Temporal range: Late Pliocene or Early Pleistocene to recent
(Panthera pardus pardus)
|Current range of the leopard, former (red), uncertain (yellow), highly fragmented (light green), and present (dark green)|
Felis pardus Linnaeus, 1758
The leopard (Panthera pardus) // is one of the five "big cats" in the genus Panthera. It is a member of the family Felidae with a wide range in sub-Saharan Africa and parts of Asia. Fossil records suggest that in the Late Pleistocene it occurred in Europe and Japan.
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 has a smaller, lighter physique. 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 is distinguished by its well-camouflaged fur, opportunistic hunting behaviour, broad diet, and strength (which it uses to move heavy carcasses into trees), as well as its ability to adapt to various habitats ranging from rainforest to steppe, including arid and montane areas, and its ability to run at speeds of up to 58 kilometres per hour (36 mph).
The leopard is listed as vulnerable on the IUCN Red List because leopard populations are threatened by habitat loss and fragmentation, and are declining in large parts of the global range. In Hong Kong, Singapore, Kuwait, Syria, Libya, Tunisia and most likely in Morocco, leopards have already been extirpated. Leopards are hunted illegally, and their body parts are smuggled in the wildlife trade for medicinal practices and decoration.
- 1 Etymology
- 2 Taxonomy
- 3 Evolution
- 4 Genetics
- 5 Characteristics
- 6 Distribution and habitat
- 7 Ecology and behaviour
- 8 Leopards and humans
- 9 References
- 10 Further reading
- 11 External links
The common name "leopard" // is a Greek compound of λέων leōn ("lion") and πάρδος pardos ("male panther"). The name reflects the fact that in antiquity, a leopard was believed to be a hybrid of a lion and a panther. The Greek word is related to Sanskrit पृदाकु pṛdāku ("snake", "tiger" or "panther"), and probably derives from a Mediterranean language, such as Egyptian. The name was first used in the 13th century. Other vernacular names for the leopard include graupanther, panther and several regional names such as tendwa in India, the term "black panther" refers to leopards with melanistic genes. A term for the leopard used in Old English and later, but now very uncommon, is "pard".
The scientific name of the leopard is Panthera pardus, the generic name Panthera derives from Latin via Greek πάνθηρ (pánthēr). 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. 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). The specific name pardus is derived from the Greek πάρδος (pardos) ("male panther").
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.
The leopard was first described by Swedish zoologist Carl Linnaeus in the 10th edition of Systema Naturae (1758). Linnaeus named the leopard 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. 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; in 1916, British zoologist Reginald Innes Pocock accorded Panthera generic rank defining Panthera pardus as species.
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.  Results of a 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. This was seconded by a 2009 study by Lars Werdelin and colleagues. 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. Results of a 2001 phylogenetic analysis of chemical secretions amongst cats suggested, however, that the leopard is closely related to the lion.
Following Linnaeus's first description, 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. Later analysis revealed a ninth valid subspecies, the Arabian leopard. However, this number might be an underestimation; limited sampling of the African leopards means that different subspecies may not have been resolved.
The nine subspecies recognised by the International Union for Conservation of Nature (IUCN) are summarised in the following table. Since 2017, the Cat Classification Task Force of the Cat Specialist Group recognises only eight subspecies, subsuming the Persian leopard to P.p. tulliana.
|African leopard (P. p. pardus) (Linnaeus, 1758), syn. P. p. panthera (Schreber, 1777), P. p. leopardus (Schreber, 1777), P. p. melanotica (Gunther, 1885), P. p. suahelicus (Neumann, 1900), P. p. nanopardus (Thomas, 1904), P. p. chui (Heller, 1913), P. p. reichenowi (Odbrera, 1918), P. p. antinorii (de Beaux, 1923), P. p. iturensis (Allen, 1924), P. p. shortridgei (Pocock, 1932), P. p. adersi (Pocock, 1932)||It lives in Sub-Saharan Africa and is the most widespread leopard subspecies.
|Indian leopard (P. p. fusca) (Meyer, 1794), syn. P. p. pernigra (Hodgson, 1863), P. p. millardi (Pocock, 1930)||It is native to the Indian Subcontinent: India, Nepal, Bhutan, Pakistan and Bangladesh.|
|Javan leopard (P. p. melas) (G. Cuvier, 1809)||It is the only subspecies native to Indonesia and lives on Java. It is Critically Endangered.|
|Arabian leopard (P. p. nimr) (Hemprich and Ehrenberg, 1833), syn. P. p. jarvisi (Pocock, 1932)||It is the smallest leopard subspecies; adult females weigh about 18 kg (40 lb). It is native to the Arabian Peninsula: Saudi Arabia, Oman, Yemen, Kuwait, the United Arab Emirates, Israel, Jordan, Lebanon, Syria; extinct in the Sinai Peninsula.|
|Anatolian leopard (P. p. tulliana) (Valenciennes, 1856), syn. Persian leopard (P. p. ciscaucasica) (Satunin, 1914), P. p. saxicolor Pocock, 1927), Baluchistan leopard (P. p. sindica Pocock, 1930), P. p. dathei Zukowsky, 1964)||Leopard populations persist in eastern Turkey, the Caucasus, Georgia, Armenia, southern Russia, Azerbaijan, Turkmenistan, Uzbekistan, Afghanistan, Iraq, and northern Iran.
