Felis pardus

Leopard

[lep-erd]

The leopard (lɛpɚd; Panthera pardus) is an Old World mammal of the Felidae family and the smallest of the four roaring cats in the genus Panthera, the other three are the tiger, lion and jaguar. Once distributed across southern Asia and Africa, from Korea to South Africa, the leopard's range of distribution has decreased radically over time because of a variety of factors, including human influence, and the leopard now chiefly occurs in sub-Saharan Africa. There are fragmented populations in India, Indochina, Malaysia, and China. Despite the loss of range and continual declines in population, the cat remains a "Least Concern" species; its numbers are greater than that of the other Panthera species, all of which face more acute conservation concerns.

The leopard has relatively short legs and a long body, with a large skull. Physically, it most closely resembles the jaguar, although it is usually smaller and of slighter build. Its fur is marked with rosettes which lack internal spots, unlike those of the jaguar. Leopards that are melanistic, either completely black or very dark in coloration, are one of the big cats known colloquially as black panthers.

The species' success in the wild owes in part to its opportunistic hunting behaviour, its adaptability to a variety of habitats and its ability to move at up to approximately 60 kilometres (37 miles) an hour. The leopard consumes virtually any animal it can hunt down and catch. Its preferred habitat ranges from rainforest to desert terrains. Its ecological role and status resembles that of the similarly-sized cougar in the Americas.

Etymology

In antiquity, it was believed that a leopard was a hybrid of a lion and a panther, as is reflected in its name, a Greek portmanteau derived from λέων léon ("lion") and πάρδος párdos ("male panther"), the latter related to Sanskrit पृदाकु pṝdāku ("snake, tiger, panther").

A panther can be any of several species of large felid; in North America, the term refers to cougars; in South America, jaguars; and everywhere else, it refers to leopards. Early naturalists distinguished between leopards and panthers not by colour (a common misconception), but by the length of the tail—panthers having longer tails than leopards.

Felis pardus was one of the many species described in Linnaeus's 18th-century work, Systema Naturae.

The generic component of its modern scientific designation, Panthera pardus, is derived from Latin via Greek πάνθηρ pánthēr. A folk etymology held that it was a compound of παν pan ("all") and θηρ ("beast"). However, it is believed instead to derive from an Indo-Iranian word meaning "whitish-yellow, pale"; in Sanskrit, this word's reflex was पाण्डर pāṇḍara, from which was derived पुण्डरीक puṇḍárīka ("tiger", among other things), then borrowed into Greek.

Taxonomy

Like the rest of the feline family, the Panthera genus has been subject to much alteration and debate and the exact relations between the four species (as well as the clouded leopard and snow leopard) have not been effectively resolved. DNA evidence shows that the lion, tiger, leopard, jaguar, snow leopard, and clouded leopard share a common ancestor nearly 11 million years ago (Ma)—the basal divergence amongst the Felidae family. The fossil record points to the emergence of Panthera just two to 3.8 million years ago.

In a mitochondrial DNA study, Yu and Zhang (2005) suggest that the leopard is most closely related to the snow leopard, and go so far as placing the latter as a fifth species of Panthera, P. uncia. Canonical works, such as the Mammal Species of the World, continue to list the snow leopard as the only species within its genus, Uncia uncia, but this could change; Johnson et al. (2006) support the placement of the snow leopard within Panthera. They suggest, however, that the snow leopard is most closely aligned with the tiger. The leopard is held to have diverged from the Panthera lineage subsequent to these two species, but before the lion and jaguar. Older research has tended to suggest that the leopard is most closely related to the lion and/or the jaguar. As recently as 2001, it was held to have split along with the lion in a phylogenetic analysis of chemical secretions amongst cats.

Panthera is believed to have emerged in Asia, with ancestors of the leopard and other cats subsequently migrating into Africa. Fossil evidence of leopard ancestors has been found from 2 to 3.5 Ma. These Pleistocene specimens resemble primitive jaguars. The modern leopard type is suggested to have evolved in Africa 470 000–825 000 years ago and radiated across Asia 170 000–300 000 years ago.

