Sunda Clouded Leopard/Indochinese Clouded Leopard
Neofelis diardi (G. Cuvier, 1823)/Neofelis nebulosa (Griffith, 1821)
IUCN RED LIST (2008): Vulnerable
Head-body length ♀ 68.6−94cm, ♂ 81.3−108cm
Tail 60−92cm
Weight ♀ 10−11.5kg, ♂ 17.7−25kg
Taxonomy and phylogeny
Until 2006, the Clouded Leopard was considered a single species classified in its own genus, Neofelis. Genetic analyses published in 2006–2007 strongly suggest that island populations on Borneo and Sumatra have been reproductively isolated from mainland populations for 1.4–2.9 million years, well within the range of species-level divisions between other large cats. These molecular distinctions are supported by differences in the pelage, and in measurements of the skulls and teeth. Taken collectively, these differences are now considered sufficient to recognise clouded leopards as two species: N. diardi (Sunda Clouded Leopard or Sundaland Clouded Leopard; Borneo and Sumatra) and N. nebulosa (Indochinese Clouded Leopard or simply Clouded Leopard; mainland Asia). Each species is provisionally divided into two subspecies. There is limited genetic data supporting the subspecific separation of the Borneo population of the Sunda Clouded Leopard (N. diardi borneensis) from that on Sumatra (N. diardi sumatrensis). The mainland population of the Indochinese Clouded Leopard is historically split into eastern (N. nebulosa nebulosa) and western (N. nebulosa macrosceloides) subspecies that have minor morphological differences yet no significant pelage or genetic differences. A third putative subspecies on Taiwan, N. nebulosa brachyura, is likely extinct. Clouded Leopards are closely related to the ‘big cats’ and are grouped with them in the Panthera lineage. Neofelis diverged very early from the line that led to the genus Panthera, with a common ancestor estimated around 6.4 million years ago.
It is unclear if the two Clouded Leopard species can interbreed. It is hypothetically possible given their relatively recent divergence, although there is no natural range overlap except perhaps in Peninsular Malaysia (where genetic analysis is yet to confirm if one or both clouded leopard species are present).
Description
Clouded Leopards of both species are very similar in general appearance and size; based on very few samples, there is limited evidence Sunda Clouded Leopard males are larger than mainland males. Clouded Leopards have a long body, relatively short robust legs with large feet and a very long tail. The head is long and heavily built, similar in overall shape and proportions to the Panthera cats. Clouded Leopards have an exceptionally large gape (almost 90° compared to 65° for the Puma) and elongated canine teeth measuring up to 4cm, relatively the longest of any living felid. This is similar to extinct sabre-toothed cats (to which all modern cats including Clouded Leopards are not closely related), the reasons for which are unclear. Sunda Clouded Leopards of both sexes have the longest canines by a small margin. Pelage differences between the two Clouded Leopards are distinct. Sunda Clouded Leopards are overall darker with grey to greyish-yellow background fur and relatively small, irregular blotches with thick, black margins and usually small black spots within each blotch. The lower legs are marked with solid, black blotches that are closely clustered. Indochinese Clouded Leopards are generally paler and brighter with buff to rich tawny background colour, and very large blotches with narrower black margins and few or no spots within the blotches. Solid black blotches on the lower legs tend to be more widely spaced than in Sunda cats. Melanism is reported anecdotally, though there is no physical evidence.
Similar species The similar Marbled Cat is sympatric with both clouded leopard species. Clouded Leopards are much larger with a distinctly longer, heavier head typical of a ‘big cat’ compared to the small rounded head of the Marbled Cat. Both Clouded Leopard species have larger, more discrete blotches with distinct edges compared to the more diffuse markings of the Marbled Cat.
A Sunda Clouded Leopard (opposite) and Indochinese Clouded Leopard (below, C), showing the distinct differences in pelage. There are also significant differences between the two species in skull morphology in which adaptations similar to those in extinct sabre-tooths are more advanced in the Sunda Clouded Leopard.