In south-western Turkey, the leopard is extinct, the Baluchistan leopard possibly evolved in southern Iran, southern Afghanistan and south-western Pakistan being separated from the northern population by Dasht-e Kavir and Dasht-e Lut deserts.
|North Chinese leopard (P. p. japonensis) (Gray, 1862)||It is native to central and northern China, where today only small and isolated populations remain.|
|Amur leopard (P. p. orientalis) (Schlegel, 1857)||It lives in the cold regions of the Russian Far East and Northeast China and is Critically Endangered. It is currently extinct in the Korean Peninsula.|
|Indochinese leopard (P. p. delacouri) Pocock, 1930||It inhabits mainland Southeast Asia, including Myanmar, Thailand, Malaysia, Cambodia, Laos, Vietnam and South China.|
|Sri Lankan leopard (P. p. kotiya) Deraniyagala, 1956||It is native to Sri Lanka.|
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.
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. Johnson and colleagues suggest that the leopard diverged next, and followed by the lion-jaguar clade.
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.
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, and in Biśnik Cave in south-central Poland. 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.
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. The chromosomes include four acrocentric, five metacentric, seven submetacentric and two telocentric pairs.
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 a juvenile leopard, the hybrid came to be known as "leopon". Unsuccessful attempts were made to mate a leopon with a tigress.
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.
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 a dwarf, smaller than either parent, the coat is variously described as sandy, tawny or greyish with brown, chestnut or faded rosettes.
Melanistic leopards are known, like melanistic jaguars, as "black panthers". Pseudomelanism (abundism) also occurs in leopards. Melanism in leopards is inherited as a trait relatively recessive to the spotted form. Interbreeding in melanistic leopards produces a significantly smaller litter size than is produced by normal pairings.
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. 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.
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. Pseudomelanism has also been reported in leopards.
The leopard is a big cat distinguished by its robust build and muscular but relatively shorter limbs, a broad head and a coat covered by spots arranged in rosettes. Males stand 60–70 cm (24–28 in) at the shoulder, while females are 57–64 cm (22–25 in) tall. The head-and-body length is typically between 90 and 190 cm (35 and 75 in). While males weigh 37–90 kg (82–198 lb), females weigh 28–60 kg (62–132 lb); these measurements vary geographically. The maximum recorded weight for a leopard is 96.5 kilograms (213 lb). Sexually dimorphic, males are larger and heavier than females.
Basically pale yellow to yellowish brown or golden (except for the melanistic forms), the coat is spotted and rosetted; spots fade toward the white underbelly and the insides and lower parts of the legs. Rosettes are most prominent on the back, flanks and hindquarters, the pattern of the rosettes is unique to each individual. Juveniles have woolly fur, and appear dark due to the densely arranged spots, the white-tipped tail, 60–100 centimetres (24–39 in) long, white underneath, displays rosettes except toward the end, where the spots form incomplete bands.
The texture and colour of the fur varies by climate and geography; leopards in forests are observed to be darker than those in deserts. The guard hairs (the layer of hairs that protect the basal hairs) are the shortest (3–4 millimetres (0.12–0.16 in)) on the face and the head, and increase in length toward the flanks and the underparts (25–30 millimetres (0.98–1.18 in)). The fur is generally soft and thick; the fur on the underparts is notably softer than that on the back. A few geographical variations have been noted in the colour and texture of the fur. Leopards in forests tend to be darker than those in deserts; the fur tends to grow longer in populations living in colder climates. The rosettes, circular in eastern African populations, tend to be squarish 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.
The leopard is often confused with the cheetah; however, the cheetah is marked with small round spots instead of the larger rosettes. Moreover, the leopard lacks the facial tear streaks characteristic of the cheetah. Other similar species are the clouded leopard and jaguar, the clouded leopard can be told apart by the diffuse "clouds" of spots compared to the smaller and distinct rosettes of the leopard, longer legs and thinner tail. The jaguar has rosettes that typically have spots within them, while those of leopards often do not. Moreover, the jaguar has larger and rounder foot pads and a larger and stronger skull.
Distribution and habitat
Leopards have the largest distribution of any wild cat, occurring widely in Africa as well as eastern and southern Asia, although populations have shown a declining trend, and are fragmented outside of sub-Saharan Africa. Within sub-Saharan Africa, the species is still numerous and even thriving in marginal habitats where other large cats have disappeared. Populations in North Africa may be extinct. 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.
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). 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; in India, leopard populations sometimes live quite close to human settlements and even in semi-developed areas. 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. Due to the leopard's superlative stealthiness, people often remain unaware that big cats live in nearby areas.
Ecology and behaviour
Leopards, like lions and tigers, tend to be nocturnal (active mainly at night). However, leopards in western African forests have been observed to be largely diurnal and hunt during twilight, when their prey animals are active; activity patterns may even vary by season. Leopards generally are active mainly from dusk till dawn, and rest for most of the day and for some hours at night in thickets, among rocks or over tree branches. Leopards have been observed walking 1–25 kilometres (0.62–15.53 mi) across their range at night; they may even wander up to 75 kilometres (47 mi) if disturbed.