Subspecies

As many as 27 leopard subspecies were once suggested, the number growing from the time of Linnaeus in the 18th century to that of Reginald Pocock in the early 20th. In 1996, Miththapala et al. revised this downward to just eight subsepecies based on DNA analysis. Uphyrina et al. would concur in 2001 but split out a ninth separately, the Arabian leopard (P. pardus nimr). The latter researchers note the number might be an underestimation because of limited sampling of African leopards. Their list as follows:

Indo-Chinese leopard (P. pardus delacouri) in Mainland Southeast Asia.
Indian leopard (P. pardus fusca) in India, southeastern Nepal, and northern Bangladesh.
North Chinese leopard (P. pardus japonensis) in China.
Sri Lankan leopard (P. pardus kotiya) in Sri Lanka.
Javan leopard (P. pardus melas) in Java.
Amur leopard (P. pardus orientalis) in the Russian Far East, northern China, and Korea.
African leopard (P. pardus pardus) in Africa.
Persian leopard or Iranian leopard (P. pardus saxicolor) in Southwest Asia.
Arabian leopard (P. pardus nimr) in Arabian Peninsula.

Older taxonomic divisions

Included in the African leopard (P. pardus pardus):

Barbary leopard (P. pardus panthera)
Cape leopard (P. pardus melanotica)
Central African leopard (P. pardus shortridgei)
Congo leopard (P. pardus ituriensis)
East African leopard (P. pardus suahelica)
Eritrean leopard (P. pardus antinorii)
Somalian leopard (P. pardus nanopardus)
Ugandan leopard ((P. pardus chui)
West African leopard (P. pardus reichinowi)
West African forest leopard (P. pardus leopardus)
Zanzibar leopard (P. pardus adersi)

Included in the Persian leopard (P. pardus saxicolor):

Anatolian leopard (P. pardus tulliana)
Baluchistan leopard (P. pardus sindica)
Caucasus leopard (P. pardus ciscaucasica)
Central Persian leopard (P. pardus dathei)
Sinai leopard (P. pardus jarvisi)

Included in the Indian leopard (P. pardus fusca):

Kashmir leopard (P. pardus millardi)
Nepal leopard (P. pardus pernigra)

Physical characteristics

The leopard is an agile and stealthy predator. Although smaller than the other members of the Panthera genus, the leopard is still able to take large prey given a massive skull that well utilizes powerful jaw muscles. Its body is comparatively long for a cat and its legs are short. Head and body length is between 90 and 190 cm (35 and 75 in), the tail reaches 60 to 110 cm (24 to 43 in). Shoulder height is 45 to 80 cm (18-31 in). Males are considerably larger than females and weigh 37 to 91 kg (82 to 200 lbs) compared to 28 to 60 kg (62 to 132 lbs) for females. The larger-bodied populations of leopard (such as the Javan leopard and the leopards from the forested mountains and tropical rainforests of Africa) are generally found in areas isolated from competing large predators, especially from dominant big cats like lions and tigers.

One of many spotted cats, a leopard may be mistaken for a cheetah or a jaguar (though, of course, there is no range overlap with the latter). The leopard has rosettes rather than cheetah's simple spots, but they lack internal spots, unlike the jaguar. The leopard is larger and less lanky than the cheetah but smaller than the jaguar. The leopard's black, irregular rosettes serve as camouflage. They are circular in East Africa but tend to be square-shaped in southern Africa.

Leopards have been reported to reach 21 years of age in captivity.

Black leopards

A melanistic morph of the leopard occurs, particularly in mountainous areas and rain forests. The black color is heritable and caused by recessive gene loci. (While they are commonly called black panthers, the term is not exclusive to leopards; it also applies to melanistic jaguars.)

Melanistic leopards are particularly common on the Malayan Peninsula: early reports suggested up to half of all leopards there are black, but a 2007 camera-trap study in Taman Negara National Park found that all specimens were melanistic. Although the benefits of melanism are difficult to interpret, it may serve as camouflage in the rainforest habitat. (Genetic research has found four independent origins for melanism in cats, suggesting that there must be some adaptive advantage.) Another possibility is that the color variation was relic adaptation to an epidemic; genes causing melanism can also affect the immune system.

In Africa, black leopards are much less common as melanism is not an adaptive advantage on the savanna: dark coloration provides poor camouflage and makes hunting difficult. Estimates are as low as one in 80 or 100. In the dense forests of the Ethiopian Highlands, however, the black leopard is much more common than in Africa generally; as many as one in five leopards may be melanistic.