Distribution and habitat
The Sunda Clouded Leopard is endemic to Borneo and Sumatra. On Borneo it occurs in Kalimantan (Indonesia), Sabah and Sarawak (Malaysia) and Brunei, potentially on approximately 50 per cent of the island where there is still good forest cover. It is absent (or likely so) in large deforested areas mainly in the south-east and west, and along a wide coastal band. On Sumatra, it occurs largely along the Bukit Barisan mountain range running the length of the island. It is unknown if Sunda Clouded Leopards occur on the offshore Batu Islands close to Sumatra. Clouded Leopards occurred on Java perhaps as recently as the Holocene (starting 11,700 years before the present) but not in modern times, and they have never occurred on Bali. The Indochinese Clouded Leopard occurs in central Nepal, Bhutan, north-east India and eastern Bangladesh (marginally), across China south of the Yangtze River, and patchily throughout Indochina including the Malay Peninsula (which is usually regarded as part of the Sundaland region, south of the Isthmus of Kra; however, based on their pelage, the Clouded Leopards on the peninsula are considered N. nebulosa). The species is considered extinct on Taiwan.
Clouded Leopards are closely associated with dense habitat and are considered forest-dependent. They occur in all kinds of dense moist and dry forest, peat-swamp forest, dry woodlands and mangroves from sea level to 1,500m (Sarawak) and 3,000m in the Himalayan foothills. They use grassland patches in forest-grassland mosaics, and a radio-collared cat in Nepal rested in dense, 4–6m-tall grass on terai floodplain grasslands. Clouded Leopards appear to tolerate some habitat modification. They may be relatively common in secondary and selectively logged forest but their ability to use forestry areas seems to decrease beyond a threshold of fairly light logging intensity. Clouded Leopards occasionally use oil-palm plantations but all camera-trap evidence indicates they largely use only the periphery of plantations. A radio-collared male in Borneo crossed oil-palm plantations under 1km-wide between forest patches and made one foray of around 2.5km through an oil palm-scrub mosaic; he did not linger in oil palm habitat and always moved rapidly through it.
During observed attacks on Proboscis Monkey troops by Sunda Clouded Leopards, adult male monkeys attempt to defend young monkeys. In one incident 10m above ground, a male Probsocis drove off a Clouded Leopard that had captured a 10-month-old infant but not before it was killed.
Feeding ecology
Clouded Leopard ecology is poorly known. Based on mostly anecdotal and incidental records, they are known to prey on a wide variety of small and medium-sized vertebrates including both terrestrial and arboreal species, and diurnal and nocturnal species. Primates and small ungulates possibly constitute the staple prey. However, as in most felids, the diet is presumably flexible and varies regionally depending on the composition of the prey community and the presence of other carnivores. Clouded Leopards are the largest felid with few competitors on Borneo whereas Tigers, Leopards and Dholes co-occur on mainland Asia and on Sumatra (excluding Leopards). It is unknown if or how these species affect Clouded Leopard feeding patterns.
Clouded Leopards are recorded killing a range of primates, from small, nocturnal slow lorises to large, diurnal adult male Proboscis Monkeys. Anecdotes of predation on Orang-utans have not been confirmed. They are capable of taking ungulates at least up to their own size, with reliable records of Barking Deer, Hog Deer (including adults) and Bearded Pig (probably young animals). Smaller prey includes oriental chevrotains, Malayan Pangolins, brush-tailed porcupines, small rodents including Indochinese Ground Squirrels, Binturongs, Common Palm Civets, and a variety of birds. Local people in Sabah and Sarawak report that fish are sometimes eaten. Clouded Leopards occasionally kill poultry; a subadult male Clouded Leopard in Nepal was trapped in a chicken coop (and subsequently released with a radio-collar). Attacks on livestock are rare. One Sunda Clouded Leopard was shot after reportedly killing goats in an enclave village surrounded by forest.
The Clouded Leopard’s morphology suggests a high degree of arborealism, with short robust legs, broad feet and a long tail. There are numerous accounts of hunting above ground, including four published observations of Clouded Leopards attacking Proboscis Monkeys in trees, one resulting in the capture of a juvenile monkey 7m above ground. However, based on a small number of data points, radio-tracked individuals travelled and hunted mostly on the ground; Clouded Leopards probably search for prey mainly from the ground and readily pursue arboreal species into trees as they are located. They hunt primarily at night with crepuscular activity peaks, although two collared individuals in Thailand were often active in the morning until midday. One of these radio-collared cats, an adult male, hunted Hog Deer and Barking Deer as they bedded down en masse on open grasslands at dusk. This Clouded Leopard rested at the forest edge before moving out onto the grasslands at night to target the deer. One of its kills, an adult male Hog Deer, was killed with a 3cm deep bite through the spine above the shoulders. The same technique was reported by local people in Sabah and Sarawak who found Clouded Leopard kills of deer and pigs. Similarly, two young Proboscis Monkeys were both attacked with bites to the back of the head and neck. This is a very unusual killing technique compared to other felids, which typically kill large prey and especially ungulates from asphyxiation by biting the throat, and it may be related to the Clouded Leopard’s unique dentition. It is unknown if Clouded Leopards scavenge.