Leopards are known for their ability to climb 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, 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.
The leopard is solitary and territorial, as are several other felids; individuals associate appreciably only in the mating season, though mothers may continue to interact with their offspring even after weaning. Mothers have been observed sharing kills with their offspring when they can not obtain any meal; in Kruger National Park, most leopards tend to keep 1 kilometre (0.62 mi) apart. Fathers may interact with their partners and cubs at times. Aggressive encounters are rare, typically limited to defending territories from intruders; in a South African reserve, a male was wounded in a male–male territorial battle over a carcass. A few instances of cannibalism have been reported.
Leopards communicate with each other in tall grass using white spots on their ears and tails, they also produce a number of vocalisations, including growls, snarls, meows and purrs. The roaring sequence in leopards consists mainly of grunts and is also known called "sawing", having been described as resembling the sound of sawing wood. Cubs are known to call their mother with a urr-urr sound.
Males occupy territories that often overlap with a few smaller female territories, probably as a strategy to enhance access to females. A radio-collar analysis in the Ivory Coast found a female home range completely enclosed within a male's. Female live with their cubs in territories that overlap extensively – probably due to the association between mothers and their offspring. There may be a few other fluctuating territories, belonging to young individuals, it is not clear if male territories tend to overlap among themselves as much as those of females do. Individuals will try to drive away intruders of the same sex.
A study of leopards in the Namibian farmlands showed that the size of territories was not significantly affected by sex, rainfall patterns or season; it concluded that the higher the prey availability in an area, the greater the population density of leopards and the smaller the size of territories, but territories tend to expand if there is human interference (which has been notably high in the study area). Territorial sizes vary geographically; they can be as small as 33–38 square kilometres (13–15 sq mi) for males and 14–16 square kilometres (5.4–6.2 sq mi) for females in forests and rocky terrain (such as in the Serengeti or Kruger National Park), or as large as 451 square kilometres (174 sq mi) for males and 188 square kilometres (73 sq mi) for females in northeastern Namibia (they might be even larger in deserts and montane areas). Territories recorded in Nepal, 48 square kilometres (19 sq mi) for males and 5–7 square kilometres (1.9–2.7 sq mi) for females, are smaller than those generally observed in Africa.
Hunting and diet
The leopard is a carnivore that prefers medium-sized prey with a body mass ranging from 10–40 kilograms (22–88 lb). A study noted that prey animals in this weight range tend to occur in dense habitat, form small herds and can be easily captured by the leopard; on the other hand, animals that prefer open areas and have developed significant anti-predator strategies are hardly preferred. Prey as heavy as 150 kilograms (330 lb) (such as greater kudu and giraffe) may be hunted if larger carnivores such as lions or tigers are absent; the largest prey killed by a leopard was reportedly a 900 kg (2,000 lb) male eland. Leopards can feed on a broad variety of prey, mainly antelopes, deer and rodents; these include: cattle, chital, duiker, dung beetle, hartebeest, hyrax, impala, gazelle, muntjac, nyala, porcupine, primates, rat, reedbuck, springbok, sambar, squirrel, waterbuck, young zebras, warthog and wildebeest. Females with cubs primarily target smaller prey.
A study at Wolong Reserve in China demonstrated variation in the leopards' diet over time; 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. A study estimated average daily consumption rates at 3.5 kg (7.7 lb) for males and 2.8 kg (6.2 lb) for females. A study of leopards in the southern Kalahari showed that water requirements are met by the bodily fluids of the prey, succulent plants and water bodies; they drink water every two to three days, and feed infrequently on moisture-rich plants such as gemsbok cucumbers (Acanthosicyos naudinianus), tsamma melon (Citrullus lanatus) and Kalahari sour grass (Schmidtia kalahariensis). A few instances of cannibalism have been reported.
The leopard depends mainly on its acute sense of hearing and vision for hunting. Hunting is primarily a nocturnal activity in most areas, though leopards in western African forests and Tsavo have been observed hunting by the day, it will stalk the prey and try to approach as close as possible (typically within 5 metres (16 ft)) to the target, and finally pounce on it with its forepaws (unlike the lion, that pounces as the prey starts escaping) and kill it by suffocation. Small prey are killed with a bite on the back of the neck, while larger animals are held strongly by the throat and strangled.
Small kills are eaten immediately, while larger carcasses are dragged over several hundred metres and safely cached to be consumed later on trees, in bushes or even caves, the way the kill is stored to be consumed later depends on the local topography and individual preferences; while trees are preferred in Kruger National Park, bushes are preferred in the plain terrain of the Kalahari. Kills are cached up to 2 kilometres (6,600 ft) apart. Although they are smaller than most other members of its genus, leopards are able to take large prey due to their massive skulls that facilitate powerful jaw muscles. Leopards are strong enough to drag carcasses heavier than themselves up trees; an individual was seen to haul a young giraffe, nearly 125 kg (276 lb), up 5.7 m (19 ft) into a tree.
Enemies and competitors
Leopards must compete for food and shelter with other large predators such as tigers, lions, cheetahs, 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, caracal, African wild cat and the cubs of lions, cheetahs, hyenas, and wild dogs. 
