Leopard

Leopard[1]
Fossil range: Late Pliocene or Early Pleistocene to Recent
Conservation status

Near Threatened (IUCN 3.1)[2]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Felidae
Genus: Panthera
Species: P. pardus
Binomial name
Panthera pardus
Linnaeus, 1758
Historic (red) and present (green) range of the leopard

The leopard (pronounced /ˈlɛpərd/), Panthera pardus, is a member of the Felidae family and the smallest of the four "big cats" in the genus Panthera, the other three being the tiger, lion and jaguar. Once distributed across eastern and southern Asia and Africa, from Siberia to South Africa, the leopard's range of distribution has decreased radically because of hunting and loss of habitat. It is now chiefly found in sub-Saharan Africa; there are also fragmented populations in Pakistan, India, Sri Lanka, Indochina, Malaysia, and China. Because of its declining range and population, it is listed as a "Near Threatened" species by the IUCN.[2]

Compared to other members of the Felidae family, 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 slightly 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 jaguars do. Both leopards and jaguars that are melanistic (completely black or very dark) are known as black panthers.

The species' success in the wild is in part due to its opportunistic hunting behavior, its adaptability to habitats, its ability to run at speeds approaching 58 kilometres per hour (36 mph), its unequaled ability to climb trees even when carrying a heavy carcass,[3] and its notorious ability for stealth. The leopard consumes virtually any animal it can hunt down and catch. Its habitat ranges from rainforest to desert terrains.

Contents

Etymology

In antiquity, it was believed that a leopard was 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.[4][5]

A panther can be any of several species of large felid: the term can refers to cougars and jaguars in the American continents; and everywhere else, to leopards.[6]

The generic component of its modern scientific designation, Panthera pardus, is derived from Latin via Greek πάνθηρ pánthēr.[7] Folk etymology held that it was a compound of παν pan ("all") and θηρ ("beast").[8] However, it is believed instead to derive from an Indo-Iranian word meaning "white-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.[5][7]

Taxonomy and evolution

Indian leopard (Panthera pardus fusca)
Northern Chinese leopard (Panthera pardus japonensis)
African leopard (Panthera pardus pardus)
Amur leopard (Panthera pardus orientalis)

Like all of the feline family, the Panthera genus has been subject to much alteration and debate, and the exact relations between the four species (as well as the clouded leopard and snow leopard) have not been effectively resolved. It is believed that the basal divergence amongst the Felidae family occurred about 11 million years ago.[9] 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.[9]

Canonical works, such as the third edition of Mammal Species of the World, list the snow leopard as the only species within its genus, Uncia uncia,[1] but more recent research could change this placement. In a mitochondrial DNA study, Yu and Zhang (2005) suggest that the leopard is most closely related to the snow leopard and place the latter as a fifth species of Panthera, P. uncia.[10] Johnson et al. (2006) also 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.[9] Older research has tended to suggest that the leopard is most closely related to the lion and/or the jaguar. As recently as 2001, it was held to have split along with the lion in a phylogenetic analysis of chemical secretions amongst cats.[11] The leopard is also considered the type species of the genus Panthera.

Panthera is believed to have emerged in Asia, with ancestors of the leopard and other cats subsequently migrating into Africa.[9] 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.[12]

Subspecies

As many as 27 leopard subspecies were once suggested, the number growing from the time of Linnaeus in the 18th century to that of Reginald Pocock in the early 20th. In 1996, Miththapala et al. revised this downward to just eight subspecies based on DNA analysis.[13] Uphyrina et al. concurred with Miththapala et al. in 2001, but split out a ninth, the Arabian leopard (P. p. nimr). The latter researchers note the number might be an underestimation because of limited sampling of African leopards.[12] A recent study on the Caucasus subspecies shows solid support for two additional subspecies, and corrected the name of a third species based on the principle of priority.[14]

The subspecies recognised by Uphyrina et al. and Khorozyan et al. are:[12][14]

Physical characteristics

Rear view of female leopard. Note ocelli (white spots on the back of the ears), used to communicate with other leopards[3]

The leopard is an agile and stealthy predator. Although smaller than the other members of the Panthera genus, the leopard is still able to take large prey given its massive skull that well utilizes powerful jaw muscles.[15] Its body is comparatively long for a cat and its legs are short.[16] Head and body length is between 125 and 165 cm (49 and 65 in) and the tail reaches 60 to 110 cm (24 to 43 in). Shoulder height is 45 to 80 cm (18 to 31 in). The muscles attached to the scapula are exceptionally strong, which enhances the leopard's ability to climb trees.[15]

