Leopard Cat

The leopard cat (Prionailurus bengalensis) is a small wild cat native to South, Southeast and East Asia. Since 2002 it has been assessed as Least Concern on the IUCN Red List as it is widely distributed but threatened by habitat loss and hunting in parts of its range.

Leopard cat subspecies differ widely in fur colour, tail length, skull shape and size of carnassials. Archaeological evidence from China indicates that the leopard cat was the first cat species to be domesticated, prior to being supplanted in favour of the wildcat.

Characteristics
A leopard cat is about the size of a domestic cat, but more slender, with longer legs and well-defined webs between its toes. Its small head is marked with two prominent dark stripes and a short and narrow white muzzle. There are two dark stripes running from the eyes to the ears, and smaller white streaks running from the eyes to the nose. The backs of its moderately long and rounded ears are black with central white spots. Body and limbs are marked with black spots of varying size and color, and along its back are two to four rows of elongated spots. The tail is about half the size of its head-body length and is spotted with a few indistinct rings near the black tip. The background color of the spotted fur is tawny, with a white chest and belly. However, in their huge range, they vary so much in coloration and size of spots as well as in body size and weight that initially they were thought to be several different species. The fur color is yellowish brown in the southern populations, but pale silver-grey in the northern ones. The black markings may be spotted, rosetted, or may even form dotted streaks, depending on subspecies. In the tropics, leopard cats weigh 0.55 to 3.8 kg, have head-body lengths of 38.8 to 66 cm, with long 17.2 to 31 cm tails. In northern China and Siberia, they weigh up to 7.1 kg, and have head-body lengths of up to 75 cm; generally, they put on weight before winter and become thinner until spring. Shoulder height is about 41 cm.

Leopard cats in the Sundaic region are darker, have smaller spots and shorter tails than leopard cats in mainland Asia.

Distribution and habitat
Leopard cats are the most widely distributed Asian small cats. Their range extends from the Amur region in the Russian Far East over the Korean Peninsula, China, Indochina, the Indian Subcontinent, to the West in northern Pakistan, and to the south in the Philippines and the Sunda islands of Indonesia. They live in tropical evergreen rainforests and plantations at sea level, in subtropical deciduous and coniferous forests in the foothills of the Himalayas at altitudes above 1000 m. They are able to tolerate human-modified landscapes with vegetation cover to some degree, and inhabit agriculturally used areas such as oil palm and sugar cane plantations.

In 2009, a leopard cat was camera trapped in Nepal's Makalu-Barun National Park at an altitude of 3254 m. At least six individuals inhabit the survey area, which is dominated by associations of rhododendron, oak and maple. The highest altitudinal record was obtained in September 2012 at 4500 m in the Kanchenjunga Conservation Area.

In the northeast of their range they live close to rivers, valleys and in ravine forests, but avoid areas with more than 10 cm of snowfall. They are rare in Pakistan's arid treeless areas. In Afghanistan, they were reported in the 1970s from Jalalkot and Norgul in the Kunar Valley, and the Waygul forest of Dare Pech.

In Sabah's Tabin Wildlife Reserve leopard cats had average home ranges of 3.5 km2. In Thailand's Phu Khieu Wildlife Reserve 20 leopard cats were radio-collared between 1999 and 2003. Home ranges of males ranged from 2.2 km2 to 28.9 km2, and of the six females from 4.4 km2 to 37.1 km2.

Leopard cats are not found in the Japanese archipelago, but fossils have been excavated there from the Pleistocene period.

Distribution of subspecies
the following subspecies are recognized:
 * Indian leopard cat P. b. bengalensis (Kerr, 1792) — ranges from India, Bangladesh, Myanmar, Thailand, the Malay Peninsula, to Indochina and Yunnan in China;
 * Javan leopard cat P. b. javanensis (Desmarest, 1816) — inhabits Java and Bali;
 * Sumatran leopard cat P. b. sumatranus (Horsfield 1821) — inhabits Sumatra and Tebingtinggi;
 * Chinese leopard cat P. b. chinensis (Gray 1837) — lives in Taiwan and China except Yunnan;
 * Himalayan leopard cat P. b. horsfieldi (Gray 1842) — ranges in Kashmir, Punjab, Kumaon, Nepal, Sikkim and Bhutan;
 * Amur leopard cat P. b. euptilura (Elliott 1871) — is distributed in eastern Siberia, Manchuria, Korea and on the Tsushima Island;
 * Bornean leopard cat P. b. borneoensis (Brongersma 1936) — inhabits Borneo;
 * P. b. trevelyani (Pocock 1939) — lives in northern Kashmir and Punjab, and in southern Baluchistan;
 * Hainan leopard cat P. b. alleni (Sody, 1949) — inhabits Hainan Island
 * Iriomote cat P. b. iriomotensis (Imaizumi, 1967) — is found exclusively on the island of Iriomote, one of the Ryukyu Islands in the Japanese Archipelago;
 * Palawan leopard cat P. b. heaneyi (Groves 1997) — inhabits the Philippine island of Palawan;
 * Visayan leopard cat P. b. rabori (Groves 1997) — inhabits the Philippine islands of Negros, Cebu, and Panay.

