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For other uses, see Leopard (disambiguation), Leopards (disambiguation), and Leopardi (disambiguation).

Leopard
Temporal range: Late Pliocene or Early Pleistocene to recent
An African leopard (Panthera pardus pardus)
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Mammalia
Order:
Carnivora
Family:
Felidae
Subfamily:
Pantherinae
Genus:
Panthera
Species:
P. pardus
Binomial name
Panthera pardus
(Linnaeus, 1758)
Subspecies

see text

Current range of the leopard, former (red), uncertain (yellow), highly fragmented (light green), and present (dark green)
Synonyms

Felis pardus Linnaeus, 1758

The leopard (Panthera pardus) (/ˈlɛpərd/) 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

The common name "leopard" (pronounced /\ˈle-pərd\/)[6] is a Greek compound of λέων leōn ("lion") and πάρδος pardos ("male panther"). The Greek word is related to Sanskrit पृदाकु pṛdāku ("snake", "tiger" or "panther"), and probably derives from a Mediterranean language, such as Egyptian.[7][8] The name was first used in the 13th century.[6] Other vernacular names for the leopard include graupanther, panther and several regional names such as tendwa in India.[9] The term "black panther" refers to leopards with melanistic genes.[10]

The scientific name of the leopard is Panthera pardus. The generic name Panthera derives from Latin via Greek πάνθηρ (pánthēr).[11] The term "panther", whose first recorded use dates back to the 13th century AD, generally refers to the leopard, and less often to the cougar and the jaguar.[10] Alternative origins suggested for Panthera include an Indo-Iranian word meaning "white-yellow" or "pale". In Sanskrit, this could have been derived from पाण्डर pāṇḍara ("tiger"), which in turn comes from पुण्डरीक puṇḍárīka (with the same meaning).[8][11] The specific name pardus is derived from the Greek πάρδος (pardos) ("male panther").[9]

Taxonomy

Two cladograms proposed for Panthera. The upper cladogram is based on the 2006 and 2009 studies, while the other is based on the 2010 and 2011 studies.

The leopard is one of the five extant species of the genus Panthera, which also includes the jaguar (P. onca), the lion (P. leo), the snow leopard (P. uncia; sometimes placed in Uncia, a separate genus of its own) and the tiger (P. tigris). This genus, along with the genus Neofelis - which consists of the clouded leopard (N. nebulosa) and the Sunda clouded leopard (N. dardi) - forms the subfamily Pantherinae of the Felidae.[12][13] The leopard was first described by Swedish zoologist Carl Linnaeus in the 10th edition of Systema Naturae (1758). Linnaeus named the leopard as Felis pardus, placing it in the genus Felis along with the domestic cat, the jaguar, the Eurasian lynx, the lion, the ocelot and the tiger.[14] 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 F. pardus as a type species. Oken's classification, however, was not widely accepted, and until the early 20th century continued using Felis or Leopardus when describing leopard subspecies.[15] In 1916, British zoologist Reginald Innes Pocock accorded Panthera generic rank defining Panthera pardus as species.[16]

The leopard is part of the Panthera lineage, one of the eight lineages of Felidae. This lineage comprises the species of Panthera and Neofelis. The clouded leopard diverged first from the lineage, followed by the snow leopard. Subsequent branching began two to three million years ago, but the details of this are disputed. [17] A 2006 phylogenetic study by Warren E. Johnson (of the National Cancer Institute) and colleagues, based on nDNA and mtDNA analysis, showed that the leopard is sister to two clades within Panthera - one consisting of the tiger and the snow leopard, and the other of the lion and the jaguar.[18] This was seconded by a 2009 study by Lars Werdelin and colleagues.[19] However, the results obtained in a 2010 study by Brian W. Davis (of the Texas A&M University) and colleagues and a 2011 study by Ji H. Mazák (of the Shanghai Science and Technology Museum) and colleagues showed a swapping between the leopard and the jaguar in the cladogram.[20][21] Results of a 2001 phylogenetic analysis of chemical secretions amongst cats suggested, however, that the leopard is closely related to the lion.[22]

Subspecies

As many as 27 leopard subspecies were subsequently described by naturalists from 1794 to 1956. Since 1996, only eight subspecies have been considered valid on the basis of mitochondrial analysis.[23] Later analysis revealed a ninth valid subspecies, the Arabian leopard. Because of limited sampling of African leopards, this number might be an underestimation.[24]

The nine subspecies recognised by IUCN are:[1][24][25]