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. 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, the mean leopard density decreased significantly (from 9.76 to 2.07 animals per 100 km2) 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. 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%). In the Primore region of the Russian Far East, Amur leopards were absent or very rarely encountered at places where Siberian tigers reside. 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. 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. 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  Usually when a tiger began to kill baits at sites formerly frequented by leopards, the leopards would no longer come and kill there; 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. 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. 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.
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. 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. Mugger crocodiles may also on rare occasions kill an adult leopard in India. Lions are occasionally successful in climbing trees and fetching leopard kills, but leopards are also known to kill or prey on cubs of lions. 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. 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.
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. Gestation lasts for 90 to 105 days. Cubs are usually born in a litter of 2–4 cubs. Mortality of cubs is estimated at 41–50% during the first year.
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, 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.
The average typical life span of a leopard is between 12 and 17 years, 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. 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. 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.
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 sculptures 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, totems and sacrifices to deities, as a result of their association with kings in Africa, the leopard's pelt is often seen today as a symbol of aristocratic rank, chiefs using it as a part of their traditional regalia.
The lion passant guardant or leopard is a frequently used charge in heraldry, most commonly appearing in groups of three, 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.
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.
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; 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.
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.
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 and conservationist Jim Corbett. 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. 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"
- Stein, A.B.; Athreya, V.; Gerngross, P.; Balme, G.; Henschel, P.; Karanth, U.; Miquelle, D.; Rostro, S.; Kamler, J.F.; Laguardia, A. (2016). "Panthera pardus". IUCN Red List of Threatened Species. Version 2016-3. International Union for Conservation of Nature. Retrieved 27 August 2016.
- Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 547. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Ghezzo, E.; Rook, L. (2015). "The remarkable Panthera pardus (Felidae, Mammalia) record from Equi (Massa, Italy): taphonomy, morphology, and paleoecology". Quaternary Science Reviews. 110 (110): 131–151. doi:10.1016/j.quascirev.2014.12.020.
- Ohdachi, Satoshi D.; Ishibashi, Yasuyuki; Iwasa, Masahiro A.; Fukui, Dai; Saitoh, Takashi (2015). The Wild Mammals of Japan (2nd ed.). Shoukadoh. ISBN 978-4-87974-691-7.[page needed]
- Nowell, K.; Jackson, P. (1996). "Leopard Panthera pardus (Linnaeus, 1758)". Wild Cats: status survey and conservation action plan. Gland, Switzerland: IUCN/SSC Cat Specialist Group.
- Jacobson, Andrew P.; Gerngross, Peter; Lemeris Jr., Joseph R.; Schoonover, Rebecca F.; Anco, Corey; Breitenmoser-Würsten, Christine; Durant, Sarah M.; Farhadinia, Mohammad S.; Henschel, Philipp; Kamler, Jan F.; Laguardia, Alice; Rostro-García, Susana; Stein, Andrew B.; Dollar, Luke (4 May 2016). "Leopard (Panthera pardus) status, distribution, and the research efforts across its range". PeerJ. 4: e1974. PMC . PMID 27168983. doi:10.7717/peerj.1974.
- Williams, Samual T.; Williams, Kathryn S.; Lewis, Bradley P.; Hill, Russell A. (19 April 2017). "Population dynamics and threats to an apex predator outside protected areas: implications for carnivore management". Royal Society Open Science. 4 (4): 161090. PMID 28484625. doi:10.1098/rsos.161090.
- Bergin, D. and Nijman, V. (2014) Open, Unregulated Trade in Wildlife in Morocco's Markets. TRAFFIC Bulletin
- Bergin, D.; Nijman, V. (2015). "Potential benefits of impending Moroccan wildlife trade laws, a case study in carnivore skins". Biodiversity and Conservation. 25 (1): 199–201. ISSN 0960-3115. doi:10.1007/s10531-015-1042-1.
- "leopard". Merriam-Webster Dictionary. Retrieved 23 March 2016.
- Partridge, Eric (1983). Origins: A Short Etymological Dictionary of Modern English. New York: Greenwich House. p. 349. ISBN 978-0-517-41425-5.
- Monier-Williams, M. (2005). A Sanskrit-English Dictionary: Etymologically and Philologically Arranged. Motilal Baransidas Publishers. ISBN 978-81-208-3105-6.
- Stein, A.B.; Hayssen, V. (2010). "Panthera pardus (Carnivora: Felidae)" (PDF). Mammalian Species. 45 (900): 30–48. doi:10.1644/900.1. Archived from the original (PDF) on 2016-03-05.
- "panther". Merriam-Webster Dictionary. Retrieved 3 April 2010.
- "pard". Oxford English Dictionary (3rd ed.). Oxford University Press. September 2005. (Subscription or UK public library membership required.) OED gives examples of "pard" from OE until 1984.
- Partridge, Eric (1983). Origins: A Short Etymological Dictionary of Modern English. New York: Greenwich House. p. 467. ISBN 978-0-517-41425-5.
- 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. Bibcode:2006Sci...311...73J. PMID 16400146. doi:10.1126/science.1122277.
