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Leopard
Temporal range: Late Pliocene or Early Pleistocene to Recent
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Mammalia
Order:
Carnivora
Family:
Felidae
Genus:
Panthera
Species:
P. pardus
Binomial name
Panthera pardus
Linnaeus, 1758

The leopard (Panthera pardus) is an Old World mammal of the Felidae family and the smallest of the four big cats of the genus Panthera, the other three being the tiger, lion and jaguar. Once distributed across southern Eurasia and Africa, from Korea to South Africa and Spain, the leopard's range of distribution has decreased radically and the leopard now chiefly occurs in sub-Saharan Africa. There are fragmented populations in Israel, Indochina, Malaysia, and western 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 sixty kilometres (37 miles) an hour. The leopard consumes virtually any animal it can catch and ranges from rainforest to desert. Its ecological role 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 compound word derived from λέων léon ("lion") and πάρδος párdos ("male panther"), the latter related to Sanskrit पृदाकु pṝdāku ("snake, tiger, panther").[2][3]

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.[4]

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.[2][5][6]

Taxonomy

Like the rest of the feline family, the Panthera genus has been subject to much alteration and debate and 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.[7] The fossil record points to the emergence of Panthera just two to 3.8 million years ago.[7][8]

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.[9] 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,[10] 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.[7] 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 seen as diverging 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.[7] 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.[11]

Subspecies

Indian leopard

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.[12] 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 mght be an underestimate because of limited sampling of African leopards. Their list as follows:[11]

File:Leo4.JPG
Sri Lankan leopard
Female leopard in the Sabi Sands area of South Africa. Note the white spot on its tail, used for communicating with cubs while hunting or in long grass

Older taxonomic divisions

Included in the African leopard (P. pardus pardus):[12]

  • 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):[12]

  • 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):[12]

  • 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 Panthera, the leopard is still able to take large prey given a massive skull that well utilizes powerful jaw muscles.[13] Its body is comparatively long for a cat and its legs are short.[14] 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 90 kg (81 to 198 lbs) compared to 28 to 60 kg (62 to 132 lbs) for females.[15]

One of many spotted cats, a leopard may be mistaken for a cheetah or a jaguar. 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.[14]

Leopards have been reported to reach 21 years of age in captivity.[16]

Black leopards

Main article: Black panther

A melanistic morph of the leopard occurs particularly in mountainous areas and rain forests. The black color is heritable and caused by a recessive gene locus.[17] In some regions, for example on the Malayan Peninsula, up to half of all leopards are black. This may be a beneficial mutation that helps them survive in their rainforest habitat. In Africa, black leopards seem to be most common in the Ethiopian Highlands. While they are commonly called black panthers, the term is not applied exclusively to leopards, as it also applies to melanistic jaguars. Black leopards are less successful on the African plains because their coloration makes them stand out. While known as panthers, there are no known cases of melanistic cougars.[18]

Biology and behavior

Female leopard. Note the ocelli (white spots on the back of the ears) used.

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.[19] 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.[20] 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 tress, underneath rocks or in the grass.[20]

Diet and hunting

Leopards are opportunistic hunters. Although mid-sized animals are preferred, the leopard will eat anything from dung beetles to 900 kg (1,984 lb) male giant elands.[13] Their diet consists mostly of ungulates and monkeys, but rodents, reptiles, amphibians, birds and fish are also eaten;[21] In Africa, mid-sized antelopes provide a majority of the leopard's prey, especially impala and Thomson's gazelles.[22] In Asia the leopard preys on deer such as chitals and muntjacs as well as various Asian antelopes and Ibex.

