts have been made in Nepal, Malaysia, and Indonesia to establish national parks in order to sustain populations of clouded leopards. Unfortunately, due to their elusive nature and dense forest habitats, data on the numbers actually surviving in parks are limited and possibly inaccurate.

US Federal List: endangered

CITES: appendix i

State of Michigan List: no special status

IUCN Red List of Threatened Species: vulnerable

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Associations

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The main predators of clouded leopards are humans, who use dogs to track and corner them. For this reason, clouded leopards avoid humans and they are rarely found near human settlements. Clouded leopards share much of their range with tigers and leopards. In these shared areas clouded leopards seem to have a more arboreal and nocturnal lifestyle. The reason for this is undocumented, but researchers suspect that tigers and leopards kill clouded leopards to eliminate competition. Therefore, clouded leopards are more active at night and spend more time in trees to avoid these large predators. Their patterned coat serves as camouflage when they are stalking their prey and attempting to remain hidden from other predators.

Known Predators:

  • tigers (Panthera tigris)
  • leopards (Panthera pardus)
  • humans (Homo sapiens)

Anti-predator Adaptations: cryptic

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Morphology

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The distinctive cloud-shaped markings of their coats make clouded leopards unmistakable. The fur is marked with elliptical blotches of a darker color than the background and the posterior edge of each blotch is partially framed in black. The blotches sit on a background field that varies from yellowish brown to dark gray. The muzzle is white and solid black spots mark the forehead and cheeks. The ventral side and limbs are marked with large, black ovals. Two solid black bars run from behind the ears along the back of the neck down to the shoulder blades and the bushy, thick tail is ringed in black. In juveniles, lateral spots are solid, not clouded. These will change by the time the animal is approximately six months old.

Adults usually weigh between 18 and 22 kilograms and stand at 50 to 60 centimeters at the shoulder. The head-body length is between 75 and 105 centimeters, and the tail length is between 79 and 90 centimeters, which is nearly as long as the body itself. There is no marked sexual dimorphism in clouded leopards, although females are slightly smaller. The legs are relatively short compared to other felids, with the hind limbs being longer than the fore limbs. The ankles have a wide range of motion and the feet are large and padded with retractile claws. As in other members of the family Felidae, the radius and the ulna are not fused, which allows for greater independence of motion. Clouded leopards have a digitigrade stance.

The skull is long and narrow compared to other felids and has well-developed crests to support the jaw muscles. Clouded leopards have the longest canine teeth relative to head and body size of any of the felids; canines can reach four centimeters or longer. A wide diastema lies between the premolars and canines, and individuals are often missing their first premolar.

The nose pad is pink and sometimes has small black spots, and the ears are short and round. The iris of the eye is usually brownish yellow or grayish green, and the pupils contract into vertical slits.

Range mass: 11 to 23 kg.

Average mass: 18-22 kg.

Range length: 123 to 200 cm.

Average length: 154-195 cm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Life Expectancy

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The average lifespan of wild clouded leopards is estimated to be 11 years. Individuals in zoos have been recorded living up to 17 years, with the average between 13 and 15 years. For wild clouded leopards, hunting or habitat destruction by humans limits lifespan. Clouded leopards also share parts of their geographic range with larger predators that kill potential competitors, such as tigers or leopards. Clouded leopards may spend a significant amount of time in trees for this reason. Studies have not been conducted regarding diseases that may limit the lifespan of clouded leopards. The number of deaths by other clouded leopards also remains unknown.

Average lifespan
Status: wild:
11 years.

Range lifespan
Status: captivity:
17 (high) years.

Average lifespan
Status: captivity:
13-15 years.

Average lifespan
Status: wild:
11 years.

Typical lifespan
Status: captivity:
17 (high) days.

Average lifespan
Status: captivity:
13-15 days.

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Habitat

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Clouded leopards occupy tropical forests at elevations up to 3000 meters. They are highly arboreal, using trees primarily for resting and also for hunting. However, they spend more time hunting on the ground than was originally believed. Sightings of clouded leopards occur most often in primary evergreen tropical forest but they have also been sighted in other habitats, such as secondary forest, logged forest, mangrove swamp, grassland, scrub land, dry tropical forest, and coastal hardwood forest.