- Werdelin, L.; Yamaguchi, N.; Johnson, W.E.; O'Brien, S.J. (2010). "Phylogeny and evolution of cats (Felidae)" (PDF). Biology and Conservation of Wild Felids: 59–82.
- Davis, B.W.; Li, G.; Murphy, W.J. (2010). "Supermatrix and species tree methods resolve phylogenetic relationships within the big cats, Panthera (Carnivora: Felidae)" (PDF). Molecular Phylogenetics and Evolution. 56 (1): 64–76. PMID 20138224. doi:10.1016/j.ympev.2010.01.036.
- Mazák, J.H.; Christiansen, P.; Kitchener, A.C.; Goswami, A. (2011). "Oldest known pantherine skull and evolution of the tiger". PLoS ONE. 6 (10): e25483. Bibcode:2011PLoSO...625483M. PMC . PMID 22016768. doi:10.1371/journal.pone.0025483.
- Wilson, D.E.; Reeder, D.M., eds. (2005). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 547–548. ISBN 978-0-8018-8221-0. OCLC 62265494.
- "Pantherinae". Integrated Taxonomic Information System. Retrieved 23 March 2016.
- Linnaeus, C. (1758). Systema Naturae. I (10th ed.). pp. 41–42.
- Ellerman, J. R.; Morrison-Scott, T. C. S. (1966). Checklist of Palaearctic and Indian mammals 1758 to 1946 (2nd ed.). London: British Museum of Natural History. pp. 315–317.
- Pocock, R.I. (1939). "Panthera pardus". The Fauna of British India, Including Ceylon and Burma. Mammalia: Volume 1. London: Taylor and Francis. pp. 222–239.
- Turner, A. (1987). "New fossil carnivore remains from the Sterkfontein hominid site (Mammalia: Carnivora)". Annals of the Transvaal Museum. 34: 319–47.
- Bininda-Emonds, O.R.P.; Decker-Flum, D.M.; Gittleman, J.L. (2001). "The utility of chemical signals as phylogenetic characters: an example from the Felidae" (PDF). Biological Journal of the Linnean Society. 72 (1): 1–15. doi:10.1111/j.1095-8312.2001.tb01297.x. Archived from the original (PDF) on 2010-01-31. Retrieved 2008-06-07.
- Miththapala, S.; Seidensticker, J.; O'Brien, S. J. (1996). "Phylogeographic subspecies recognition in leopards (Panthera pardus): molecular genetic variation" (PDF). Conservation Biology. 10 (4): 1115–1132. doi:10.1046/j.1523-1739.1996.10041115.x.
- Uphyrkina, O.; Johnson, E.W.; Quigley, H.; Miquelle, D.; Marker, L.; Bush, M.; O'Brien, S. J. (2001). "Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus" (PDF). Molecular Ecology. 10 (11): 2617–2633. PMID 11883877. doi:10.1046/j.0962-1083.2001.01350.x.
- Khorozyan, I. G.; Gennady, F.; Baryshnikov, G. F.; Abramov, A. V. (2006). "Taxonomic status of the leopard, Panthera pardus (Carnivora, Felidae) in the Caucasus and adjacent areas" (PDF). Russian Journal of Theriology. 5 (1): 41–52. Archived from the original (PDF) on 2016-03-03.
- Kitchener, A.C., Breitenmoser-Würsten, C., Eizirik, E., Gentry, A., Werdelin, L., Wilting, A. and Yamaguchi, N. (2017). "A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group" (PDF). Cat News. Special Issue 11.
- Soultan, A.; et al. (2016). "Recent observation for leopard Panthera pardus in Egypt". Mammalia. 81. doi:10.1515/mammalia-2015-0089.
- Spalton, J.A. and Al Hikmani, H. M. (2006). "The Leopard in the Arabian Peninsula – Distribution and Subspecies Status" (PDF). Cat News Special Issue 1: 4–8.
- Dou, H., Feng, L., Xiao, W., & Wang, T. (2014). The complete mitochondrial genome of the North Chinese Leopard (Panthera pardus japonensis). Mitochondrial DNA: 1-2.
- Brakefield, T. (1993). Big Cats: Kingdom of Might. ISBN 978-0-89658-329-0.
- Schmid, E. (1940). "Variationstatistische Untersuchungen am Gebiss pleistozäner und rezenter Leoparden und anderer Feliden". Zeitschrift für Säugetierkunde (in German). 15: 1–179.
- Marciszak, A.; Stefaniak, K. (2010). "Two forms of cave lion: Middle Pleistocene Panthera spelaea fossilis Reichenau, 1906 and Upper Pleistocene Panthera spelaea spelaea Goldfuss, 1810 from the Bísnik Cave, Poland". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 258 (3): 339–351. ISSN 0077-7749. doi:10.1127/0077-7749/2010/0117.
- Diedrich, C.G. (2013). "Late Pleistocene leopards across Europe – northernmost European German population, highest elevated records in the Swiss Alps, complete skeletons in the Bosnia Herzegowina Dinarids and comparison to the Ice Age cave art". Quaternary Science Reviews. 76: 167–193. Bibcode:2013QSRv...76..167D. doi:10.1016/j.quascirev.2013.05.009.
