Leopard

The leopard (Panthera pardus) is one of the five "big cats" in the genus Panthera. It is a member of the family Felidae with a wide range in regions of sub-Saharan Africa, West Asia, the Middle East, South and Southeast Asia to Siberia.^[2] Fossil records found in Italy suggest that in the Pleistocene it ranged as far as Europe.^[3]

Compared to other members of Felidae, the leopard has relatively short legs and a long body with a large skull. It is similar in appearance to the jaguar, but is smaller and more lightly built. Its fur is marked with rosettes similar to those of the jaguar, but the leopard's rosettes are smaller and more densely packed, and do not usually have central spots as the jaguar's do. Both leopards and jaguars that are melanistic are known as black panthers.

The leopard's success in the wild is due to its well camouflaged fur; its opportunistic hunting behaviour, broad diet, and strength to move heavy carcasses into trees; its ability to adapt to various habitats ranging from rainforest to steppe and including arid and montane areas; and to run at speeds up to 58 kilometres per hour (36 mph).^[4]

It is listed as Near Threatened on the IUCN Red List because leopard populations are declining in large parts of their range. They are threatened by habitat loss and pest control. Their habitats are fragmented and they are illegally hunted so that their pelts may be sold in wildlife trade for medicinal practices and decoration.^[5] They have been extirpated in Hong Kong, Singapore, Kuwait, Syria, Libya, Tunisia and most likely Morocco.^[1]

Etymology

In antiquity, a leopard was believed to be a hybrid of a lion and a panther, as is reflected in its name, which is a Greek compound of λέων leōn (lion) and πάρδος pardos (male panther). The Greek word is related to Sanskrit पृदाकु pṛdāku (snake, tiger, panther), and probably derives from a Mediterranean language, such as Egyptian.^[6][7]

A panther can be any of several species of large felids: the term can refer to cougars and jaguars in the American continents but (given the European origins of the word) it is largely thought to define the leopard at its source. Black panther refers to leopards with melanistic genes, which are not uncommon in rainforest habitats.^[8]

The generic component of its modern scientific designation, Panthera pardus, derives from Latin via Greek πάνθηρ (pánthēr).^[9] Folk etymology saw the name as a compound of παν (pan, all) and θηρ (beast).^[10] However, it is believed instead to be derived from an Indo-Iranian word meaning "white-yellow, pale"; in Sanskrit, this word's reflex was पाण्डर pāṇḍara, which was derived from पुण्डरीक puṇḍárīka (tiger, among other things), then borrowed into Greek.^[7][9]

Characteristics

Leopards show a great diversity in coat colour and rosettes patterns. In general, the coat colour varies from pale yellow to deep gold or tawny, and is patterned with black rosettes. The head, lower limbs and belly are spotted with solid black. Coat colour and patterning are broadly associated with habitat type. Their rosettes are circular in East Africa but tend to be squarer in southern Africa and larger in Asian populations. Their yellow coat tends to be more pale and cream coloured in desert populations, more gray in colder climates, and of a darker golden hue in rainforest habitats. Overall, the fur under the belly tends to be lighter coloured and of a softer, downy type. Solid black spots in place of open rosettes are generally seen along the face, limbs and underbelly.^[4]

Leopards are agile and stealthy predators. Although they are smaller than most other members of the genus Panthera, they are able to take large prey due to their massive skulls that facilitate powerful jaw muscles. Head and body length is usually between 90 and 165 cm (35 and 65 in). The tail reaches 60 to 110 cm (24 to 43 in) long, around the same length as the tiger's tail and proportionately long for the genus (though snow leopards and the much smaller marbled cats have relatively longer tails).^[11][12] Shoulder height is from 45 to 80 cm (18 to 31 in). The muscles attached to the scapula are exceptionally strong, which enhance their ability to climb trees. They are very diverse in size. Males are about 30% larger than females, weighing 30 to 91 kg (66 to 201 lb) compared to 23 to 60 kg (51 to 132 lb) for females. Large males of up to 91 kg (201 lb) have been documented in Kruger National Park in South Africa; however, males in South Africa's coastal mountains average 31 kg (68 lb) and the females from the desert-edge in Somalia average 23 to 27 kg (51 to 60 lb). This wide variation in size is thought to result from the quality and availability of prey found in each habitat.

The leopard's body is comparatively long, and its legs are short.^[13] The largest verified leopard weighed 96.5 kg (213 lb) and reached 190 cm (75 in) in head-and-body length. Larger sizes have been reported but are generally considered unreliable.^[12][14]

Variant colouration

Melanistic leopards are commonly called black panthers, a term that also applies to melanistic jaguars. Pseudomelanism (abundism) also occurs in leopards.^[15] Melanism in leopards is inherited as a Mendelian, single gene recessive trait relative to the spotted form.^[16] Pairings of black animals inter se have a significantly smaller litter size than other possible pairings.^[17]

The black panther is common in the equatorial rainforest of Malaya and the tropical rainforest on the slopes of some African mountains such as Mount Kenya.^[18] Between January 1996 and March 2009, Indochinese leopards were photographed at 16 sites in the Malay Peninsula in a sampling effort of more than 1000 trap nights. Of 445 photographs of melanistic leopards taken, 410 came from study sites south of the Kra Isthmus, where the non-melanistic morph was never photographed. These data suggest the near fixation of the dark allele in the region. The expected time to fixation of this recessive allele due to genetic drift alone ranged from about 1,100 years to about 100,000 years.^[19][20]

Melanism in leopards has been hypothesized to be causally associated with a selective advantage for ambush.^[21]

A rare "strawberry" leopard was photographed in South Africa's Madikwe Game Reserve.^[22] It is thought the leopard has erythrism, a little-understood genetic condition that causes either an overproduction of red pigments or an underproduction of dark pigments.