Range elevation: 0 to 3000 m.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: savanna or grassland ; forest ; rainforest ; scrub forest ; mountains

Wetlands: swamp

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Distribution

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Clouded leopards, Neofelis nebulosa, are found south of the Himalayas in Nepal, Bhutan, and some areas of northeastern India. Myanmar, southern China, Taiwan, Vietnam, Laos, Cambodia, Thailand, and mainland Malaysia make up the southern parts of its geographic range. Three subspecies are recognized, occupying different regions within the range. Neofelis nebulosa nebulosa is found from southern China to mainland Malaysia; Neofelis nebulosa brachyura formerly lived in Taiwan but is now probably extinct; and Neofelis nebulosa macrosceloides is found from Myanmar to Nepal. Until recently, Neofelis diardi was classified as a subspecies of Neofelis nebulosa, but researchers studying molecular evidence now consider it to be a separate species. Neofelis diardi inhabits the islands of Borneo and Sumatra.

Biogeographic Regions: oriental (Native )

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Trophic Strategy

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Little is known about the feeding behavior of clouded leopards. Like other felids, they are strict carnivores. They are also solitary hunters, preying on birds, fish, monkeys, deer, and rodents. Prey species include argus pheasant, stump-tailed macaque, slow loris, silvered leaf monkey, sambar, hog deer, Indian muntjac, lesser mouse-deer, wild boar, bearded pig, Malayan pangolin, Indochinese ground squirrel, Asiatic brush-tailed porcupine, and masked palm civet. They have also been known to kill domestic animals, including calves, pigs, goats, and poultry. Fish remains have been found in the excrement of wild clouded leopards. Clouded leopards kill prey with a bite to the back of the neck, which snaps the spine. They pull flesh off of the carcass by stabbing the meat with its incisors and large canines and then abruptly jerking the head back.

Animal Foods: birds; mammals; fish

Primary Diet: carnivore (Eats terrestrial vertebrates, Piscivore )

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Associations

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Clouded leopards are one of the top predators in their range, especially where tigers and leopards are absent. They play a role in controlling populations of prey species, which effectively limits the impact which these populations have on the ecosystem. For example, by preying on cervids and keeping population size low, clouded leopards prevent excessive stress on plant populations. Like all other mammals, clouded leopards can be hosts for many internal parasites, as well as ectoparasites. Internal parasites found in the feces of clouded leopards include liver flukes (Dicrocoeliidae), intestinal flukes (Echinostomatidae), Paragonimus westermanni, Gnathostoma spinigerum, pseudophyllid cestodes (Pseudophyllidea), cyclophyllidean tapeworms (Mesocestoididae, Hymenolepididae, Taeniidae), Toxoplasma gondii, Mammomonogamus, Toxascaris, Oncicola, Sarcocystis, and Giardia. Many of these parasites are probably acquired from prey species. Ectoparasites of clouded leopards include several tick species: Amblyomma testudinarium, Haemaphysalis asiatica, Haemaphysalis hystricis, Haemaphysalis semermis, Rhipicephalus haemaphysaloides, and Ixodes granulatus.

Commensal/Parasitic Species:

  • Gnathostoma spinigerum
  • Paragonimus westermani
  • Echinostomatidae
  • Dicrocoeliidae
  • Pseudophyllidea
  • Mesocestoididae
  • Hymenolepididae
  • Taeniidae
  • Toxoplasma gondii
  • Mammomonogamus
  • Toxascaris
  • Oncicola
  • Sarcocystis
  • Giardia
  • Amblyomma testudinarium
  • Haemaphysalis asiatica
  • Haemaphysalis hystricis
  • Haemaphysalis semermis
  • Rhipicephalus haemaphysaloides
  • Ixodes granulatus
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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Benefits

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Clouded leopards have been hunted extensively for their pelts, which may be bought on the wildlife black market. The smuggling of skins from mainland China has increased as the demand for clouded leopard pelts in Taiwan has been renewed. Prior to the conversion of tribal peoples in Taiwan to Christianity, clouded leopard skins were used in ceremonies and the hunter was considered heroic for killing these animals. Today, ownership of a clouded leopard pelt is a status symbol among men in some Asian countries. Authorities have found pelts for sale in many markets throughout mainland Southeast Asia as well. Body parts, especially claws, teeth and bones, are still used in traditional medicine practices. Clouded leopard occasionally appears on menus at upscale restaurants in Asia. In addition, live animals are traded illegally as pets.