- Heptner, V.G. Heptner (1992). Mammals of the Soviet Union. Leiden: Brill. p. 61. ISBN 978-90-04-08876-4.
- Tanomtong, A.; Khunsook, S.; Keawmad, P.; Pintong, K. (2008). "Cytogenetic study of the leopard, Panthera pardus (Carnivora, Felidae) by conventional staining, G-banding and high-resolution staining technique" (PDF). Cytologia. 73 (1): 81–90. doi:10.1508/cytologia.73.81.
- Kisling, V.N., ed. (2001). Zoo and Aquarium History : Ancient Animal Collections to Zoological Gardens. Boca Raton, Florida (USA): CRC Press. p. 314. ISBN 978-0-8493-2100-9.
- Eberhart, G. M. (2002). Mysterious Creatures : A Guide to Cryptozoology. Oxford, UK: ABC-Clio. pp. 514–516. ISBN 978-1-57607-283-7.
- "Geocites – Liger & Tigon Info". Archived from the original on October 15, 2007. Retrieved June 9, 2008.
- Gamble, C.; Griffiths, R. (2004). Leopards: Natural History and Conservation. Voyageur Press. ISBN 978-0-89658-656-7.
- Eizirik, E.; Yuhki, N.; Johnson, W.E.; Menotti-Raymond, M.; Hannah, S.S.; O'Brien, S.J. (2003). "Molecular genetics and evolution of melanism in the cat family" (PDF). Current Biology. 13 (5): 448–53. PMID 12620197. doi:10.1016/S0960-9822(03)00128-3.[permanent dead link]
- Robinson, R. (1970). "Inheritance of the black form of the leopard Panthera pardus". Genetica. 41 (1): 190–7. PMID 5480762. doi:10.1007/BF00958904.
- Searle, A.G. (1968). Comparative Genetics of Coat Colour in Mammals. London: Logos Press.
- Kawanishi, K.; Sunquist, M. E.; Eizirik, E.; Lynam, A. J.; Ngoprasert, D.; Wan Shahruddin, W. N.; Rayan, D. M.; Sharma, D. S. K.; Steinmetz, R. (2010). "Near fixation of melanism in leopards of the Malay Peninsula". Journal of Zoology. 282 (3): 201–206. doi:10.1111/j.1469-7998.2010.00731.x.
- Sunquist, F. (2007). "Malaysian Mystery Leopards". National Wildlife Magazine. 45 (1).
- ""Strawberry" Leopard Discovered—A First". National Geographic. 2012.
- Shuker, K.P.N. (2003). The Beasts that Hide from Man : Seeking the World's Last Undiscovered Animals. New York, USA: Paraview Press. p. 273. ISBN 978-1-931044-64-6.
- Estes, R. (1991). The Behavior Guide to African Mammals, Including Hoofed Mammals, Carnivores, Primates. Los Angeles: The University of California Press. ISBN 978-0-520-08085-0.
- Nowak, R.M. (1999). Walker's Mammals of the World (6th ed.). Baltimore, USA: Johns Hopkins University Press. pp. 828–31. ISBN 978-0-8018-5789-8.
- Burnie, D; Wilson, D.E., eds. (2001). Animal: The Definitive Visual Guide to the World's Wildlife. DK Adult. ISBN 978-0-7894-7764-4.
- Brain, C.K. (1983). The Hunter or the Hunted: An Introduction to African Cave Taphonomy. University of Chicago Press. ISBN 978-0-226-07090-2.
- Hoath, R. (2009). Field Guide to the Mammals of Egypt. Cairo, Egypt: American University in Cairo Press. pp. 106–7. ISBN 978-977-416-254-1.
- Schütze, H. (2002). Field Guide to the Mammals of the Kruger National Park. Cape Town, South Africa: Struik Publishers. pp. 92–3. ISBN 978-1-86872-594-6.
- Menon, V. (2014). Indian Mammals: A Field Guide. Gurgaon, India: Hachette India. ISBN 978-93-5009-761-8.
- Skinner, J.D.; Chimimba, C.T. (2005). The Mammals of the Southern African Subregion (3rd ed.). Cambridge, UK: Cambridge University Press. pp. 397–401. ISBN 978-0-521-84418-5.
- Foley, C.; Foley, L.; Lobora, A.; De Luca, D.; Msuha, M.; Davenport, T.R.B.; Durant, S.M. (2014). A Field Guide to the Larger Mammals of Tanzania. Princeton, USA: Princeton University Press. pp. 132–3. ISBN 978-1-4008-5280-2.
- Mivart, St. G.J. (1900). The Cat: An Introduction to the Study of Backboned Animals, Especially Mammals. London, UK: John Murray. pp. 427–9.
- Francis, C.M. (2008). A Field Guide to the Mammals of Southeast Asia. London, UK: New Holland. p. 296. ISBN 978-1-84537-735-9.
- Murphey, Rhoads (1951). "The Decline of North Africa Since the Roman Occupation: Climatic or Human?" (PDF). Annals of the Association of American Geographers. XLI (2): 116–132. doi:10.1080/00045605109352048.
- Gavashelishvili, A.; Lukarevskiy, V. (2008). "Modelling the habitat requirements of leopard Panthera pardus in west and central Asia". Journal of Applied Ecology. 45 (2): 579–588. doi:10.1111/j.1365-2664.2007.01432.x.