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,[22] 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 25 kg (55 lb) 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.[23]

Anatolian leopard resting on a tree

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.[24] Cubs are usually born in a litter of 2–4,[25] 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.[26] 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.[13] 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.[27] 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.[28] 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.[26] 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.[29]

The leopard is solitary and, aside from mating, interactions between individuals appear to be infrequent.[29] 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.[30]

Ecology

Distribution and habitat

Data from 1996 found that the leopard has the largest distribution of any wild cat,[13] occurring most in certain parts of southern Asia and widely in eastern and central Africa,[31] 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; and in the Indian subcontinent, Southeast Asia, and China, the leopard is still relatively abundant.[1]

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.[32] 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° celcius.[11]

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.[33]

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.[13] 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.[34]

Variant coloration

A melanistic leopard, or "black panther".

A pseudo-melanistic leopard has a normal background colour, 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.

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:

There is, however, a peculiar dark phase in South Africa, a specimen of which was obtained in 1885 in hilly land covered with scrub-jungle, near Grahamstown. The ground-colour of this animal was a rich tawny, with an orange tinge; but the spots, instead of being of the usual rosette-like form, were nearly all small and solid, like those on the head of an ordinary leopard; while from the top of the head to near the root of the tail the spots became almost confluent, producing the appearance of a broad streak of black running down the back. A second skin had the black area embracing nearly the whole of the back and flanks, without showing any trace of the spots, while in those portions of the skin where the latter remained they were of the same form as in the first specimen. Two other specimens are known; the whole four having been obtained from the Albany district. These dark-coloured South African leopards differ from the black leopards of the northern and eastern parts of Africa and Asia in that while in the latter the rosette-like spots are always retained and clearly visible, in the former the rosettes are lost – as, indeed, is to a considerable extent often the case in ordinary African leopards – and all trace of spots disappears from the blacker portions of the skin.

— Lydekker, R. (1910), Harmsworth Natural History

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.

The wide black portion, which glistens like the sheen of silk velvet, extends from the top of the head to the extremity of the tail entirely free from any white or tawny hairs … In the tiger, the stripes are black, of a uniform character, upon a tawny background, and they run in parallel lines from the centre of the back to the belly. In this skin, the stripes are almost golden yellow, without the uniformity and parallelism of the tiger characteristics, and they extend along the sides in labyrinthine graceful curls and circles, several inches below the wide shimmering black continuous course of the back. The extreme edges around the legs and belly are white and spotted like the skin of a leopard … The skin is larger than that of a leopard but smaller than that of a full grown tiger.

— Collins, Holdridge Ozro (1915)

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

Main article: Pumapard
Pumapard, Rothschild Museum, Tring.

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

File:Dionysus on a leopard, pella, greece.jpg
A mosaic depicting Dionysus on a leopard. 4th century BC. Pella, Greece.

Leopards have been known to humans since antiquity and have featured in the art, mythology and folklore of many countries where they have occurred historically, such as ancient Greece, Persia and Rome, as well as some where they haven't such as England. The modern use of the leopard as an emblem for sport or 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. Leopeard 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 animals were given to Henry III by the Holy Roman Emperor Frederick II.[35]

A female leopard in the Sabi Sands of South Africa illustrating just how close tourists can get to these wild cats.

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.

Coat of arms of the German state of Baden-Württemberg

Heraldry

Main article: Leopard (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.

The Leopard Men

The Leopard men were a West African society that practiced cannibalism during the 1900s. They were centred in Sierra Leone, Liberia and Côte d'Ivoire.

Members would dress in leopard skins, waylaying travelers with sharp claw-like weapons in the form of leopards' claws and teeth. The victims' flesh would be cut from their bodies and distributed to members of the society. They held a belief that this ritual cannibalism would strengthen both members of the society as well as their entire tribe.[36]

Man-eating

Although most leopards will tend to avoid humans, 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.[citation needed] 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.[citation needed] Kenneth Anderson, who had first hand experience with many man-eating leopards, 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...