Positive Impacts: pet trade ; food ; body parts are source of valuable material

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Benefits

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As agricultural lands continue to encroach on clouded leopard habitat, incidences of clouded leopard attacks on livestock have increased. Clouded leopards prey on calves, goats, pigs, and poultry. Villagers use poison to kill predators such as clouded leopards.

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Behavior

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Like other felids, clouded leopards have keen vision as well as good senses of smell and hearing. Captive clouded leopards mark their territories by clawing trees, urine spraying, scraping, and head rubbing, all of which are typical scent-marking behaviors. Vocalizations made by captive animals are characteristic of members of the family Felidae, which include growling, mewing, hissing, and spitting. Clouded leopards do not purr, but they do make a low-intensity snorting noise called “prusten” when they have friendly interactions with other individuals. Clouded leopards, tigers, snow leopards, and jaguars are the only felids that use this type of vocalization. They also have a long moaning call that can be heard across distances. The purpose of this call is unknown, but observers think it is a form of communication between animals in different territories, perhaps as a mating call or to warn other cats away from their territory. Clouded leopards also have vibrissae on their muzzles, which detect tactile stimuli, especially at night.

Communication Channels: visual ; tactile ; acoustic ; chemical

Other Communication Modes: scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Untitled

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A subspecies of clouded leopards, Neofelis nebulosa diardi, is now considered a separate species, Neofelis diardi. They are found on Borneo.

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Reproduction

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All that is known about the mating behavior of clouded leopards comes from observations of captive animals. This lack of knowledge concerning wild mating behavior has made it extremely difficult to breed these animals in captivity. Arranged mating encounters at zoos often conclude with aggression between the two individuals, and the male often kills the female with a bite to the back of the neck. For this reason, many experts believe that compatibility between a male and female is important for productive matings. The most successful matings have occurred between a male and female that were raised together from only a few weeks of age. However, researchers do not believe that clouded leopards are monogamous in the wild. In zoos, mating usually occurs between December and March, but it can occur at any time throughout the year. Because clouded leopards occupy tropical habitats, breeding may be less seasonal in the wild. The mating pair copulates many times over the course of several days. The male typically grasps the female with a bite to the back of the neck before an intromission, and the female vocalizes once the intromission occurs. In the wild, clouded leopards use elevated areas to deliver a long moaning call that travels well. This call is suspected to be a mating call, but it may be a territorial call instead.

The gestation period for captive clouded leopards normally lasts between 88 and 95 days, although it can last anywhere from 85 to 109 days. Females most often give birth to two kittens per pregnancy, but litters of one to five kittens have been documented as well. Kittens are born with the large spots that are characteristic of their adult counterparts, but these spots are solid black until approximately six months of age. A newborn kitten weighs between 140 and 280 grams, depending on the size of the litter. Kittens first open their eyes between two and eleven days of age. Clouded leopard kittens begin walking at 20 days of age, and they can climb trees as early as six weeks old. They start to consume flesh between 7 and 10 weeks old, and they are weaned shortly thereafter at 10 to 14 weeks. It has been reported that clouded leopard kittens are able to kill chickens at 10 weeks old. At zoos, clouded leopard kittens are typically taken away from their mothers to be hand-reared but, in the wild, kittens normally stay with their mothers for about ten months. Little is known about the interbirth interval of female clouded leopards. The length of time between births for captive cats has ranged from 10 to 16 months. Clouded leopards in captivity arrive at sexual maturity between 20 and 30 months of age, with the average being 23 to 24 months.

Breeding interval: The length of time between matings for captive cats has ranged from 10 to 16 months.

Breeding season: In captivity, breeding usually occurs between December and March, but it can occur year round.