Similar species

Leopards may sometimes be confused with two other large spotted cats, the cheetah, with which it may co-exist in Africa, and the jaguar, a neotropical species that it does not naturally co-exist with. However, the patterns of spots in each are different: the cheetah has simple black spots, evenly spread; the jaguar has small spots inside the polygonal rosettes; while the leopard normally has rounder, smaller rosettes than those of the jaguar. The cheetah has longer legs and a thinner build that makes it look more streamlined and taller but less powerfully built than the leopard. The jaguar is more similar in build to the leopard but is generally larger in size and has a more muscular, bulky appearance.^[23]

Distribution and habitat

Leopards have the largest distribution of any wild cat, occurring widely in Africa as well as eastern and southern Asia, although populations have shown a declining trend and are fragmented outside of sub-Saharan Africa. Within sub-Saharan Africa, the species is still numerous and even thriving in marginal habitats where other large cats have disappeared. Populations in North Africa may be extinct.^[4] Data on their distribution in Asia are not consistent. Populations in southwest and central Asia are small and fragmented; in the northeast, they are critically endangered. In the Indian subcontinent, Southeast Asia, and China, leopards are still relatively abundant. Of the species as a whole, its numbers are greater than those of other Panthera species, all of which face more acute conservation concerns.^[1][25]

Leopards are exceptionally adaptable, although associated primarily with savanna and rainforest. Populations thrive anywhere in the species range where grasslands, woodlands, and riverine forests remain largely undisturbed. In the Russian Far East, they inhabit temperate forests where winter temperatures reach a low of −25 °C (−13 °F).^[26] They are equally adept surviving in some of the world's most humid rainforests and even semi-arid desert edges.

Leopards in west and central Asia try to avoid deserts, areas with long-duration snow cover and areas that are near urban development.^[25] In India, leopard populations sometimes live quite close to human settlements and even in semi-developed areas.^[27] Although occasionally adaptable to human disturbances, leopards require healthy prey populations and appropriate vegetative cover for hunting for prolonged survival and thus rarely linger in heavily developed areas.^[25][27] Due to the leopard's superlative stealthiness, people often remain unaware that big cats live in nearby areas.^[27]

Distribution of subspecies

Since Carl Linnaeus published his description of a leopard in the Systema Naturae in 1758, as many as 27 leopard subspecies were subsequently described by naturalists from 1794 to 1956. Since 1996, according to DNA analysis carried out in the 1990s, only eight subspecies are considered valid.^[28] Later analysis revealed a ninth valid subspecies, the Arabian leopard. Because of limited sampling of African leopards, this number might be an underestimation.^[26]

The nine subspecies recognised by IUCN are:^[1][26][29]

Subspecies of Leopard

Subspecies	Description	Image
African leopard (P. p. pardus)	Lives in Sub-Saharan Africa. It is the most widespread subspecies of leopards. North Africa: extinct in Algeria, Egypt, Libya,and Tunisia with relict population in Morocco West Africa: Benin, Burkina Faso, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Mali, Mauritania, Niger, Nigeria, Senegal, Togo East Africa: Burundi, Djibouti, Eritrea, Ethiopia, Kenya, Tanzania, Somalia, South Sudan, Sudan, Rwanda, Uganda Central Africa: Cameroon, Central African Republic, Chad, Democratic Republic of the Congo, Gabon, Republic of the Congo, Southern Africa: Angola, Botswana, Malawi, Mozambique, Namibia, South Africa, Swaziland, Tanzania, Zambia, and Zimbabwe
Indian leopard (P. p. fusca)	Native to the Indian Subcontinent. It is widespread in India, Nepal, Bangladesh and Bhutan. South Asia: Bangladesh, Bhutan, India, Nepal, and Pakistan; extinct in China
Arabian leopard (P. p. nimr), also known as Erythrean leopard	Native to the Arabian Peninsula. It lives in arid areas of Saudi Arabia, Israel, Jordan, and the United Arab Emirates. It is the smallest leopard subspecies. West Asia: Israel, Kuwait, Lebanon, Jordan, Oman, Saudi Arabia, Syria, United Arab Emirates, and Yemen; extinct in the Sinai Peninsula of Egypt
Persian leopard (P. p. saxicolor), also known as Central Asian leopard or Caucasian leopard	Inhabits the Caucasus, Turkmenistan, Afghanistan, and northern Iran.[29] It is the largest leopard subspecies. West Asia: Armenia, Azerbaijan, Iran, Iraq, Georgia, Russia, and Turkey Central Asia: Turkmenistan and Uzbekistan South Asia: Afghanistan and Pakistan
North-Chinese leopard (P. p. japonensis), also simply known as the Chinese leopard	Only native to central to northern China. It is among the medium-sized leopard subspecies. East Asia: China
Amur leopard (P. p. orientalis), also known as Far Eastern leopard or Siberian leopard	Found today only in the cold regions of Russian Far East and Northeast China. It is the most Critically Endangered leopard subspecies, and one of the most endangered animals in the world. It is currently extinct in the Korean Peninsula. North Asia: China and Siberia; extinct in North Korea and South Korea
Indochinese leopard (P. p. delacouri), also known as South-Chinese leopard	Widespread in mainland Southeast Asia and South China. Southeast Asia: China, Cambodia, Laos, Malaysia, Myanmar, Thailand, Vietnam
Javan leopard (P. p. melas)	The only subspecies native to Indonesia. It is found in the Indonesian Island of Java. It is among the most Critically Endangered leopard subspecies. Southeast Asia: Indonesia
Sri Lankan leopard (P. p. kotiya)	Found only in Sri Lanka. South Asia: Sri Lanka