Biology and behavior

The leopard is known for its ability in climbing, and it has been observed resting on tree branches during the day and descending from trees headfirst. It is a powerful swimmer, although, not as strong as some other big cats, such as the tiger. The leopard is also very agile, and can run over sixty kilometres an hour, leap over six metres and jump up to three metres vertically. The leopard is primarily a nocturnal creature, and many of its operations are done by night. However, there have been recorded instances of leopards hunting during the light, especially when the sky is overcast. It spends much of its day resting and sleeping, up in the branches of trees, underneath rocks or in the grass.

Diet and hunting

Leopards are opportunistic hunters. Although mid-sized animals are preferred, the leopard will eat anything from dung beetles to male giant elands. Their diet consists mostly of ungulates and monkeys, but rodents, reptiles, amphibians, birds and fish are also eaten. In Africa, mid-sized antelopes provide a majority of the leopard's prey, especially impala and Thomson's gazelles. In Asia the leopard preys on deer such as chitals and muntjacs as well as various Asian antelopes and Ibex. One study at the Wolong Reserve in China revealed how adaptable the leopard's hunting behaviour is: over the course of seven years vegetative cover receded, and the animals opportunistically shifted from primarily consuming tufted deer to instead pursuing bamboo rats and other smaller prey.

The leopard stalks its prey silently and at the last minute pounces on its prey and strangles its throat with a quick bite. Leopards often hide their kills in dense vegetation or take them up trees, and are capable of carrying animals up to three times their own weight this way.

One survey of nearly 30 research papers found preferred prey weights of 10 to 40 kg (22-88 lb), with most preferred. Along with impala and chital, a preference for bushbuck and common duiker was found. Other prey selection factors include a preference for prey in small herds, in dense habitat, and those that afford the predator a low risk of injury.

Reproduction

A male may follow a female who catches his attention. Eventually, a fight for reproductive rights may take place. Depending on the region, leopards may mate all year round (Asia and Africa) or seasonally during January to February (Manchuria and Siberia). The estrous cycle lasts about 46 days and the female usually is in heat for 6–7 days. Gestation lasts for 90-105 days. Cubs are usually born in a litter of 2–4, but infant mortality is high and usually no more than 1–2 cubs survive beyond their infancy. The pregnant females find a cave, crevice among boulders, hollow tree, or thicket to give birth and make a den. Cubs open their eyes after a period of 10 days. The fur of the young tends to be longer and thicker than that of adults. Their pelage is also more gray in color with less defined spots. Around three months the infants begin to follow the mother out on hunts. At one year of age leopard young can probably fend for themselves but they remain with the mother for 18–24 months.

Social structure and home range

Studies of leopard home range size have tended to focus on protected areas, which may have led to skewed data; as of the mid-1980s, only 13% of the leopard range actually fell within a protected area. In their IUCN survey of the literature, Nowell and Jackson suggest male home territories vary between 30-78 square kilometers, but just 15-16 km² for females. Research in a conservation area in Kenya shows similar territory sizes and sex differential: 32.8 km² ranges for males, on average, and 14 km² for females. In Nepal, somewhat larger male ranges have been found at about 48 km², while female ranges are in-keeping with other research, at 17 km²; female home territories were seen to decrease to just five to seven km² when young cubs were present, while the sexual difference in range size seemed to be in positive proportion to overall increase. However, significant variations in size of home territories have been suggested across the leopard's range. In Namibia, for instance, research that focussed on spatial ecology in farmlands outside of protected areas found ranges that were consistently above 100 km², with some more than 300 km²; admitting that their data were at odds with others', the researchers also suggested little or no sexual variation in the size of territories. Virtually all sources suggest that males do have larger ranges. There seems to be little or no overlap in territory amongst males, although overlap exists between the sexes; one radio-collar analysis in the Ivory Coast found a female home range completely enclosed within a male's.

The leopard is solitary and, aside from mating, interactions between individuals appear to be infrequent. Aggressive encounters have been observed, however. Two of five males studied over a period of a year at a game reserve in South Africa died, both violently. One was initially wounded in a male-male territorial battle over a carcass; taken in by researchers, it was released after a successful convalescence only to be killed by a different male a few months later. A second was killed by another predator, possibly a spotted hyena. A third of the five was badly wounded in intraspecific fighting, but recovered.