Range number of offspring: 1 to 5.

Average number of offspring: 2.

Range gestation period: 85 to 109 days.

Average gestation period: 88-95 days.

Range weaning age: 10 to 14 weeks.

Average time to independence: 10 months.

Range age at sexual or reproductive maturity (female): 20 to 30 months.

Average age at sexual or reproductive maturity (female): 23-24 months.

Range age at sexual or reproductive maturity (male): 20 to 30 months.

Average age at sexual or reproductive maturity (male): 23-24 months.

Key Reproductive Features: iteroparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous

Average birth mass: 170 g.

Average number of offspring: 2.

After mating, male and female clouded leopards separate, and the male does not take part in the rearing of offspring. The gestation period is typically between 88 and 95 days. The female does not appear pregnant until the third trimester, at which time her abdomen and nipples become larger. When the kittens are born, the mother licks them to keep them clean and warm. She continues to clean them until they learn to do so themselves. It is unknown where a female keeps her young while she is hunting, but she probably hides them in dense vegetation. Females produce milk for the kittens, which is their sole source of nutrition until they are between 7 and 10 weeks old. They are completely weaned when they are between 10 and 14 weeks of age. Until they are approximately 10 months old, the mother continues to provide them with prey while they grow and learn to hunt for themselves. At this age, they leave their mothers to find their own territories.

Parental Investment: altricial ; pre-fertilization (Provisioning, Protecting: Female); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female); extended period of juvenile learning

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Holmes, K. 2009. "Neofelis nebulosa" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Neofelis_nebulosa.html
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Katie Holmes, University of Michigan-Ann Arbor
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Phil Myers, Museum of Zoology, University of Michigan-Ann Arbor
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Biology

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The clouded leopard has amazing tree climbing abilities, indeed in Malaysia its local name actually means 'branch-of-a-tree tiger' (4). It has been seen running head-first down tree trunks, climbing about on the underside of branches, and hanging upside down by its hind feet with the tail providing balance (6). The ability to climb trees allows it to forage for food in the canopy although it mainly uses the tree branches for resting. This species also swims well and has been found on small islands off the mainland in the past (6). It hunts by day or night, either stalking its prey on the ground or by ambushing it from the trees (7). It was originally thought that the long canines were for preying on large ungulates, though recent studies show that it feeds mainly on primates, birds, small mammals, porcupines, deer, and wild boar, as well as domestic livestock (5). These leopards are believed to be solitary animals except during the breeding season, when the males seek out the females (2), though little is known about their biology due to their elusiveness and so most information comes from captive individuals (7). The gestation period is between 86 and 93 days, and the female bears between one and five cubs, each weighing around 150 – 280 grams (4). Born with much darker side markings than the adults they are nursed for up to five months and achieve independence at nine months (7). It is not known how long the clouded leopard lives for in the wild, but captive individuals have lived for up to seventeen years (2).
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Conservation

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The clouded leopard is generally protected under game laws and fully protected in parks and reserves, but there is not enough knowledge about wild populations to draw firm conclusions about the success of these conservation measures. The Formosan clouded leopard of Taiwan, (N. nebulosa brachyurus) a subspecies of Neofelis is thought to be extinct, and is an indication of the possible fate of clouded leopards elsewhere (2). Captive breeding programmes were initially unsuccessful due to the small gene pool available in captive as well as wild individuals (2). In addition, there has been high incidence of males killing females in captivity (5). The Clouded Leopard Species Survival Plan (SSP) recognised these problems and in the late 1990s set up the Thailand Clouded Forest Foundation to research the behaviour of these cats and improve captive management and artificial reproduction (8). In 1998, 22 compatible pairs were given breeding recommendations and from January 1998 to August 2001, a total of 52 births occurred from 12 pairs (8). These results are positive, but there are still problems of reduced genetic diversity among the captive leopards and the small numbers of breeding individuals, which the Foundation is trying to address (8). The clouded leopard's survival depends on these conservation measures. It is hoped that one day we can attribute the rare sightings of this beautiful leopard to its elusive nature rather than diminishing numbers (5).
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Description