A morphological analysis of characters of leopard skulls implies the validity of two more subspecies:^[29]

• Anatolian leopard (P. p. tulliana) (Valenciennes, 1856) inhabits Western Turkey
• Balochistan leopard (P. p. sindica) (Pocock, 1930) inhabits Pakistan, and possibly also parts of Afghanistan and Iran

The following African leopard populations used to be considered subspecies until 1996:^[26][28]

• Barbary leopard (P. p. panthera) (Schreber, 1777)
• Sinai leopard (P. p. jarvasi) (Pocock, 1932)
• Zanzibar leopard (P. p. adersi) (Pocock, 1932)

The smallest leopard subspecies is the Arabian leopard. Adult females weigh as little as 18 kg (40 lb).^[30] Large subspecies, in which males weigh up to 91 kg (201 lb), are the Sri Lankan leopard and the Persian leopard. Such larger leopards inhabit areas which lack tigers and lions, so that leopards are at the top of the food chain with no competitive restriction from large prey.^[31]

Ecology and behaviour

Leopards are elusive, solitary, and largely nocturnal.^[32][33] They have primarily been studied in open savanna habitats, which may have biased common descriptions. Activity level varies depending on the habitat and the type of prey that they hunt. Radio-tracking and scat analysis in West Africa showed that rainforest leopards are more likely to be diurnal and crepuscular. Forest leopards are also more specialised in prey selection and exhibit seasonal differences in activity patterns.^[34]

Leopards are known for their ability in climbing, and have been observed resting on tree branches during the day, dragging their kills up trees and hanging them there, and descending from trees headfirst.^[35] They are powerful swimmers, although are not as disposed to swimming as some other big cats, such as the tiger. They are very agile, and can run at over 58 kilometres per hour (36 mph), leap over 6 metres (20 ft) horizontally, and jump up to 3 metres (9.8 ft) vertically.^[36] They produce a number of vocalizations, including grunts, roars, growls, meows, and purrs.^[37]

Social structure and home range

Home ranges of male leopards vary between 30 km² (12 sq mi) and 78 km² (30 sq mi), and of females between 15 to 16 km² (5.8 to 6.2 sq mi).^[4] Virtually all sources suggest that males do have larger home ranges. There seems to be little or no overlap in territory among 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.^[39]

Research in a conservation area in Kenya showed similar territory sizes and sex differential: 32.8 km² (12.7 sq mi) average ranges for males, and 14 km² (5.4 sq mi) for females.^[40]

In Nepal, somewhat larger male ranges have been found at about 48 km² (19 sq mi), while female ranges at 17 km² (6.6 sq mi); female home ranges decreased to 5 to 7 km² (1.9 to 2.7 sq mi) when young cubs were present, while the sexual difference in range size seemed to be in positive proportion to overall increase.^[41]

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. However, significant variations in the size of home ranges have been suggested across the leopard's range. Research in Namibia that focused on spatial ecology in farmlands outside of protected areas revealed ranges that were consistently above 100 km² (39 sq mi) with some more than 300 km² (120 sq mi). Admitting that their data were at odds with others, the researchers found little or no sexual variation in the size of territories.^[42]

Aggressive encounters have been observed. 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.^[32]

Hunting and diet

Leopards are versatile, opportunistic hunters, and have a very broad diet. They feed on a greater diversity of prey than other members of the genus Panthera, and are reported to eat anything from dung beetles to common elands, though medium-sized prey species in the 20–80 kg (44–176 lb) range are usually taken.^[4] The largest prey reported killed by a leopard was a 900 kg (2,000 lb) male eland.^[43] although leopards generally do not prey on such large animals.^[44] Their diet consists mostly of ungulates, followed by primates, primarily monkeys of various species, including the Vervet monkey. However, they will also opportunistically eat rodents, reptiles, amphibians, insects, birds (especially ground-based types like the vulturine guineafowl), fish and sometimes smaller predators (such as foxes, jackals, martens and smaller felid species). In at least one instance, a leopard has predated a sub-adult Nile crocodile that was crossing over land.^[45] Leopards are the only natural predators of adult chimpanzees and gorillas, although the cat may sometimes choose to avoid these as they are potentially hazardous prey, especially large male silverback gorillas.^[46][47] They stalk their prey silently, pounce on it at the last minute, and strangle its throat with a quick bite. In Africa, mid-sized antelopes provide a majority of their prey, especially impala and Thomson's gazelles.^[48]

In the open savanna of Tsavo National Park, they kill most of their prey while hunting between sunset and sunrise.^[49] In Kruger National Park, males and females with cubs are more active at night. At least 92 prey species have been documented in their diet. They focus their hunting activity on locally abundant medium-sized ungulate species in the 20 to 80 kg (44 to 176 lb) range, while opportunistically taking other prey. Analysis of leopard scats found that 67% contained ungulate remains, of which 60% were impala, the most abundant antelope, with adult weights of 40 to 60 kg (88 to 132 lb). Small mammal remains were found most often in scats of sub-adult leopards, especially females. Average daily consumption rates was estimated at 3.5 kg (7.7 lb) for adult males and 2.8 kg (6.2 lb) for females.^[50]

In Asia, the leopard primarily preys on deer such as chitals and muntjacs, as well as various Asian antelopes and ibex. Prey preference estimates in southern India showed that the most favored prey of the leopard were chitals.^[51] A study at the Wolong Reserve in China revealed how adaptable their hunting behaviour is. Over the course of seven years, the vegetative cover receded, and the animals opportunistically shifted from primarily consuming tufted deer to pursuing bamboo rats and other smaller prey.^[52]