Subspecies of leopard
Subspecies Description Image
African leopard (P. p. pardus) Lives in Sub-Saharan Africa. It is the most widespread subspecies of leopards.
Indian leopard (P. p. fusca) Native to the Indian Subcontinent. It is widespread in India, Nepal, Bangladesh and Bhutan.
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.
Persian leopard (P. p. saxicolor), also known as Central Asian leopard or Caucasian leopard Inhabits the Caucasus, Turkmenistan, Afghanistan, and northern Iran.[25] It is the largest leopard subspecies.
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.
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.
Indochinese leopard (P. p. delacouri), also known as South-Chinese leopard Widespread in mainland Southeast Asia and South China.
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.
Sri Lankan leopard (P. p. kotiya) Found only in Sri Lanka.
Map of approximate distribution of leopard subspecies

A morphological analysis of characters of leopard skulls implies the validity of two more subspecies:[25]

The following African leopard populations used to be considered subspecies until 1996:[24][23]

The smallest leopard subspecies is the Arabian leopard. Adult females weigh as little as 18 kg (40 lb).[26] 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.[27]

Evolution

The last common ancestor of the Panthera and Neofelis species is believed to have occurred about 6.37 million years ago. The clouded leopard was the first to diverge from the rest of the Panthera lineage, followed by the snow leopard. The genus Panthera is believed to have emerged in Asia, from where they subsequently emigrated to Africa. The tiger-snow leopard clade diverged from the rest of Panthera around 2.9 million years ago.[20][21] Johnson and colleagues suggest that the leopard diverged next, and followed by the lion-jaguar clade.[18]

Fossils of ancestors of the leopard have been found in East Africa and South Asia, dating back to the Pleistocene between 2 and 3.5 million years ago. The modern leopard is suggested to have evolved in Africa 0.5 to 0.8 million years ago and to have radiated across Asia 0.2 to 0.3 million years ago.[24]

In Europe, the leopard is known at least since the Pleistocene. Fossil bones and teeth dating from the Pliocene were found in Perrier in France, northeast of London, and in Valdarno (Italy). Similar fossils dating back to the Pleistocene were excavated mostly in loess and caves at 40 sites in the continent - 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.[28] The Pleistocene leopards of Europe can be divided into four subsequent subspecies. The first European leopard subspecies P. p. begoueni is known from the beginning of the early Pleistocene and was replaced about 0.6 million years ago by P. p. sickenbergi, which in turn was replaced by P. p. antiqua around 0.3 million years ago. The most recent 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 several parts of Europe.[29]

Genetics

The diploid number of chromosomes in the leopard is 38, the same as in any other felid, save for the ocelot and the margay, whose diploid number of chromosomes is 36.[30] The chromosomes include four acrocentric, five metacentric, seven submetacentric and two telocentric pairs.[31]

Hybrids

Pumapard, 1904
Main article: Panthera hybrid

Crossbreeding between the leopard and the other members of the Panthera has been documented. In 1953, a lioness and a male leopard were mated in the Hanshin Park in Nishinomiya, Japan. The first litter from this pairing was born on 2 November 1959, consisting of a male and a female. Another litter was born in 1961, in which all the offspring were spotted and bigger than juvenile leopard. The hybrid came to be known as "leopon". Unsuccessful attempts were made to mate a leopon with a tigress.[32]

Although lions and leopards may come into contact in sub-Saharan Africa, they are generally not known to interbreed naturally. However, there have been anecdotal reports of felids larger than the cheetah but smaller than the lion, with a lion-like face, from the Central African Republic, Kenya, Rwanda and Uganda. This animal, known as the marozi and by several other names, is covered with grayish spots or rosettes on the back, the flanks and the legs. However, there have been no confirmed sightings of the marozi since the 1930s.[33]

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. The pumapard is characterised by a long body like the puma's, but with shorter legs. The hybrid is in general dwarf, smaller than either parent. The coat is variously described as sandy, tawny or greyish with brown, chestnut or faded rosettes.[34]

Variant colouration

Melanistic leopard or "black panther"
Main article: Black panther

Melanistic leopards, known, like the melanistic jaguars, as "black panthers". Pseudomelanism (abundism) also occurs in leopards.[35] Melanism in leopards is inherited as a trait relatively recessive to the spotted form.[36] Interbreeding in melanistic leopard produces a significantly smaller litter size than produced by normal pairings.[37]

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.[38] 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 the 445 photographs of melanistic leopards, 410 came from study sites south of the Kra Isthmus, where the non-melanistic morph was never photographed. This data suggests the near fixation of the dark allele in the region. The expected time for the fixation of this recessive allele due to genetic drift alone ranged from about 1,100 years to about 100,000 years.[39][40]