Leopards show a great diversity in size. Males are about 30% larger than females,[17] weighing 30 to 91 kg (66 to 200 lb) compared to 23 to 60 kg (51 to 130 lb) for females. Large males of up to 91 kg have been documented in Kruger National park in South Africa; however, males in the South Africa's coastal mountains average a much smaller 31 kg.[15] This wide variation in size is thought to result from the quality and availability of prey found in each habitat. Smaller sized leopards also are known in the deserts of the Middle East.[18]

Leopards may sometimes be confused with two other large spotted cats, the cheetah and the jaguar. However, the patterns of spots in each are different: the cheetah has simple 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 leopard is larger and much more muscular than the cheetah, but slightly smaller and more lightly built than the jaguar.[19]

Leopards show a great diversity in physical appearance, particularly because of the wide variations in color coat and rosette patterns. The leopard's rosettes are circular in East Africa but tend to be squarer in southern Africa[16] and larger in Asian populations.[15] The leopard's yellow coat tends to more pale and cream colored 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 colored and of a softer, downy type. Solid black spots in place of open rosettes are generally seen along the face, limbs and underbelly.

Variant coloration

A melanistic leopard, or "black panther"
Male Persian leopard with an atypical coat pattern (Wilhelma, Germany)
See also: Black panther

A melanistic morph of the leopard occurs, particularly in mountainous areas and rain forests. The black color is heritable and caused by recessive gene loci.[20] (While they are commonly called black panthers, the term is not exclusive to leopards; it also applies to melanistic jaguars.)

Melanistic leopards are particularly common on the Malayan Peninsula: early reports suggested up to half of all leopards there are black, but a 2007 camera-trap study in Taman Negara National Park found that all specimens were melanistic.[21] Although the benefits of melanism are difficult to interpret, it may serve as camouflage in the rainforest habitat. Genetic research has found four independent origins for melanism in cats, suggesting that there may be an adaptive advantage.[20] Another possibility is that the color variation is a relic adaptation to an epidemic; genes causing melanism can also affect the immune system.[21]

In Africa, black leopards are much less common as melanism is not an adaptive advantage on the savanna: dark coloration provides poor camouflage and makes hunting difficult. In the dense forests of the Ethiopian Highlands, however, the black leopard is much more common than in Africa generally; as many as one in five leopards may be melanistic.[22]

Pseudo-melanism (abundism) occurs in leopards. A pseudo-melanistic leopard has a normal background color, but the spots are more densely packed than normal and merge to obscure the golden-brown 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.[23]

Biology and behavior

Leopard resting on a tree
Leopard standing in a tree
Leopard kill hanging from a tree

The leopard is known for its ability in climbing, and it has been observed resting on tree branches during the day, dragging its kills up trees and hanging them there, and descending from trees headfirst.[24] 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 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.[25] Leopards produce a number of vocalizations, including grunts, roars, growls, meows and "sawing" sounds.[26]

Leopards are generally considered to be nocturnal; however, the animal has primarily been studied in open savannah habitats, which may have biased common descriptions. Activity level may vary depending on the habitat and the type of prey it hunts. For instance, 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.[27]

Diet and hunting

Leopards are versatile, opportunistic hunters. In the open savannah, they are most successful when hunting between sunset and sunrise, though they may hunt during the day, especially in forest areas when they have the advantage of being hidden by dense brush or cloudy skies.[15] 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,[28] and are capable of carrying animals up to three times their own weight this way.[15] The leopard is the only big cat known to carry its prey up into a tree.

Leopards have relatively flexible dietary needs and generally feed on a greater diversity of prey compared to other members of the Panthera species.[15] Although mid-sized animals are preferred, the leopard will eat anything from dung beetles to 900 kg (1,984 lb) male giant elands.[15] Their diet consists mostly of ungulates and monkeys, but rodents, reptiles, amphibians, insects, birds and fish are also eaten.[29] One survey of nearly 30 research papers conducted by Hayward et al. (2006) 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.[30]

In Africa, mid-sized antelopes provide a majority of the leopard's prey, especially impala and Thomson's gazelles.[28] In Asia the leopard 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 favoured prey of the leopard was the chital.[31] A study at the Wolong Reserve in China revealed how adaptable the leopard's hunting behaviour is: over the course of seven years the vegetative cover receded, and the animals opportunistically shifted from primarily consuming tufted deer to instead pursuing bamboo rats and other smaller prey.[32]