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The clouded leopard is named after the distinctive 'clouds' on its coat - ellipses partially edged in black, with the insides a darker colour than the background colour of the pelt (4). The base of the fur is a pale yellow to rich brown, making the darker cloud-like markings look even more distinctive (5). The limbs and underbelly are marked with large black ovals, and the back of its neck is conspicuously marked with two thick black bars (6). The clouded leopard is about the size of a small Labrador retriever with a body length of up to 90 cm, an intermediate size between the large and small cats of the world (4). It does, however, have an exceptionally long tail for balancing, which can be as long as the body itself, thick with black ring markings (2). In proportion to its body size it also has the largest canines of all the cats, a feature that has earned this cat the reputation of being the 'modern day sabre-tooth' (5). Well adapted to forest life, the clouded leopard also has relatively short legs and broad paws which make it excellent at climbing trees and creeping through thick forest (7).
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Habitat

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These shy and elusive cats are usually associated with tropical forests, but they also make use of other habitats. They have been seen in primary and secondary logged forest as well as grassland and scrub, mangrove swamps and even dry tropical forest (6).
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Range

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The clouded leopard is found in tropical and sub-tropical forests from India, South China, Burma, and Indochina to Sumatra and Borneo (5). Despite this wide distribution, it actually has a very low population and is very thinly dispersed (5). The clouded leopard's total effective population size is estimated at below 10,000 mature breeding individuals; no subpopulation contains more than 1,000 mature breeding individuals (1).
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Status

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Classified as Vulnerable (VU C2a(i)) on the IUCN Red List 2002 (1) and is listed on Appendix I of CITES (3).
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Threats

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Deforestation resulting from commercial logging and the growth of human settlements is thought to be the foremost threat to this species, (7). Not only does deforestation remove the clouded leopard's own shelter and habitat, but it reduces the number of prey species (4). As humans have increasingly encroached on their habitats, these leopards have been known to prey on livestock which puts them at risk of being killed by the owners (2). Another major threat is the hunting of this cat for its beautiful pelt and decorative teeth as well as its bones, which are prized in the traditional Asian medicinal trade (6). Clouded leopards have even featured on the menu of restaurants in Thailand and China which cater to wealthy Asian tourists (6).
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Brief Summary

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Smithsonian Channel Video: Ghost Cat: Saving the Clouded Leopard (Full Episode)
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Clouded leopard

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The clouded leopard (Neofelis nebulosa), also called mainland clouded leopard, is a wild cat inhabiting dense forests from the foothills of the Himalayas through 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 first cat that genetically diverged 9.32 to 4.47 million years ago from the common ancestor of the pantherine cats. Today, the clouded leopard is locally extinct in Singapore, Taiwan, and possibly also in Hainan Island and Vietnam. Its total population 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. 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 Canton Province 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 estimated to have been the first cat that genetically diverged from the common ancestor of the Felidae 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.[2]

In Nepal, the clouded leopard was thought to be extinct 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 Pokhara Valley. These findings extended the known range westward, suggesting it is able to survive and breed in degraded woodlands that previously harboured moist subtropical semideciduous forest.[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

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 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 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 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.

Many of the remaining forest areas throughout the range are too small to ensure the long-term persistence of clouded leopard populations.[57] 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 body parts of the tiger in traditional Asian medicine and tonics, and live animals for the pet 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. Three of the surveyed markets are situated on international borders with China and Thailand, and cater to international buyers. Implementation and enforcement of CITES is considered inadequate.[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 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.[53] These bans, however, are poorly enforced in India, Malaysia and Thailand.[58]

In the United States, the clouded leopard is listed as endangered under the Endangered Species Act, prohibiting trade in live animals or body parts.[1]

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, including six zoos in India:[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

References

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Clouded leopard: Brief Summary

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The clouded leopard (Neofelis nebulosa), also called mainland clouded leopard, is a wild cat inhabiting dense forests from the foothills of the Himalayas through 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 first cat that genetically diverged 9.32 to 4.47 million years ago from the common ancestor of the pantherine cats. Today, the clouded leopard is locally extinct in Singapore, Taiwan, and possibly also in Hainan Island and Vietnam. Its total population 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. 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.

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