They select their prey focusing on small herds, dense habitat, and low risk of injury, preferring prey weights of 10 to 40 kg (22 to 88 lb) such as impala, chital, bushbuck, and common duiker with an average body weight of 25 kg (55 lb).^[53]

In search of safety, leopards often stash their young or recent kills high up in a tree, which can be a great feat of strength considering that they may be carrying prey heavier than themselves in their mouth while they climb vertically.^[48] One leopard was seen to haul a young giraffe, estimated to weigh up to 125 kg (276 lb), more than twice the weight of the cat, up 5.7 m (19 ft) into a tree.^[49]

Interspecific predatory relationships

Leopards must compete for food and shelter with other large predators such as tigers, lions, spotted hyenas, striped hyenas, brown hyenas, up to five species of bear and both African and Asiatic wild dogs. These animals may steal the leopard's kill, devour its young or even kill adult leopards. Leopards co-exist alongside these other large predators by hunting for different types of prey and by avoiding areas frequented by them. Leopards may also retreat up a tree in the face of direct aggression from other large carnivores but leopards have been seen to either kill or prey on competitors such as black-backed jackal, African wild cat and the cubs of lions, cheetahs, hyenas, and wild dogs.^[4]

Resource partitioning occurs where leopards share their range with tigers. Leopards tend to take smaller prey, usually less than 75 kg (165 lb), where tigers are present.^[4] In areas where the leopard is sympatric with the tiger, coexistence is reportedly not the general rule, with leopards being few where tigers are numerous.^[54] Mean leopard density decreased significantly (from 9.76 animals/100km2 to 2.07 animals/100km2) when the mean density of tigers increased (from 3.31 animals/100km2 to 5.81 animals/100km2) from 2004-5 to 2007-8 in the Rajaji National Park in India following the relocation of pastoralists out of the park.^[55] There, the two species have high dietary overlap, and an increase in the tiger population resulted in a sharp decrease in the leopard population and a shift in the leopard diet to small prey (from 9% to 36%) and domestic prey (from 6.8% to 31.8%).^[55] In the Primore region of the Russian Far East, Amur leopards were absent or very rarely encountered at places where Siberian tigers reside.^[56] However, in the Chitwan National Park in Nepal, both species coexist because there is a large prey biomass, a large proportion of prey is of the smaller sizes, and dense vegetation exists.^[57] Here leopards killed prey ranging from less than 25 kg (55 lb) to 100 kg (220 lb) in weight with most kills in the 25–50 kg (55–110 lb) range; tigers killed more prey in the 50–100 kg (110–220 lb) range.^[57] There were also differences in the microhabitat preferences of the individual tiger and leopard followed over a 5-month (December to April) period in this study - the tiger used roads and (except in February) forested areas more frequently, while the leopard used recently burned areas and open areas more frequently ^[58] Usually when a tiger began to kill baits at sites formerly frequented by leopards, the leopards would no longer come and kill there.^[59] In the tropical forests of India’s Nagarhole National Park, tigers selected prey weighing more than 176 kg (388 lb), whereas leopards selected prey in the 30–175 kg (66–386 lb) range.^[60] In tropical forest they do not always avoid the larger cats by hunting at different times. With relatively abundant prey and differences in the size of prey selected, tigers and leopards seem to successfully coexist without competitive exclusion or inter-species dominance hierarchies that may be more common to the leopard's co-existence with the lion in savanna habitats.^[61] In areas with high tiger populations, such as in the central parts of India’s Kanha National Park, leopards are not permanent residents, but transients. They were common near villages at the periphery of the park and outside the park.^[59]

In some areas of Africa, troops of large baboon species (potential leopard prey themselves) will kill and sometimes eat leopard young if they discover them.^[62] Occasionally, Nile crocodiles may prey on leopards of any age. For example, one large adult leopard was grabbed and consumed by a large crocodile while attempting to hunt along a bank in Kruger National Park.^[63][64][65] Mugger crocodiles may also on rare occasion kill an adult leopard in India.^[66] Lions are occasionally successful in climbing trees and fetching leopard kills,^[48] but leopards are also known to kill or prey on cubs of large lions.^[4] In the Kalahari desert, leopards frequently lose kills to the brown hyena, if the leopard is unable to move the kill into a tree. Single brown hyenas have been observed charging at and displacing male leopards from kills.^[67][68] Burmese pythons have been known to prey on leopards, with an adult cat having been recovered from the stomach of an 5.5 m (18 ft) specimen.^[69]

Two cases of leopards killing cheetahs have been reported in 2014.^[70][71]

Reproduction and life cycle

Depending on the region, leopards may mate all year round. In Manchuria and Siberia, they mate during January and February. The estrous cycle lasts about 46 days and the female usually is in heat for 6–7 days.^[72] Gestation lasts for 90 to 105 days.^[73] Cubs are usually born in a litter of 2–4 cubs.^[74] Mortality of cubs is estimated at 41–50% during the first year.^[50]

Females give birth in a cave, crevice among boulders, hollow tree, or thicket to make a den. Cubs are born with closed eyes, which open four to nine days after birth.^[43] The fur of the young tends to be longer and thicker than that of adults. Their pelage is also more gray in colour with less defined spots. Around three months of age, the young begin to follow the mother on hunts. At one year of age, leopard young can probably fend for themselves, but remain with the mother for 18–24 months.^[35]

The average typical life span of a leopard is between 12 and 17 years.^[75] The oldest recorded Spotted Leopard was a female named Roxanne living in captivity at McCarthy’s Wildlife Sanctuary in The Acreage, Palm Beach County, Florida. She died August 8, 2014 at the age of 24 years, 2 months and 13 days. This has been verified by the Guinness World Records.^[76] Previously, the oldest recorded Leopard was a female named Bertie living in captivity in Warsaw Zoo. She died in December 2010 at the age of 24.^[77] The oldest recorded male Leopard was Cezar, who reached the age of 23. He also lived at Warsaw Zoo and was Bertie's lifelong companion.^[78]

Taxonomy

Like all of the feline family, the genus Panthera 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.