A rare "strawberry" leopard was photographed in South Africa's Madikwe Game Reserve. This condition was probably caused by erythrism, a little-understood genetic condition that causes either an overproduction of red pigments or an underproduction of dark pigments.[41] Pseudomelanism has also been reported in leopard.[42]

Characteristics

African leopard at Serengeti National Park, Tanzania

The leopard is one of the big cats. Males stand 60–70 cm (24–28 in) at the shoulder; females are 57–64 cm (22–25 in) tall. The head-and-body length is typically between 90–190 cm (35–75 in). While males weigh 37–90 kg (82–198 lb), females weigh 28–60 kg (62–132 lb). The tail is nearly 58–110 cm (23–43 in) long.[43][44] 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.[45][46] 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.[citation needed]

The leopard's body is comparatively long, and its legs are short.[47] 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.[46][48]

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.[49]

Distribution and habitat

Leopards on the Magerius Mosaic from modern Tunisia. Numerous Roman mosaics from North African sites depict fauna now found only in tropical Africa.[50]
A leopard and her cub on the tree in the Serengeti savanna.

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][51]

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).[24] 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.[51] In India, leopard populations sometimes live quite close to human settlements and even in semi-developed areas.[52] 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.[51][52] Due to the leopard's superlative stealthiness, people often remain unaware that big cats live in nearby areas.[52]

Ecology and behaviour

Leopard resting on a tree

Leopards are elusive, solitary, and largely nocturnal.[53][54] 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.[55]

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.[56] 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.[57] They produce a number of vocalizations, including grunts, roars, growls, meows, and purrs.[43]

Social structure and home range

Leopard visual communication
Female showing white spots on the back of the ears (ocelli) used to communicate with other leopards.[58]
Female leopard showing the white spot on the tail used for communicating with cubs while hunting or in long grass.[58]

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.[59]

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.[60]

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.[61]

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.[62]

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.[53]

Hunting and diet

Stages of leopard hunting prey
Stalking
Killing young bushbuck
Dragging kill
Caching kill in a tree

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.[63] although leopards generally do not prey on such large animals.[64] 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.[65] 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.[66][67] 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.[68]

In the open savanna of Tsavo National Park, they kill most of their prey while hunting between sunset and sunrise.[69] 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.[70]

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.[71] 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.[72]

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).[73]

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.[68] 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.[69]

Interspecific predatory relationships

Video of a leopard in the wild

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.[74] The 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.[75] 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%).[75] In the Primore region of the Russian Far East, Amur leopards were absent or very rarely encountered at places where Siberian tigers reside.[76] 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.[77] 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.[77] There were also differences in the microhabitat preferences of the individual tiger and leopard followed over 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 [78] Usually when a tiger began to kill baits at sites formerly frequented by leopards, the leopards would no longer come and kill there.[79] 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.[80] 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.[81] 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.[79]

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.[82] 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.[83][84][85] Mugger crocodiles may also on rare occasions kill an adult leopard in India.[86] Lions are occasionally successful in climbing trees and fetching leopard kills,[68] 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.[87][88] Burmese pythons have been known to prey on leopards, with an adult cat having been recovered from the stomach of a 5.5 m (18 ft) specimen.[89]

Two cases of leopards killing cheetahs have been reported in 2014.[90][91]

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.[92] Gestation lasts for 90 to 105 days.[93] Cubs are usually born in a litter of 2–4 cubs.[94] Mortality of cubs is estimated at 41–50% during the first year.[70]

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.[63] 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.[56]

The average typical life span of a leopard is between 12 and 17 years.[95] 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.[96] 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.[97] 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.[98]

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 commonly represented on engravings and was used to symbolise the power of the king or oba; since the leopard was considered the king of the forest. Leopard were also kept and paraded as mascots and sacrificed to deities.[99]

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.[100]

Heraldry

Main article: Leopard (heraldry)
Three leopards (loggerheads) on the flag of Shropshire, England.

The lion passant guardant or leopard is a frequently used charge in heraldry, most commonly appearing in groups of three.[101] 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.[102]

Tourism

A female leopard in the Sabi Sands of South Africa near a game vehicle
Road sign in Namibia

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.[103] 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.[104]

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.[105]

Man-eating

Main article: Leopard attack
The Panar Leopard, shot by Jim Corbett in 1910 after allegedly killing 400 people

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.[106] 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.[107] 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"

See also

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Bibliography

Further reading

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