Reproduction and life cycle

Depending on the region, leopards may mate all year round (Asia and Africa) or seasonally during January and February (Manchuria and Siberia). The estrous cycle lasts about 46 days and the female usually is in heat for 6–7 days.[33] Gestation lasts for 90–105 days.[17] Cubs are usually born in a litter of 2–4 cubs,[34] but usually no more than 1–2 cubs survive their first year as the infant mortality rate is between 40 to 50 percent.[15]

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.[35] 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.[24]

Leopards have been reported to reach 21 years of age in captivity.[36]

Social structure and home range

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[3]

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.[37] In their IUCN survey of the literature, Nowell and Jackson suggest male home territories vary between 30–78 square kilometers (km2), but just 15–16 km2 for females.[15] 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.[38] 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 5 to 7 km2 when young cubs were present, while the sexual difference in range size seemed to be in positive proportion to overall increase.[39] However, significant variations in the 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.[37] 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.[40]

The leopard is solitary and, aside from mating, interactions between individuals appear to be infrequent.[40] 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.[41]

Ecology

Distribution and habitat

Data from 1996 found that the leopard has the largest distribution of any wild cat,[15] occurring most in certain parts of southern Asia and widely in eastern and central Africa, although populations before and since have shown a declining trend and are fragmented outside of subsaharan Africa. The IUCN notes that within sub-Saharan Africa the species is "still numerous and even thriving in marginal habitats" where other large cats have disappeared, but that populations in North Africa may be extinct. In Asia, data on distribution is not consistent: populations in Southwest and Central Asia are small and fragmented; in the northeast portion of the range, they are critically endangered; but in the Indian subcontinent, Southeast Asia, and China, the leopard is still relatively abundant.[2] Of the species as a whole, its numbers are greater than those of other Panthera species, all of which face more acute conservation concerns.[2]

Leopards live mainly in grasslands, woodlands and riverside forests. While they are usually associated with the savanna and rainforest, leopards are exceptionally adaptable: in the Russian Far East, the animal inhabits temperate forests where winter temperatures reach a low of −25 °C.[12]

Ecological role

Leopards must compete for food and shelter with other large predators like lions, tigers, spotted hyenas and both African and Asiatic wild dogs. These animals may steal the leopard's kill or devour its young.[15] A single lion or tiger can kill an adult leopard. 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. [42]. [43]. Leopards have adapted to live alongside these other predators by hunting for different types of prey and by avoiding areas frequented by them.[15] 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.[44]

Nowell and Jackson note that resource portioning occurs where the leopard shares its range with the lion or tiger: the leopard tends to take smaller prey (usually less than 75 kg) where its large cousins are present.[15] 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.[45]

Hybrids

Pumapard, 1904

Crossbreeding between leopards and other members of the Panthera genus has been documented resulting in hybrid species. A cross between a female 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.[46] Although lions and leopards may come in to contact in sub-Saharan Africa, they are not widely believed to interbreed naturally. However, there have been anecdotal reports of lion-leopard crosses, known as "marozis", in several African countries.

Crossbreeding between jaguars and leopards in captivity has also been documented.[47][48] A cross between a female leopard and a male jaguar is referred to as a jagupard, the reverse is known a leguar; however, a 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 the jaguar live. Results from leopard-tiger matings have not been known to produce live offspring.

A pumapard is a hybrid animal resulting from a mating between a leopard and a puma (a member of the Puma genus, not the Panthera genus). 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.[49]

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.[49]

Leopards and humans

Leopards have been known to humans since prehistory 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.

Leopard domestication has also been recorded – several leopards were kept in a menagerie established by King John at the Tower of London in the 13th century; in around 1235 three of these animals were given to Henry III by the Holy Roman Emperor Frederick II.[50]

Heraldry

Coat of arms of the German state of Baden-Württemberg

The lion passant guardant or leopard is a frequently used charge in heraldry, most commonly appearing in groups of three.[51] 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 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.[52]

Tourism

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

Park reserves in several countries operate wildlife touring programs that allow visitors to observe leopards in their natural habitat. The Sabi Sands Private Game Reserve in South Africa is one such establishment that offers safari ventures. Sri Lanka offers two leopard habitats, Yala National Park and Wilpattu National Park, where wildlife tours are available. In India, leopards can be seen in the Madhya Pradesh and Uttarakhand national parks.