The leopard was among the first animals named under the modern system of biological classification, since it was described by Carl Linnaeus in 1758 in the 10th edition of Systema Naturae. Linnaeus placed the leopard under the genus Felis as the binominal Felis pardus.^[79] In the 18th and 19th centuries, most naturalists and taxonomists followed his example. In 1816, Lorenz Oken proposed a definition of the genus Panthera, with a subgenus Panthera using Linnaeus' Felis pardus as a type species. But most disagreed with his definition, and until the beginning of the 20th century continued using Felis or Leopardus when describing leopard subspecies.^[80] In 1916, Reginald Innes Pocock accorded Panthera generic rank defining Panthera pardus as species.^[81]

It is believed that the basal divergence amongst the family Felidae occurred about 11 million years ago. The last common ancestor of the lion, tiger, leopard, jaguar, snow leopard, and clouded leopard is believed to have occurred about 6.37 million years ago. Panthera is believed to have emerged in Asia, with ancestors of the leopard and other cats subsequently migrating into Africa. The researchers suggest that the snow leopard is most closely aligned with the tiger, whereas the leopard possibly has diverged from the Panthera lineage subsequent to these two species, but before the lion and jaguar.^[82]

Results of phylogenetic analyses of chemical secretions amongst cats has suggested that the leopard is closely related to the lion.^[83] Results of a mitochondrial DNA study carried out later suggest that the leopard is closely related to the snow leopard, which is placed as a fifth Panthera species, Panthera uncia.^[84]

Evolution

Fossils of early leopard ancestors have been found in East Africa and South Asia from the Pleistocene of 2 to 3.5 Ma. The modern leopard is suggested to have evolved in Africa 470,000–825,000 years ago and radiated across Asia 170,000–300,000 years ago.^[26]

In Europe, the leopard is known at least since the Pleistocene. Fossil leopard bones and teeth dating from the Pliocene were found in Perrier in France, northeast of London, and in Valdarno in Italy. At 40 sites in Europe fossil bones and dental remains of leopards dating from the Pleistocene were excavated mostly in loess and caves. The sites of these fossil records range from near Lisbon, near Gibraltar, and Santander Province in northern Spain to several sites in France, Switzerland, Italy, Austria, Germany, in the north up to Derby in England, in the east to Přerov in the Czech Republic, and the Baranya in southern Hungary.^[85] The Pleistocene leopards of Europe can be divided into four subsequent subspecies. The first European leopard subspecies P. p. begoueni is known since the beginning of the early Pleistocene and was replaced about 600,000 years ago by P. p. sickenbergi, which in turn was replaced by P. p. antiqua at around 300,000 years ago. The last form, the Late Pleistocene Ice Age leopard (P. p. spelaea) appeared at the beginning of the Late Pleistocene and survived until about 24,000 years ago in large parts of Europe.^[86]

Leopards and humans

Leopards have been known to humans throughout history, 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 for several millennia, such as 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. During the Benin Empire, the leopard was used to symbolise the power of the king or oba.^[87]

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; around 1235, three of these animals were given to Henry III by Holy Roman Emperor Frederick II.^[88]

Heraldry

The lion passant guardant or leopard is a frequently used charge in heraldry, most commonly appearing in groups of three.^[89] The heraldic leopard lacks spots and sports a mane, making it visually almost identical to the heraldic lion, and the two are often used interchangeably. These traditional lions passant guardant 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, the Democratic Republic of the Congo and Gabon, which uses a black panther.^[90]

Tourism

In protected areas of several countries, wildlife touring programs and safari ventures offer sightings of leopards in their natural habitat. While luxury establishments may boast the fact that wild animals can be seen at close range on a daily basis, the leopard's camouflage and propensity to hide and stalk prey typically make leopard sightings rare.^[91] In Sri Lanka's Yala National Park, leopards have been ranked by visitors to be among the least visible of all animals in the park despite their high concentration in the reserve.^[92]

In South Africa, safaris are offered in the Sabi Sand Game Reserve. In Sri Lanka, wildlife tours are available in the Yala and Wilpattu National Parks. In India, safaris are offered in the Madhya Pradesh and Uttarakhand national parks as well as in the Pali district of western Rajasthan.^[93]

Man-eating

Most leopards avoid people, but humans may occasionally be targeted as prey. Most healthy leopards prefer wild prey to humans, but injured, sickly, or struggling cats or those with a shortage of regular prey may resort to hunting humans and become habituated to it. Although usually slightly smaller than humans, an adult leopard is much more powerful and easily capable of killing them. Two extreme cases occurred in India: the first leopard, "the Leopard of Rudraprayag", killed more than 125 people; the second, the "Panar Leopard", was believed to have killed more than 400. Both were killed by the renowned hunter Jim Corbett.^[94] Man-eating leopards are considered bold and difficult to track by feline standards and may enter human settlements for prey, more so than lions and tigers.^[95] Author and big game hunter Kenneth Anderson had first-hand experience with many man-eating leopards, and described them as far more threatening than tigers:

Although examples of such animals are comparatively rare, when they do occur they depict the panther [leopard] as an engine of destruction quite equal to his far larger cousin, the tiger. Because of his smaller size he can conceal himself in places impossible to a tiger, his need for water is far less, and in veritable demoniac cunning and daring, coupled with the uncanny sense of self-preservation and stealthy disappearance when danger threatens, he has no equal.