- Athreya, Vidya (2012-08-16) Living with Leopards Outside Protected Areas in India. conservationindia.org
- Tilson, R. (2010). Tigers of the World: The Science, Politics and Conservation of Panthera tigris. Elsevier. p. 22. ISBN 978-0-8155-1570-8.
- Schaller, G.B. (1972). The Serengeti Lion: A Study of Predator-Prey Relations. Chicago, USA: University of Chicago Press. ISBN 978-0-226-73639-6.
- Hunter, L.; Balme, G.; Walker, C.; Pretorius, K.; Rosenberg, K. (2003). "The landscape ecology of leopards (Panthera pardus) in northern KwaZulu-Natal, South Africa: a preliminary project report" (PDF). Ecological Journal. 5: 24–30. Archived from the original (PDF) on February 13, 2010.
- Spalton, J.A.; al Hikmani, H.M.; Willis, D.; Said, A.S.B. (2006). "Critically endangered Arabian leopards Panthera pardus nimr persist in the Jabal Samhan Nature Reserve, Oman". Oryx. 40 (3): 287–94. doi:10.1017/S0030605306000743.
- Jenny, D.; Zuberbuhler, K. (2005). "Hunting behaviour in west African forest leopards" (PDF). African Journal of Ecology. 43 (3): 197–200. doi:10.1111/j.1365-2028.2005.00565.x.
- "Leopard (Panthera pardus); Physical characteristics and distribution". Comparative Mammalian Brain Collections.
- "Animal bytes – Panthera pardus". Sea World. Archived from the original on 24 June 2008. Retrieved 6 June 2008.
- "Animal bytes: Leopard". Zoological Society of San Diego. Retrieved 13 February 2010.
- Kingdon, J.; Happold, D.; Butynski, T.; Hoffmann, M.; Happold, M.; Kalina, J. (2013). Mammals of Africa. London: Bloomsbury Publishing. pp. 159–68. ISBN 978-1-4081-8996-2.
- Hunter, L.; Balme, G.; Walker, C.; Pretorius, K.; Rosenberg, K. (2003). "The landscape ecology of leopards (Panthera pardus) in northern KwaZulu-Natal, South Africa: A preliminary project report" (PDF). Ecological Journal. 5: 24–30. Archived from the original (PDF) on 2010-02-13.
- Steyn, V.; Funston, P.J. (2006). "A case of cannibalism in leopards" (PDF). South African Journal of Wildlife Research. 36 (2): 189–90.
- Bodendorfer, T.; Hoppe-Dominik, B.; Fischer, F.; Linsenmair, K.E. (2006). "Prey of the leopard (Panthera pardus) and the lion (Panthera leo) in the Comoé and Marahoué National Parks, Côte d'Ivoire, West Africa". Mammalia. 70 (3/4): 231–46. doi:10.1515/MAMM.2006.037.
- Sunquist, Mel; Sunquist, Fiona (2002). Wild Cats of the World. University of Chicago Press. p. 423. ISBN 978-0-226-77999-7.
- 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. doi:10.1111/j.1469-7998.1996.tb05296.x.
- Marker, L.L.; Dickman, A.J. (2005). "Factors affecting leopard (Panthera pardus) spatial ecology, with particular reference to Namibian farmlands" (PDF). South African Journal of Wildlife Research. 35 (2): 105–15.
- Mizutani, F.; Jewell, P. A. (1998). "Home-range and movements of leopards (Panthera pardus) on a livestock ranch in Kenya". Journal of Zoology. 244 (2): 269–286. doi:10.1017/S0952836998002118.
- Bertram, B.C.R. (1982). "Leopard ecology as studied by radio tracking". Symposia of the Zoological Society of London. 49: 341–52.
- Stander, P.E.; Haden, P.J.; Kaqece, II.; Ghau, II. (1997). "The ecology of asociality in Namibian leopards". Journal of Zoology. 242 (2): 343–64. doi:10.1111/j.1469-7998.1997.tb05806.x.
- Odden, M.; Wegge, P. (2005). "Spacing and activity patterns of leopards Panthera pardus in the Royal Bardia National Park, Nepal" (PDF). Wildlife Biology. 11 (2): 145–52. doi:10.2981/0909-6396(2005)11[145:SAAPOL]2.0.CO;2.
- Hayward, M.W.; Henschel, P.; O'Brien, J.; Hofmeyr, M.; Balme, G.; Kerley, G.I.H. (2006). "Prey preferences of the leopard (Panthera pardus)" (PDF). Journal of Zoology. 270: 298–313. doi:10.1111/j.1469-7998.2006.00139.x.
- Scheepers, J.L.; Gilchrist, D. (1991). "Leopard predation on giraffe calves in the Etosha National Park" (PDF). Madoqua. 18 (1): 49.
- Sunquist, M.E.; Sunquist, F. (2002). Wild Cats of the World. Chicago: University of Chicago Press. p. 325. ISBN 978-0-226-77999-7.