A Clouded Leopard attacks a Hog Deer. With so little known about the Clouded Leopard’s killing technique, it is still unclear whether the species routinely uses the deep neck bite observed on a handful of carcasses.
The Clouded Leopard’s blade-like upper canines and wide gape are among many features it shares with extinct sabretooths including a lowered jaw joint and a reinforced mandible that is resistant to bending.
Social and spatial behaviour
Only 12 Clouded Leopards have ever been radio-collared: five in Borneo and seven in Nepal and Thailand. The latter two studies provided a limited amount of data over a short period; the former study is ongoing with GPS telemetry that is expected to provide greater detail. Based on the little available information, Clouded Leopards are thought to follow a typical felid pattern of overlapping territories with exclusive core areas. Limited data suggest the sexes have similar territory sizes although it is likely that males have larger ranges than females and overlap multiple female territories; this awaits more research. The published range sizes are 16.1−40km2 (two subadult females and two adult females) and 35.5−43.5km2 (one subadult male and two adult males). Density estimates appear to be quite low compared to other cats, especially on Borneo despite the lack of competing large carnivores. Density estimates for Sunda Clouded Leopards include 0.84−1.04 per 100 km2 in degraded secondary lowland forest (Tangkulap-Pinangah and Segaliud Lokan Forest Reserves, Sabah); 1.29 per 100km2 (Tesso Nilo-Bukit Tigapuluh Conservation Landscape, Sumatra); 1.9 per 100km2 for primary forest (Maliau Basin Conservation Area, Sabah) which drops to 0.8 per 100km2 when adjacent logged forest is included in the estimate; 1.76 per 100km2 in well-protected primary forest (Danum Valley Conservation Area, Sabah); and 2.55 per 100km2 in secondary forest undergoing rehabilitation (Ulu Segama Forest Reserve, Sabah). The only rigorous density estimate for Indochinese Clouded Leopard is 4.7 per 100km2 (Manas National Park, India).
Reproduction and demography
This is virtually unknown from the wild. Most information comes from captive animals (all of them Indochinese Clouded Leopards) and is based on few records. Breeding occurs year-round in captivity and is likely to be aseasonal in the wild. Unusually, males quite frequently kill females in captivity (with a similar spinal nape bite as for large prey), the reasons for which are unknown but assumed to be related to a captive setting; it is extremely unlikely that this behaviour is common for wild individuals. Oestrus lasts about one week and gestation lasts 85−95 days (rarely to 109 days). Litters average two to three kittens, exceptionally reaching five. Weaning begins around 7−10 weeks. Sexual maturity is 20−30 months.
Mortality People are responsible for most deaths in studied areas, otherwise mortality is unknown. Large carnivores are potential predators, though there are no records. Clouded Leopards take refuge in trees when pursued by domestic dogs.
Lifespan Unknown in the wild; up to 17 years in captivity.
The Clouded Leopard is half the size of the next smallest species of the Panthera lineage, but its big cat ancestry is clear in the elongated, heavily built head and jaws with a relatively smaller braincase compared to small cats.
STATUS AND THREATS
Clouded Leopards are relatively widespread but their status is poorly known in most of the range. Although they are often regarded as more resilient than larger felids, they do not appear to attain high densities anywhere and they are closely associated with forested habitats which are undergoing extremely rapid conversion by people in much of the range. South-east Asia has the world’s fastest deforestation rate due to logging and conversion for settlement and agriculture, including plantations especially of oil palm and rubber. Forest loss is the main factor driving declines and has contributed to extirpation of the Sunda Clouded Leopard from an estimated 50 per cent of Borneo and around two-thirds of Sumatra. The Indochinese Clouded Leopard has undergone similar range loss and decline, particularly in China and in Indochina where the distribution is very patchy, and there are few recent records for most of Cambodia, China, Laos and Vietnam. Clouded Leopard skins, bones and meat have commercial value, and are illegally traded in wildlife markets; for example, 13 market surveys between 2001 and 2010 at two border towns in Burma recorded parts of at least 149 individuals. Clouded Leopards are vulnerable to snares and are hunted opportunistically in much of their range including in protected areas; a minimum number of seven were killed inside Kerinci Seblat National Park (Sumatra) in 2000−2001.
Both species: CITES Appendix I. Red List: Vulnerable. Population trend: Decreasing.