— Kenneth Anderson, Nine Man-Eaters and One Rogue, Chapter II "The Spotted Devil of Gummalapur"

There is something very terrifying in the angry grunt of a charging leopard, and I have seen a line of elephants that were staunch to a tiger, turn and stampede from a charging leopard.

— Jim Corbett, The Temple Tiger and More Man-Eaters of Kumaon, chapter "The Panar Man-Eater"

Hybrids

Crossbreeding between leopards and other members of the genus Panthera has been documented, resulting in hybrids. A cross between a lioness and a male leopard is known as a leopon (or a lipard if the sex of the parents is reversed). Leopons have been bred in captivity; a well-documented case occurred at the Koshien Hanshin Park in Nishinomiya, Japan, in the late 1950s.^[96] Although lions and leopards may come into contact in sub-Saharan Africa, they are not widely believed to interbreed naturally. However, there have been anecdotal reports of small lions with exceptionally pronounced spotting, known as "marozis" and various other names, in several African countries, for which there has been cryptozoological speculation that they may be naturally occurring lion-leopard hybrids.^[97]

Crossbreeding between jaguars and leopards in captivity has also been documented.^[98][99] A cross between a female leopard and a male jaguar is referred to as a jagupard, the reverse is known a leguar; however, crosses between either have also been called lepjags. Such crosses can only occur in captivity because leopards do not exist in the wild on the American continents where jaguars live. There have also been a few claims of crosses between tigers and leopards.^[100]

A pumapard is a hybrid animal resulting from a mating between a leopard and a puma (a member of the genus Puma, not the genus Panthera). Three sets of these hybrids were bred in the late 1890s and early 1900s by Carl Hagenbeck at his animal park in Hamburg, Germany. While most of these animals did not reach adulthood, one of these was purchased in 1898 by the Berlin Zoo. A similar hybrid in the Berlin Zoo purchased from Hagenbeck was a cross between a male leopard and a female puma. A specimen in the Hamburg Zoo (in the photo at right) was the reverse pairing, fathered by a puma bred to an Indian leopardess.^[101]

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.^[101]

In popular culture

• The 1938 film Bringing up Baby prominently features a pet leopard.
• Rudyard Kipling's novel The Jungle Book features a black leopard named Bagheera.
• In Disney's Tarzan, the character Sabor is a leopardess. She kills both Tarzan's biological parents and Kala's first child, before an adult Tarzan kills her in a fierce fight. She is voiced by Frank Welker, using sounds of big cats such as leopards, lions, tigers, jaguars, and cougars.
• The Island of Doctor Moreau by H.G. Wells features a human-leopard hybrid known as Leopard-Man, and the film The Island of Dr. Moreau (1996 film) features a human-leopard hybrid named Lo-Mai.
• The mascot of the 2010 World Cup in South Africa was an anthropomorphic leopard called Zakumi.

See also

• List of solitary animals

References

Bibliography

• Partridge, Eric (1983). Origins: A Short Etymological Dictionary of Modern English. New York: Greenwich House. ISBN 0-517-41425-2. 

Further reading

External links

Wikispecies has information related to: Panthera pardus

Wikimedia Commons has media related to: Panthera pardus (category)

• IUCN/SSC Cat Specialist Group : Panthera pardus in Africa and Panthera pardus in Asia
• Leopard Anthology: Research and conservation of leopards in Asia
• Asian Leopard Specialist Society: Research, conservation and management of Asian leopard subspecies
• The Animal Files: Leopard