Ecology

Distribution and habitat

Data from 1996 found that the leopard has the largest distribution of any wild cat, occurring most in certain parts of southern Asia and widely in eastern and central Africa, although populations before and since have shown a declining trend and are fragmented outside of subsaharan Africa. The IUCN notes that within sub-Saharan Africa the species is "still numerous and even thriving in marginal habitats" where other large cats have disappeared, but that populations in North Africa may be extinct. In Asia, data on distribution is not consistent: populations in Southwest and Central Asia are small and fragmented; in the northeast portion of the range, they are critically endangered; but in the Indian subcontinent, Southeast Asia, and China, the leopard is still relatively abundant.

Leopards live mainly in grasslands, woodlands and riverside forests. The animal has primarily been studied in open savannah habitats, which may have biased common descriptions. It is generally considered nocturnal, for instance, but radio-tracking and scat analysis in West Africa has found that rainforest leopards are more likely to be diurnal and crepuscular. Forest leopards are also more specialized in prey selection and exhibit seasonal differences in activity patterns. While associated with the savanna and rainforest, the leopard is exceptionally adaptable: in the Russian Far East, the animal inhabits temperate forests where winter temperatures reach a low of –25 °C.

Ecological role

Because of their wide habitat range, leopards must compete for food and safety with other large predators such as lions, tigers, spotted hyenas and wild dogs. These competitors sometimes may steal the leopard's kill or devour its young. A single lion or tiger is capable of killing an adult leopard. Leopards have adapted to live alongside these other predators by hunting at different times of the day, and by avoiding areas frequented by them. In search of safety, the leopard will often stash its young or a recent kill high up in a tree. Lions are occasionally successful in climbing trees and fetching leopard kills, and if motivated, an adult tiger might also scale a tree to acquire food.

Nowell and Jackson note that resource portioning occurs where the leopard shares range with the lion or tiger: the leopard tends to take smaller prey (usually less than 75 kg) where its large feline cousins are present. One tropical forest study suggests that leopards do not always avoid the larger cats by hunting at different times. With relatively abundant prey, tigers and leopards were seen to successfully coexist without competitive exclusion or inter-species dominance hierarchies that may be more common to the savanna.

Variant coloration

A pseudo-melanistic leopard has a normal background color, but its excessive markings have coalesced so that its back seems to be entirely black. In some specimens, the area of solid black extends down the flanks and limbs; only a few lateral streaks of golden-brown indicate the presence of normal background colour. Any spots on the flanks and limbs that have not merged into the mass of swirls and stripes are unusually small and discrete, rather than forming rosettes. The face and underparts are paler and dappled like those of ordinary spotted leopards. These melanistic leopards are often incorrectly referred to by the general population as "black panthers".

In a paper about panthers and ounces of Asia, Reginald Innes Pocock used a photo of a leopard skin from southern India; it had large black-rimmed blotches, each containing a number of dots and it resembled the pattern of a jaguar or clouded leopard. Another of Pocock's leopard skins from southern India had the normal rosettes broken up and fused and so much additional pigment that the animal looked like a black leopard streaked and speckled with yellow.

Most other colour morphs of leopards are known only from paintings or museum specimens. There have been very rare examples where the spots of a normal black leopard have coalesced to give a jet black leopard with no visible markings. Pseudo-melanism (abundism) occurs in leopards. The spots are more densely packed than normal and merge to largely obscure the background colour. They may form swirls and, in some places, solid black areas. Unlike a true black leopard the tawny background colour is visible in places. One pseudo-melanistic leopard had a tawny orange coat with coalescing rosettes and spots, but white belly with normal black spots (like a black-and-tan dog).

A 1910 description of a pseudo-melanistic leopard:

Another pseudo-melanistic leopard skin was described in 1915 by Holdridge Ozro Collins who had purchased it in 1912. It had been killed in Malabar, India that same year.