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

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.[citation needed]

Notes

  1. ^ a b Template:IUCN2006 Database entry includes justification for why this species is of least concern
  2. ^ a b ""Leopard"". Online Etymology Dictionary. Douglas Harper. Retrieved 2007-11-30.
  3. ^ Monier-Williams, Sir Monier. "pṝdāku". A Sanskrit-English Dictionary (JPEG). pp. p. 647. Retrieved 2007-11-30. {{cite book}}: |pages= has extra text (help); External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  4. ^ Template:La icon Linnaeus, C (1758). Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. (Editio decima, reformata. ed.). Holmiae. (Laurentii Salvii).
  5. ^ Monier-Williams, Sir Monier. "pāṇḍara". A Sanskrit-English Dictionary (JPEG). pp. p. 616. Retrieved 2007-11-30. {{cite book}}: |pages= has extra text (help); External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  6. ^ Monier-Williams, Sir Monier. "puṇḍárīka". A Sanskrit-English Dictionary (JPEG). pp. p. 631. Retrieved 2007-11-30. {{cite book}}: |pages= has extra text (help); External link in |chapterurl= (help); Unknown parameter |chapterurl= ignored (|chapter-url= suggested) (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
  7. ^ a b c d Johnson, W.E., Eizirik, E., Pecon-Slattery, J., Murphy, W.J., Antunes, A., Teeling, E. & O'Brien, S.J. (2006). "The Late Miocene radiation of modern Felidae: A genetic assessment". Science. 311: 73–77. doi:10.1126/science.1122277.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  8. ^ Turner, A. (1987). "New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora)". Annals of the Transvaal Museum. 34: 319–347. ISSN 0041-1752. {{cite journal}}: Cite has empty unknown parameters: |month= and |coauthors= (help)
  9. ^ Yu L & Zhang YP (2005). "Phylogenetic studies of pantherine cats (Felidae) based on multiple genes, with novel application of nuclear beta-fibrinogen intron 7 to carnivores" (PDF). Molecular Phylogenetics and Evolution. 35 (2): 483–495. doi:10.1016/j.ympev.2005.01.017.
  10. ^ Wozencraft, W. C. (2005). "Order Carnivora". In Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 548. ISBN 978-0-8018-8221-0. OCLC 62265494.
  11. ^ a b c Uphyrkina, O. (2001). "Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus". Molecular Ecology. 10 (11): 2617–2633. Retrieved 2008-08-06. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  12. ^ a b c d Miththapala, Sriyanie (1996). "Phylogeographic Subspecies Recognition in Leopards (P. pardus): Molecular Genetic Variation". Conservation Biology. 10 (4): 1115–1132. Retrieved 2008-06-06. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  13. ^ a b c d e Nowell, K.; Jackson, P. eds. (1996). Wild Cats. Status Survey and Conservation Action Plan. IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland. (see Panthera Pardus, pp. 24 – 29.)
  14. ^ a b "Leopard". African Wildlife Foundation. Retrieved 2007-09-21..
  15. ^ Ronald M. Nowak: Walker's Mammals of the World. Johns Hopkins University Press, 1999 ISBN 0-8018-5789-9
  16. ^ Crandall, L (1964). The management of wild animals in captivity. Chicago: University of Chicago Press.
  17. ^ "The Black Panther". Retrieved 3 April. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
  18. ^ "Black leopard general information". Retrieved 3 April. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
  19. ^ "Leopard biology and behavior". Retrieved 6 June. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
  20. ^ a b "Animal bytes - Panthera pardus". Retrieved 6 June. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
  21. ^ Schaller, p. 290
  22. ^ a b Schaller, p. 291
  23. ^ Hayward, M. W. (2006). "Prey preferences of the leopard (Panthera pardus)". Journal of Zoology. 270 (2): 298–313. Retrieved 2007-09-13. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  24. ^ Sadleir R (1966). "Notes on the Reproduction of the larger Felidae". Int. Zoo Yearbook: Vol 6. London: Zool. Soc. London. pp. 184–87.
  25. ^ "Leopard". wildcatconservation.org. 2008. Retrieved 6 June. {{cite web}}: Check date values in: |accessdate= (help)
  26. ^ a b Marker, L. L. (2005). "Factors affecting leopard (Panthera pardus) spatial ecology, with particular reference to Namibian farmlands" (PDF). South African Journal of Wildlife Research. 35 (2). Retrieved 2007-09-13. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  27. ^ Mizutani, F. (1998). "Home-range and movements of leopards (Panthera pardus) on a livestock ranch in Kenya". Journal of Zoology. 244: 269–286. doi:10.1017/S0952836998002118. Retrieved 2007-09-13. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  28. ^ Odden, Morten (2005). "Spacing and activity patterns of leopards Panthera pardus in the Royal Bardia National Park, Nepal". Wildlife Biology. 11: 145–152. doi:10.2981/0909-6396(2005)11[145:SAAPOL]2.0.CO;2. Retrieved 2007-09-13. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  29. ^ a b Jenny, D. (1996). "Spatial organization of leopards Panthera pardus in Tai National Park, Ivory Coast: Is rainforest habitat a "tropical haven"?". Journal of Zoology. 240 (3): 427–440. Retrieved 2007-09-13. {{cite journal}}: Unknown parameter |month= ignored (help)
  30. ^ Hunter, Luke (2003). "The landscape ecology of leopards (Panthera pardus) in northern KwaZulu-Natal, South Africa: A preliminary project report" (PDF). Ecological Journal. 5: 24–30. Retrieved 2007-09-16. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  31. ^ LeeAnn Bies. "Leopard distribution". Retrieved 5 June. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)
  32. ^ Jenny, David (2005). "Hunting behaviour in West African forest leopards". African Journal of Ecology. 43 (3): 197–200. Retrieved 2008-05-23. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
  33. ^ Schaller, p. 293
  34. ^ Karanth, K. Ullas (2000). "Behavioural correlates of predation by tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India". Journal of Zoology. 250: 255–265. Retrieved 2008-06-05. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  35. ^ Owen, James (November 3, 2005). "Medieval Lion Skulls Reveal Secrets of Tower of London "Zoo"". National Geographic Magazine. National Geographic. Retrieved 2007-09-05.
  36. ^ "The Leopard Society - Africa in the mid 1900s". Retrieved 3 April. {{cite web}}: Check date values in: |accessdate= (help); Unknown parameter |accessyear= ignored (|access-date= suggested) (help)