Extant Carnivora species Kingdom: Animalia Phylum: Chordata Class: Mammalia Infraclass: Eutheria Superorder: Laurasiatheria   Suborder Feliformia Nandiniidae Nandinia African palm civet (N. binotata) Herpestidae (Mongooses) Atilax Marsh mongoose (A. paludinosus) Bdeogale Bushy-tailed mongoose (B. crassicauda) Jackson's mongoose (B. jacksoni) Black-footed mongoose (B. nigripes) Crossarchus Alexander's kusimanse (C. alexandri) Angolan kusimanse (C. ansorgei) Common kusimanse (C. obscurus) Flat-headed kusimanse (C. platycephalus) Cynictis Yellow mongoose (C. penicillata) Dologale Pousargues's mongoose (D. dybowskii) Galerella Angolan slender mongoose (G. flavescens) Black mongoose (G. nigrata) Somalian slender mongoose (G. ochracea) Cape gray mongoose (G. pulverulenta) Slender mongoose (G. sanguinea) Helogale Ethiopian dwarf mongoose (H. hirtula) Common dwarf mongoose (H. parvula) Herpestes Short-tailed mongoose (H. brachyurus) Indian gray mongoose (H. edwardsii) Indian brown mongoose (H. fuscus) Egyptian mongoose (H. ichneumon) Small Asian mongoose (H. javanicus) Long-nosed mongoose (H. naso) Collared mongoose (H. semitorquatus) Ruddy mongoose (H. smithii) Crab-eating mongoose (H. urva) Stripe-necked mongoose (H. vitticollis) Ichneumia White-tailed mongoose (I. albicauda) Liberiictus Liberian mongoose (L. kuhni) Mungos Gambian mongoose (M. gambianus) Banded mongoose (M. mungo) Paracynictis Selous' mongoose (P. selousi) Rhynchogale Meller's mongoose (R. melleri) Suricata Meerkat (S. suricatta) Hyaenidae (Hyenas) Crocuta Spotted hyena (C. crocuta) Hyaena Brown hyena (H. brunnea) Striped hyena (H. hyaena) Proteles Aardwolf (P. cristatus) Felidae Large family listed below Viverridae Large family listed below Eupleridae Small family listed below   Family Felidae Felinae Acinonyx Cheetah (A. jubatus) Caracal Caracal (C. caracal) African golden cat (C. aurata) Catopuma Bay cat (C. badia) Asian golden cat (C. temminckii) Felis Chinese mountain cat (F. bieti) Domestic cat (F. catus) Jungle cat (F. chaus) Sand cat (F. margarita) Black-footed cat (F. nigripes) Wildcat (F. silvestris) Leopardus Pantanal cat (L. braccatus) Colocolo (L. colocolo) Geoffroy's cat (L. geoffroyi) Kodkod (L. guigna) Southern tigrina (L. guttulus) Andean mountain cat (L. jacobita) Pampas cat (L. pajeros) Ocelot (L. pardalis) Oncilla (L. tigrinus) Margay (L. wiedii) Leptailurus Serval (L. serval) Lynx Canadian lynx (L. canadensis) Eurasian lynx (L. lynx) Iberian lynx (L. pardinus) Bobcat (L. rufus) Otocolobus Pallas's cat (O. manul) Pardofelis Marbled cat (P. marmorata) Prionailurus Leopard cat (P. bengalensis) Flat-headed cat (P. planiceps) Rusty-spotted cat (P. rubiginosus) Fishing cat (P. viverrinus) Puma Cougar (P. concolor) Jaguarundi (P. yagouaroundi) Pantherinae Panthera Lion (P. leo) Jaguar (P. onca) Leopard (P. pardus) Tiger (P. tigris) Snow leopard (P. uncia) Neofelis Clouded leopard (N. nebulosa) Sunda clouded leopard (N. diardi)   Family Viverridae (includes Civets) Paradoxurinae Arctictis Binturong (A. binturong) Arctogalidia Small-toothed palm civet (A. trivirgata) Macrogalidia Sulawesi palm civet (M. musschenbroekii) Paguma Masked palm civet (P. larvata) Paradoxurus Golden wet-zone palm civet (P. aureus) Asian palm civet (P. hermaphroditus) Jerdon's palm civet (P. jerdoni) Golden palm civet (P. zeylonensis) Hemigalinae Chrotogale Owston's palm civet (C. owstoni) Cynogale Otter civet (C. bennettii) Diplogale Hose's palm civet (D. hosei) Hemigalus Banded palm civet (H. derbyanus) Prionodontinae (Asiatic linsangs) Prionodon Banded linsang (P. linsang) Spotted linsang (P. pardicolor) Viverrinae Civettictis African civet (C. civetta) Genetta (Genets) Abyssinian genet (G. abyssinica) Angolan genet (G. angolensis) Bourlon's genet (G. bourloni) Crested servaline genet (G. cristata) Common genet (G. genetta) Johnston's genet (G. johnstoni) Rusty-spotted genet (G. maculata) Pardine genet (G. pardina) Aquatic genet (G. piscivora) King genet (G. poensis) Servaline genet (G. servalina) Haussa genet (G. thierryi) Cape genet (G. tigrina) Giant forest genet (G. victoriae) Poiana African linsang (P. richardsonii) Leighton's linsang (P. leightoni) Viverra Malabar large-spotted civet (V. civettina) Large-spotted civet (V. megaspila) Malayan civet (V. tangalunga) Large Indian civet (V. zibetha) Viverricula Small Indian civet (V. indica)   Family Eupleridae Euplerinae Cryptoprocta Fossa (C. ferox) Eupleres Falanouc (E. goudotii) Western falanouc (E. major) Fossa Malagasy civet (F. fossana) Galidiinae Galidia Ring-tailed mongoose (G. elegans) Galidictis Broad-striped Malagasy mongoose (G. fasciata) Grandidier's mongoose (G. grandidieri) Mungotictis Narrow-striped mongoose (M. decemlineata) Salanoia Brown-tailed mongoose (S. concolor) Durrell's vontsira (S. durrelli)   Suborder Caniformia (cont. below) Ursidae (Bears) Ailuropoda Giant panda (A. melanoleuca) Helarctos Sun bear (H. malayanus) Melursus Sloth bear (M. ursinus) Tremarctos Spectacled bear (T. ornatus) Ursus American black bear (U. americanus) Brown bear (U. arctos) Polar bear (U. maritimus) Asian black bear (U. thibetanus) Mephitidae (Skunks) Conepatus (Hog-nosed skunks) Molina's hog-nosed skunk (C. chinga) Humboldt's hog-nosed skunk (C. humboldtii) American hog-nosed skunk (C. leuconotus) Striped hog-nosed skunk (C. semistriatus) Mephitis Hooded skunk (M. macroura) Striped skunk (M. mephitis) Mydaus Sunda stink badger (M. javanensis) Palawan stink badger (M. marchei) Spilogale (Spotted skunks) Southern spotted skunk (S. angustifrons) Western spotted skunk (S. gracilis) Eastern spotted skunk (S. putorius) Pygmy spotted skunk (S. pygmaea) Procyonidae Bassaricyon (Olingos) Eastern lowland olingo (B. alleni) Northern olingo (B. gabbii) Western lowland olingo (B. medius) Olinguito (B. neblina) Bassariscus Ring-tailed cat (B. astutus) Cacomistle (B. sumichrasti) Nasua (Coatis inclusive) White-nosed coati (N. narica) South American coati (N. nasua) Nasuella (Coatis inclusive) Western mountain coati (N. olivacea) Eastern mountain coati (N. meridensis) Potos Kinkajou (P. flavus) Procyon Crab-eating raccoon (P. cancrivorus) Raccoon (P. lotor) Cozumel raccoon (P. pygmaeus) Ailuridae Ailurus Red panda (A. fulgens)   Suborder Caniformia (cont. above) Otariidae (Eared seals) (includes fur seals and sea lions) (Pinniped inclusive) Arctocephalus South American fur seal (A. australis) Australasian fur seal (A. forsteri) Galápagos fur seal (A. galapagoensis) Antarctic fur seal (A. gazella) Juan Fernández fur seal (A. philippii) Brown fur seal (A. pusillus) Guadalupe fur seal (A. townsendi) Subantarctic fur seal (A. tropicalis) Callorhinus Northern fur seal (C. ursinus) Eumetopias Steller sea lion (E. jubatus) Neophoca Australian sea lion (N. cinerea) Otaria South American sea lion (O. flavescens) Phocarctos New Zealand sea lion (P. hookeri) Zalophus California sea lion (Z. californianus) Galápagos sea lion (Z. wollebaeki) Odobenidae (Pinniped inclusive) Odobenus Walrus (O. rosmarus) Phocidae (Earless seals) (Pinniped inclusive) Cystophora Hooded seal (C. cristata) Erignathus Bearded seal (E. barbatus) Halichoerus Gray seal (H. grypus) Histriophoca Ribbon seal (H. fasciata) Hydrurga Leopard seal (H. leptonyx) Leptonychotes Weddell seal (L. weddellii) Lobodon Crabeater seal (L. carcinophagus) Mirounga (Elephant seals) Northern elephant seal (M. angustirostris) Southern elephant seal (M. leonina) Monachus Mediterranean monk seal (M. monachus) Hawaiian monk seal (M. schauinslandi) Ommatophoca Ross seal (O. rossi) Pagophilus Harp seal (P. groenlandicus) Phoca Spotted seal (P. largha) Harbor seal (P. vitulina) Pusa Caspian seal (P. caspica) Ringed seal (P. hispida) Baikal seal (P. sibirica) Canidae Large family listed below Mustelidae Large family listed below   Family Canidae (includes dogs) Atelocynus Short-eared dog (A. microtis) Canis Side-striped jackal (C. adustus) African golden wolf (C. anthus) Golden jackal (C. aureus) Coyote (C. latrans) Gray wolf (C. lupus) Black-backed jackal (C. mesomelas) Red wolf (C. rufus) Ethiopian wolf (C. simensis) Cerdocyon Crab-eating fox (C. thous) Chrysocyon Maned wolf (C. brachyurus) Cuon Dhole (C. alpinus) Lycalopex Culpeo (L. culpaeus) Darwin's fox (L. fulvipes) South American gray fox (L. griseus) Pampas fox (L. gymnocercus) Sechuran fox (L. sechurae) Hoary fox (L. vetulus) Lycaon African wild dog (L. pictus) Nyctereutes Raccoon dog (N. procyonoides) Otocyon Bat-eared fox (O. megalotis) Speothos Bush dog (S. venaticus) Urocyon Gray fox (U. cinereoargenteus) Island fox (U. littoralis) Vulpes (Foxes) Bengal fox (V. bengalensis) Blanford's fox (V. cana) Cape fox (V. chama) Corsac fox (V. corsac) Tibetan sand fox (V. ferrilata) Arctic fox (V. lagopus) Kit fox (V. macrotis) Pale fox (V. pallida) Rüppell's fox (V. rueppelli) Swift fox (V. velox) Red fox (V. vulpes) Fennec fox (V. zerda)   Family Mustelidae Lutrinae (Otters) Aonyx African clawless otter (A. capensis) Oriental small-clawed otter (A. cinerea) Enhydra Sea otter (E. lutris) Hydrictis Spotted-necked otter (H. maculicollis) Lontra North American river otter (L. canadensis) Marine otter (L. felina) Neotropical otter (L. longicaudis) Southern river otter (L. provocax) Lutra European otter (L. lutra) Hairy-nosed otter (L. sumatrana) Lutrogale Smooth-coated otter (L. perspicillata) Pteronura Giant otter (P. brasiliensis) Mustelinae (including badgers) Arctonyx Hog badger (A. collaris) Eira Tayra (E. barbara) Galictis Lesser grison (G. cuja) Greater grison (G. vittata) Gulo Wolverine (G. gulo) Ictonyx Saharan striped polecat (I. libyca) Striped polecat (I. striatus) Lyncodon Patagonian weasel (L. patagonicus) Martes (Martens) American marten (M. americana) Yellow-throated marten (M. flavigula) Beech marten (M. foina) Nilgiri marten (M. gwatkinsii) European pine marten (M. martes) Japanese marten (M. melampus) Fisher (M. pennanti) Sable (M. zibellina) Meles Japanese badger (M. anakuma) Asian badger (M. leucurus) European badger (M. meles) Mellivora Honey badger (M. capensis) Melogale (Ferret-badgers) Bornean ferret-badger (M. everetti) Chinese ferret-badger (M. moschata) Javan ferret-badger (M. orientalis) Burmese ferret-badger (M. personata) Mustela (Weasels and Ferrets) Amazon weasel (M. africana) Mountain weasel (M. altaica) Stoat (M. erminea) Steppe polecat (M. eversmannii) Colombian weasel (M. felipei) Long-tailed weasel (M. frenata) Japanese weasel (M. itatsi) Yellow-bellied weasel (M. kathiah) European mink (M. lutreola) Indonesian mountain weasel (M. lutreolina) Black-footed ferret (M. nigripes) Least weasel (M. nivalis) Malayan weasel (M. nudipes) European polecat (M. putorius) Siberian weasel (M. sibirica) Back-striped weasel (M. strigidorsa) Egyptian weasel (M. subpalmata) Neovison (Minks) American mink (N. vison) Poecilogale African striped weasel (P. albinucha) Taxidea American badger (T. taxus) Vormela Marbled polecat (V. peregusna)