In May 1936, the British Natural History Museum exhibited the mounted skin of an unusual Somali leopard. The pelt was richly decorated with an intricate pattern of swirling stripes, blotches, curls and fine-line traceries. This is different from a spotted leopard, but similar to a king cheetah hence the modern cryptozoology term king leopard. Between 1885 and 1934, six pseudo-melanistic leopards were recorded in the Albany and Grahamstown districts of South Africa. This indicated a mutation in the local leopard population. Other king leopards have been recorded from Malabar in southwestern India. Shooting for trophies may have wiped out these populations.

Hybrids

A pumapard is a hybrid animal resulting from a union between a leopard and a puma. Three sets of these hybrids were bred in the late 1890s and early 1900s by Carl Hagenbeck at his animal park in Hamburg, Germany. Most did not reach adulthood. One of these was purchased in 1898 by Berlin Zoo. A similar hybrid in Berlin Zoo purchased from Hagenbeck was a cross between a male leopard and a female puma. Hamburg Zoo's specimen was the reverse pairing, the one in the black and white photo, fathered by a puma bred to an Indian leopardess. Whether born to a female puma mated to a male leopard, or to a male puma mated to a female leopard, pumapards inherit a form of dwarfism. Those reported grew to only half the size of the parents. They have a puma-like long body (proportional to the limbs, but nevertheless shorter than either parent), but short legs. The coat is variously described as sandy, tawny or greyish with brown, chestnut or "faded" rosettes.

Leopards and humans

Leopards have been known to humans since antiquity and have featured in the art, mythology and folklore of many countries where they have historically occurred, such as ancient Greece, Persia and Rome, as well as some where they have not existed since for several millenniums, such as in England. The modern use of the leopard as an emblem for sport or a coat of arms is much more restricted to Africa, though numerous products worldwide have used the name.

Leopards and humans have many relations, involving tourism, heraldry and modern culture. Leopard domestication has also been recorded - several leopards were kept in a menagerie established by King John at the Tower of London in the 13th century; in around 1235 three of these animals were given to Henry III by the Holy Roman Emperor Frederick II.

Tourism

Despite its size, this largely nocturnal and arboreal predator is difficult to see in the wild. The best location to see leopards in Africa is in the Sabi Sand Private Game Reserve in South Africa, where leopards are habituated to safari vehicles and are seen on a daily basis at very close range. In Asia, one can see leopards at Yala National Park in Sri Lanka, which has one of the world's highest densities of wild leopards, but even here sightings are by no means guaranteed because more than half the park is closed to the public, allowing the animals to thrive. Another good destination for leopard watching is the recently reopened Wilpattu National Park, also in Sri Lanka. In India leopards are found all over the country and this wide distribution leads to maximum man-animal conflict. Among the best places to observe leopards in India are national parks in Madhya Pradesh and Uttarakhand.

Heraldry

The lion passant guardant or "leopard" is a frequently used charge in heraldry, most commonly appearing in groups of three. The heraldric leopard lacks spots and sports a mane, making it visually almost identical to the heraldric lion, and the two are often used interchangeably. These traditional lion passant guardants appear in the coat of arms of England and many of its former colonies; more modern naturalistic (leopard-like) depictions appear on the coat of arms of several African nations including Benin, Malawi, Somalia, Democratic Republic of the Congo and Gabon which uses a black panther.

Man-eating

Despite being predators of man's hominid ancestors, most leopards will tend to avoid humans. Still, people are occasionally targeted as prey. Most healthy leopards prefer wild prey to humans, but cats who are injured, sickly or struggling with a shortage of regular prey often turn to hunting people and may become habituated to it. In the most extreme cases, both in India, a leopard dubbed "the Leopard of Rudraprayag" is claimed to have killed over 125 people and the infamous leopard called "Panar Leopard" killed over 400 after being injured by a poacher and thus being made unable to hunt normal prey. The "Leopard of Rudraprayag" and the "Panar Leopard" were both killed by the famed hunter Jim Corbett. Man-eating leopards are considered bold by feline standards and commonly enter human settlements for prey, more so than their lion and tiger counterparts. Kenneth Anderson, who had first hand experience with many man-eating leopards, described them as far more threatening than tigers:

Because they can subsist on small prey and are less dependent on large prey, leopards are less likely to turn to man-eating than either lions or tigers. However, leopards might be attracted to human settlements by livestock or pets, especially domestic dogs, and they may resort to the eating of humans should conditions demand it, and no other food is available.

Notes

References

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Leopards and spots on ears and tail
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