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 make leopard sightings rare.[53] For example, 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.[54]

Man-eating

The Panar Leopard, shot by Jim Corbett in 1910 after killing 400 people

While most leopards avoid people, humans may occasionally be targeted as prey. Most healthy leopards prefer wild prey to humans, but injured, sickly or struggling cats with a shortage of regular prey may resort to hunting humans and become habituated to it. Two extreme cases occurred in India: the first leopard, "the Leopard of Rudraprayag", may have killed over 125 people; the second, the "Panar Leopard", was believed to have killed more than 400, after injury by a poacher made it unable to hunt normal prey.[55][56] Both were killed by the hunter Jim Corbett.[57] Man-eating leopards are considered bold by feline standards and may enter human settlements for prey, more so than lions and tigers.[58] Author and big game hunter 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.
—Kenneth Anderson, Nine Man-Eaters and One Rogue, Chapter II The Spotted Devil of Gummalapur

Notes

  1. ↑ 1.0 1.1 Wozencraft, W. Christopher (16 November 2005). "Order Carnivora (pp. 532-628)". In Wilson, Don E., and Reeder, DeeAnn M., eds. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press, 2 vols. (2142 pp.). p. 548. ISBN 978-0-8018-8221-0. OCLC 62265494. http://www.bucknell.edu/msw3/browse.asp?id=14000250. 
  2. ↑ 2.0 2.1 2.2 2.3 Henschel, P., Hunter, L., Breitenmoser, U., Purchase, N., Packer, C., Khorozyan, I., Bauer, H., Marker, L., Sogbohossou, E., Breitenmoser-Wursten, C. (2008). "Panthera pardus". IUCN Red List of Threatened Species. Version 2010.1. International Union for Conservation of Nature. http://www.iucnredlist.org/apps/redlist/details/15954. 
  3. ↑ 3.0 3.1 3.2 "Animal bytes: Leopard". Zoological Society of San Diego. http://www.sandiegozoo.org/animalbytes/t-leopard.html. Retrieved 13 February 2010. 
  4. ↑ Partridge, 1983, p. 349
  5. ↑ 5.0 5.1 Monier-Williams, M. (2005). A Sanskrit-English dictionary: etymologically and philologically arranged. Motilall Baransidass Publishers. ISBN 81-208-3105-5. http://books.google.com/?id=zUezTfym7CAC&pg=PA1&dq=Monier-Williams&q=. 
  6. ↑ "Panther". Merriam-Webster Online Dictionary. Retrieved 3 April 2010.
  7. ↑ 7.0 7.1 Partridge, 1983, p. 467
  8. ↑ "Panther". Online Etymology Dictionary. Douglas Harper. http://www.etymonline.com/index.php?term=panther. Retrieved 3 April 2010. 
  9. ↑ 9.0 9.1 9.2 9.3 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 (5757): 73–77. doi:10.1126/science.1122277. PMID 16400146. 
  10. ↑ 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. PMID 15804417. http://202.203.208.84/SCI/2005SCI/2005/Li%20Yu%EF%BC%882005M.P.E).pdf. 
  11. ↑ Bininda-Emonds, Olaf R. P.; Decker-Flum, Denise M.; et al. (2001). "The utility of chemical signals as phylogenetic characters: an example from the Felidae" (PDF). Biological Journal of the Linnean Society 72: 1–15. doi:10.1111/j.1095-8312.2001.tb01297.x. http://www.personal.uni-jena.de/~b6biol2/Publications/FatCats.pdf. Retrieved 2008-06-07. 
  12. ↑ 12.0 12.1 12.2 12.3 Uphyrkina, O.; Johnson, E.W.; et al. (November 2001). "Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus". Molecular Ecology 10 (11): 2617–2633. doi:10.1046/j.0962-1083.2001.01350.x. PMID 11883877. 
  13. ↑ Miththapala, Sriyanie; Seidensticker, John; O'Brien, Stephen J. (August 1996). "Phylogeographic Subspecies Recognition in Leopards (P. pardus): Molecular Genetic Variation.". Conservation Biology 10 (4): 1115–1132. doi:10.1046/j.1523-1739.1996.10041115.x. http://www.blackwell-synergy.com/links/doi/10.1046/j.1523-1739.1996.10041115.x. Retrieved 2008-06-06. 