The Iriomote cat has been proposed as a species since 1967, but following mtDNA analysis in the 1990s it is now considered a subspecies of the leopard cat.

The Tsushima leopard cat (Japanese: Tsushima yamaneko) lives exclusively on Tsushima Island. Initially regarded as belonging to the Chinese leopard cat subspecies, it is now considered an isolated population of the Amur leopard cat, P. b. euptilura.

Ecology and behavior
Leopard cats are solitary, except during breeding season. Some are active during the day, but most hunt at night, preferring to stalk murids, tree shrews and hares. They are agile climbers and quite arboreal in their habits. They rest in trees, but also hide in dense thorny undergrowth on the ground. In the oil palm plantations of Sabah, they have been observed up to 4 m above ground hunting rodents and beetles. In this habitat, males had larger home ranges than females, averaging 3.5 km2 and 2.1 km2 respectively. Each male's range overlapped one or more female ranges. There is evidence to suggest that rats are abundant and may be easy to catch in oil palm plantations. There, leopard cats feed on a large proportion of rats compared to forested areas.

Leopard cats can swim, but seldom do so. They produce a similar range of vocalisations to the domestic cat. Both sexes scent mark their territory by spraying urine, leaving faeces in exposed locations, head rubbing, and scratching.

Diet
Leopard cats are carnivorous, feeding on a variety of small prey including mammals, lizards, amphibians, birds and insects. In most parts of their range, small rodents such as rats and mice form the major part of their diet, which is often supplemented with grass, eggs, poultry, and aquatic prey. They are active hunters, dispatching their prey with a rapid pounce and bite. Unlike many other small cats, they do not "play" with their food, maintaining a tight grip with their claws until the animal is dead. This may be related to the relatively high proportion of birds in their diet, which are more likely to escape when released than are rodents.

Reproduction and development
The breeding season of leopard cats varies depending on climate. In tropical habitats, kittens are born throughout the year. In colder habitats farther north, females give birth in spring. Their gestation period lasts 60–70 days. Litter size varies between two and three kittens. Captive born kittens weighed 75 to 130 g at birth and opened their eyes by latest 15 days of age. Within two weeks, they doubled their weight and were four times their birth weight at the age of five weeks. At the age of four weeks, their permanent canines break through, and they begin to eat meat. Captive females reach sexual maturity earliest at the age of one year and have their first litter at the age of 13 to 14 months. Captive leopard cats have lived for up to thirteen years.

The estrus period lasts 5–9 days.

Threats
In China, leopard cats are hunted mainly for their fur. Between 1984 and 1989, about 200,000 skins were exported yearly. A survey carried out in 1989 among major fur traders revealed more than 800,000 skins on stock. Since the European Union imposed an import ban in 1988, Japan has become the main buyer, and imported 50,000 skins in 1989. Although commercial trade is much reduced, the species continues to be hunted throughout most of its range for fur, for food, and as pets. They are also widely viewed as poultry pests and killed in retribution.

In Myanmar, 483 body parts of at least 443 individuals were observed in four markets surveyed between 1991 and 2006. Numbers were significantly larger than non-threatened species. Three of the surveyed markets are situated on international borders with China and Thailand, and cater to international buyers, although the leopard cat is completely protected under Myanmar's national legislation. Implementation and enforcement of CITES is considered inadequate.

Conservation
Prionailurus bengalensis is listed in CITES Appendix II. In Hong Kong, the species is protected under the Wild Animals Protection Ordinance Cap 170. The population is well over 50,000 individuals and, although declining, the cat is not endangered.

The Tsushima leopard cat is listed as Critically Endangered on the Japanese Red List of Endangered Species, and has been the focus of a conservation program funded by the Japanese government since 1995.