References

  • Allsen, Thomas T. (2006). "Natural History and Cultural History: The Circulation of Hunting Leopards in Eurasia, Seventh-Seventeenth Centuries." In: Contact and Exchange in the Ancient World. Ed. Victor H. Mair. University of Hawai'i Press. Pp. 116-135. ISBN-13: ISBN 978-0-8248-2884-4; ISBN-10: ISBN 0-8248-2884-4
  • Khalaf-von Jaffa, Norman Ali Bassam Ali Taher (2005). The Arabian Leopard (Panthera pardus nimr). Gazelle: The Palestinian Biological Bulletin. Number 42, June 2005. pp. 1-8. (in German).
  • Khalaf-Sakerfalke von Jaffa, Norman Ali Bassam Ali Taher (2006). The Chinese leopard (Panthera pardus japonensis, Gray 1862) in Neunkirchen Zoo, Neunkirchen, Saarland, Germany. Gazelle: The Palestinian Biological Bulletin. Number 60, December 2006. pp. 1-10.
  • Schaller, George B. (1972). The Serengeti Lion. Chicago: University of Chicago Press. ISBN 0-226-73639-3.
  • Leopards and spots on ears and tail [1]
  • DeRuiter, D.J. and Berger, L.R. (2000) Leopards as Taphonomic Agents in dolomitic Caves - Implications for bone Accumulations in the Hominid-bearing Deposits of South Africa. J. Arch. Sci. 27, 665-684.
  • Taylor, Peter (2005). Beyond Conservation: A Wildland Strategy. Earthscan. ISBN 1844071979. {{cite book}}: Unknown parameter |coathors= ignored (help)
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