  14. ↑ 14.0 14.1 Khorozyan, I. G., Gennady, F., Baryshnikov, G. F. and Abramov, A. V. 2006. Taxonomic status of the leopard, Panthera pardus (Carnivora, Felidae) in the Caucasus and adjacent areas. Russian Journal of Theriology 5(1): 41–52.
  15. ↑ 15.00 15.01 15.02 15.03 15.04 15.05 15.06 15.07 15.08 15.09 15.10 15.11 15.12 15.13 Nowell, K. and Jackson, P., (1996), p. 24 – 29.
  16. ↑ 16.0 16.1 "Leopard". African Wildlife Foundation. http://www.awf.org/content/wildlife/detail/leopard. Retrieved 2007-09-21. 
  17. ↑ 17.0 17.1 Kindersley, Dorling (2001, 2005). Animal. New York City: DK Publishing. ISBN 0-7894-7764-5. 
  18. ↑ Nowell, K. and Jackson, P. (1996) p. 44-47.
  19. ↑ "Jaguar (panthera onca)". Our animals. Akron Zoo. http://www.akronzoo.org/learn/jaguar.asp. Retrieved 27 March 2010. 
  20. ↑ 20.0 20.1 Eizirik, Eduardo; Yuhki, Naoya; et al. (May 2003). "Molecular genetics and evolution of melanism in the cat family". Current Biology 13 (5): 448–453. doi:10.1016/S0960-9822(03)00128-3. PMID 12620197. http://cat.inist.fr/?aModele=afficheN&cpsidt=14600002. Retrieved 2008-06-07. 
  21. ↑ 21.0 21.1 Sunquist, Fiona (Dec/Jan 2007). "Malaysian Mystery Leopards". National Wildlife Magazine 45 (1). http://www.nwf.org/nationalwildlife/printerFriendly.cfm?issueID=112&articleID=1413. Retrieved 2008-06-07. 
  22. ↑ "Melanism in Felines". Wildlife of Ethiopia. Biological Society of Ethiopia. April 2001. 
  23. ↑ Gamble, Cyndi; Rodney Griffiths (2004). Leopards: Natural History & Conservation. Voyageur Press. ISBN 0896586561. 
  24. ↑ 24.0 24.1 "Leopard (Panthera pardus); Physical characteristics and distribution". Comparative Mammalian Brain Collections. http://brainmuseum.org/Specimens/carnivora/leopard/index.html. Retrieved 7 June 2008. 
  25. ↑ "Animal bytes – Panthera pardus". Sea World. http://www.seaworld.org/animal-info/Animal-bytes/animalia/eumetazoa/coelomates/deuterostomes/chordata/craniata/mammalia/carnivora/leopard.htm. Retrieved 6 June 2008. 
  26. ↑ Estes, R. (1991). The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores, Primates Los Angeles, The University of California Press.
  27. ↑ Jenny, David; Zuberbühler, Klaus (September 2005). "Hunting behaviour in West African forest leopards". African Journal of Ecology 43 (3): 197–200. doi:10.1111/j.1365-2028.2005.00565.x. http://www.blackwell-synergy.com/doi/abs/10.1111/j.1365-2028.2005.00565.x. Retrieved 2008-05-23. 
  28. ↑ 28.0 28.1 Schaller, 1972, p. 291
  29. ↑ Schaller, 1972, p. 290
  30. ↑ Hayward, M. W.; Henschel, P.; et al. (October 2006). "Prey preferences of the leopard (Panthera pardus)". Journal of Zoology 270 (2): 298–313. doi:10.1111/j.1469-7998.2006.00139.x. http://www.blackwell-synergy.com/doi/abs/10.1111/j.1469-7998.2006.00139.x. Retrieved 2007-09-13. 
  31. ↑ Arivazhagan C., Arumugam R., and Thiyagesan K. (2005). JBNHS Vol. 104(2), Food habits of Leopard (Panthera Pardus Fusca) Dhole (Cuon Alpinus) and striped Hyena (Hyaena Hyaena) in a tropical dry thorn forest of Southern India. Journal of the Bombay National Historical Society. Vol, 104, issue 2.
  32. ↑ Johnson, Kenneth G.; Weng, Wei; et al. (August 1993). "Food Habits of Asiatic Leopards (Panthera pardus fusea) in Wolong Reserve, Sichuan, China". Journal of Mammalogy 74 (3): 646–650. doi:10.2307/1382285. http://www.jstor.org/stable/1382285. Retrieved 2008-06-13. 
  33. ↑ Sadleir R (1966). "Notes on the Reproduction of the larger Felidae". Int. Zoo Yearbook: Vol 6. London: Zool. Soc. London. pp. 184–87. 