In the United States, P. bengalensis is listed as Endangered under the Endangered Species Act since 1976; except under permit, it is prohibited to import, export, sell, purchase and transport leopard cats in interstate commerce.

Taxonomic history
In 1792, Robert Kerr first described a leopard cat under the binominal Felis bengalensis in his translation of Carl von Linné's Systema Naturae as being native to southern Bengal. Between 1829 and 1922, different authors of 20 more descriptions classified the cat either as Felis or Leopardus. Owing to the individual variation in fur colour, leopard cats from British India were described as Felis nipalensis from Nepal, Leopardus ellioti from the area of Bombay, Felis wagati and Felis tenasserimensis from Tenasserim. In 1939, Reginald Innes Pocock subordinated them to the genus Prionailurus. The collection of the Natural History Museum in London comprised several skulls and large amounts of skins of leopard cats from various regions. Based on this broad variety of skins, he proposed to differentiate between a southern subspecies Prionailurus bengalensis bengalensis from warmer latitudes to the west and east of the Bay of Bengal, and a northern Prionailurus bengalensis horsfieldi from the Himalayas, having a fuller winter coat than the southern. His description of leopard cats from the areas of Gilgit and Karachi under the trinomen Prionailurus bengalensis trevelyani is based on seven skins that had longer, paler and more greyish fur than those from the Himalayas. He assumed that trevelyani inhabits more rocky, less forested habitats than bengalensis and horsfieldi.

Between 1837 and 1930, skins and skulls from China were described as Felis chinensis, Leopardus reevesii, Felis scripta, Felis microtis, decolorata, ricketti, ingrami, anastasiae and sinensis, and later grouped under the trinomen Felis bengalensis chinensis. In the beginning of the 20th century, a British explorer collected wild cat skins on the island of Tsushima. Oldfield Thomas classified these as Felis microtis, which had been first described by Henri Milne-Edwards in 1872.

Two skins from Siberia motivated Daniel Giraud Elliot to write a detailed description of Felis euptilura in 1871. One was depicted in Gustav Radde's illustration cum description of a wild cat; the other was part of a collection at the Regent's Park Zoo. The ground colour of both was light brownish-yellow, strongly mixed with grey and covered with reddish-brown spots, head grey with a dark-red stripe across the cheek. In 1922, Tamezo Mori described a similar but lighter grey spotted skin of a wild cat from the vicinity of Mukden in Manchuria that he named Felis manchurica. Later both were grouped under the trinomen Felis bengalensis euptilura. In the 1970s and 1980s, the Russian zoologists Geptner, Gromov and Baranova disagreed with this classification. They emphasized the differences of skins and skulls at their disposal and the ones originating in Southeast Asia, and coined the term Amur forest cat, which they regarded as a distinct species. In 1987, Chinese zoologists pointed out the affinity of leopard cats from northern China, Amur cats and leopard cats from southern latitudes. In view of the morphological similarities they did not support classifying the Amur cat as a species.

The initial binomial euptilura given by Elliott has been incorrectly changed to "euptilurus" by some later authors, but under the International Code of Zoological Nomenclature Article 31.2.1, nouns and noun phrases are not subject to gender agreement; at present, both terms appear in use, but only the spelling "euptilura" is correct.

Molecular analysis of 39 leopard cat tissue samples clearly showed three clades: a northern lineage and southern lineages 1 and 2. The northern lineage comprises leopard cats from Tsushima Islands, the Korean Peninsula, the continental Far East, Taiwan, and Iriomote Island. Southern lineage 1, comprising Southeast Asian populations, showed higher genetic diversity. Southern lineage 2 is genetically distant from the other lineages.

Leopard cats and hybrids as pets
Archeological and morphometric studies have indicated that the first cat species to become a human commensal or domesticate in Neolithic China was the leopard cat, beginning at least 5000 years ago. However, these cats were ultimately replaced over time with cats descended from Felis sylvestris lybica from the Middle East.

The Asian leopard cat (P. b. bengalensis) has been mated with a domestic cat since the 1960s to produce hybrid offspring known as the Bengal cat. This hybrid is usually permitted to be kept as a pet without a license. For the typical pet owner, a Bengal cat should be at least four generations (F4) removed from the leopard cat. The "foundation cats" from the first three filial generations of breeding (F1–F3) are usually reserved for breeding purposes or the specialty-pet home environment.

Keeping a leopard cat as a pet may require a license in some localities. License requirements vary.