  34. ↑ "Leopard". International Society Endangered Cats. 2008. http://www.wildcatconservation.org/Leopard_(Panthera_pardus).html. Retrieved 6 June 2008. 
  35. ↑ Sunquist and Sunquist, 2002, p. 331
  36. ↑ Crandall, L (1964). The management of wild animals in captivity. Chicago: University of Chicago Press. 
  37. ↑ 37.0 37.1 Marker, L. L.; Dickman, A. J. (March 2005). "Factors affecting leopard (Panthera pardus) spatial ecology, with particular reference to Namibian farmlands" (PDF). South African Journal of Wildlife Research 35 (2). http://www.cheetah.org/ama/orig/leopard.pdf. Retrieved 2007-09-13. 
  38. ↑ Mizutani, F.; Jewell, P. A. (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. http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=40975. Retrieved 2007-09-13. 
  39. ↑ Odden, Morten; Wegge, Per (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. http://wild-cat.org/pardus/infos/Odden+Wegge2005-Leopards-spacing+activity-BardiaNP.pdf. Retrieved 2007-09-13. 
  40. ↑ 40.0 40.1 Jenny, D. (November 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. doi:10.1111/j.1469-7998.1996.tb05296.x. http://md1.csa.com/partners/viewrecord.php?requester=gs&collection=ENV&recid=4026878&q=Panthera+pardus+habitat&uid=1040057&setcookie=yes. Retrieved 2007-09-13. 
  41. ↑ Hunter, Luke; Balme, Guy; et al. (2003). "The landscape ecology of leopards (Panthera pardus) in northern KwaZulu-Natal, South Africa: A preliminary project report." (PDF). Ecological Journal 5: 24–30. http://www.biolsci.monash.edu.au/research/leopards/docs/ecojournal-2003.pdf. Retrieved 2007-09-16. 
  42. ↑ Owens, Mark and Owens, Delia. Cry of the Kalahari. 1984. p133-5.
  43. ↑ "Hyenas of the Kalahari". Owens, Delia and Owens, Mark. Natural History. 89(2):p50 (February 1980)
  44. ↑ Schaller, 1972, p. 293
  45. ↑ Karanth, K. Ullas; Sunquist, Melvin E. (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. doi:10.1111/j.1469-7998.2000.tb01076.x. 
  46. ↑ McCarthy, Eugene M. Leopon: Lioness x Leopard Macroevolution.net. Accessed 18 March 2010.
  47. ↑ Jaguar Panthera onca. Zoological Society of San Diego Zoo. Accessed 28 March 2010.
  48. ↑ Hartwell, S. Hybrids involving leopards and jaguars.Messybeast.com. Britain. Accessed 28 March 2010.
  49. ↑ 49.0 49.1 Jen Stevenson. "Le Pumapard". The Strand. http://media.www.thestrand.ca/media/storage/paper404/news/2005/03/17/Features/Le.Pumapard-891758.shtml. Retrieved 16 June 2008. 
  50. ↑ Owen, James (November 3, 2005). "Medieval Lion Skulls Reveal Secrets of Tower of London "Zoo"". National Geographic Magazine. National Geographic. http://news.nationalgeographic.com/news/2005/11/1103_051103_tower_lions.html. Retrieved 2007-09-05. 
  51. ↑ Strickland, Debra Higgs; Debra Hassig (1999). The Mark of the Beast: The Medieval Bestiary in Art, Life, and Literature. Taylor & Francis. ISBN 0815329520. 
  52. ↑ Pedersen, Christian Fagd (1971). The International Flag Book in Color. Morrow. 
  53. ↑ "In South Africa, It’s All in the Game". The New York Times. 24 October 2004. http://travel.nytimes.com/2004/10/24/travel/24timbavati.html. Retrieved 20 March 2010. 
  54. ↑ U.M.I.R.K. Weerasinghe, Dayananda Kariyawasm and Mangala De Zoysa “Ruhuna (Yala) National Park in Sri Lanka: Visitors, Visitation, and Eco-Tourism”. University of Ruhuna
  55. ↑ Hart, Donna; Robert W. Sussman (2005). Man the Hunted: Primates, Predators, and Human Evolution. Westview Press. ISBN 0813339367. 
  56. ↑ Tougias, p.147
  57. ↑ Capstick, Peter Hathaway (1978). Death in the Long Grass. St. Martin's Press. ISBN 0312186134. 
  58. ↑ Anderson, Kenneth. Nine Man-Eaters and one Rogue, Allen & Unwin Ltd, 1954.

References

External links