- Zuberbühler, K.; Jenny, D. (2002). "Leopard predation and primate evolution". Journal of Human Evolution. 43 (6): 873–86. PMID 12473487. doi:10.1006/jhev.2002.0605.
- Arivazhagan, C.; Arumugam, R.; Thiyagesan, K. (2005). "Food habits of leopard (Panthera pardus fusca), dhole (Cuon alpinus) and striped hyena (Hyaena hyaena) in a tropical dry thorn forest of southern India" (PDF). Journal of the Bombay National Historical Society. 104 (2). Archived from the original (PDF) on 4 March 2009.
- Johnson K. G.; Wei W.; Reid D. G.; Jinchu H. (1993). "Food habits of Asiatic leopards (Panthera pardus fusca) in Wolong Reserve, Sichuan, China". Journal of Mammalogy. 74 (3): 646–50. JSTOR 1382285. doi:10.2307/1382285.
- Bailey, T.N. (1993). The African leopard: a study of the ecology and behaviour of a solitary felid. New York, US: Columbia University Press. ISBN 978-1-932846-11-9.
- Bothma, J. du P. (2005). "Water-use by southern Kalahari leopards" (PDF). South African Journal of Wildlife Research. 35: 131–7.
- Mills, M.G.L.; Hes, L. (1997). The Complete Book of Southern African Mammals. Cape Town, South Africa: Struik Publishers. pp. 178–80. ISBN 978-0-947430-55-9.
- Hamilton, P.H. (1976). The movements of leopards in Tsavo National Park, Kenya, as determined by radio-tracking (M.Sc. thesis). Nairobi: University of Nairobi.
- Schaller, G. (1972). Serengeti: a kingdom of predators. New York: Knopf. ISBN 978-0-394-47242-3.
- Boitani, L. (1984). Simon & Schuster's Guide to Mammals. Simon & Schuster/Touchstone Books. ISBN 978-0-671-42805-1.
- "Cat fight: leopard vs caracal", Africa Geographic, 8 July 2016
- Seidensticker, J (1976). "On the ecological separation between tigers and leopards" (PDF). Biotropica. 8 (4): 225. JSTOR 2989714. doi:10.2307/2989714.
- Harihar, Abishek; Pandav, Bivash; Goyal, Surendra P (2011). "Responses of leopard Panthera pardus to the recovery of a tiger Panthera tigris population". Journal of Applied Ecology. 48 (3): 806–814. doi:10.1111/j.1365-2664.2011.01981.x.
- Hepnter, V.G.; and Sluskii, A.A. (1992) Mammals of the Soviet Union. Vol 2, Part 2. (Carnivores: Hyaenas and Cats) New Delhi: Amerind Publishing. pp. 266–7.
- Seidensticker, J (1976). "On the ecological separation between tigers and leopards" (PDF). Biotropica. 8 (4): 225–234. JSTOR 2989714. doi:10.2307/2989714.
- Seidensticker, J (1976). "On the ecological separation between tigers and leopards" (PDF). Biotropica. 8: 229–230.
- Seidensticker, J (1976). "On the ecological separation between tigers and leopards" (PDF). Biotropica. 8: 232.
- Karanth, K. U.; Sunquist, M. E. (1995). "Prey selection by tiger, leopard and dhole in tropical forests". Journal of Animal Ecology. 64 (4): 439–450. JSTOR 5647. doi:10.2307/5647.
- Karanth, U. K.; Sunquist, M. E. (2000). "Behavioural correlates of predation by tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India". Journal of Zoology. 250 (2): 255–265. doi:10.1111/j.1469-7998.2000.tb01076.x.
- Leopard left for dead by baboon troop. wilderness-safaris.com (2006-10-25).
- "Nile Crocodile". Crocodilian Species List.
- Bailey, T. N. (1993). The African leopard: ecology and behavior of a solitary felid. Columbia University Press.
- Kingdon, J., Happold, D., Butynski, T., Hoffmann, M., Happold, M., & Kalina, J. (2013). Mammals of Africa (Vol. 1). A&C Black.
- Bhatnagar, C., & Mahur, M. (2010). "Observations on feeding behavior of a wild population of marsh crocodile in Baghdarrah Lake, Udaipur, Rajasthan". Reptile Rap. 10: 16–18.
- Owens, M.; Owens, D. (1984). Cry of the Kalahari. Boston: Houghton Mifflin. ISBN 978-0-395-32214-7.
- Owens, D.; Owens, M. (February 1980). "Hyenas of the Kalahari". Natural History. 89 (2): 50.
- Gower, David; Garrett, Katherine; Stafford, Peter (2012). Snakes. Firefly Books. p. 60. ISBN 978-1-55407-802-8.
- Perowne, J. (2014). Blog: Leopard kills cheetah in the Mara and hoists it up a tree. The Safari Collection, Nairobi.
- Hagen, M. (2014). Leopard kills Cheetah. youtube.com
- Sadleir, R. (1966). "Notes on the Reproduction of the larger Felidae". International Zoo Yearbook. 6: 184–87. doi:10.1111/j.1748-1090.1966.tb01746.x.
- Hemmer, H. (1976). "Gestation period and postnatal development in felids". In Eaton, R.L. The world's cats. 3. Carnivore Research Institute, Univ. Washington, Seattle. pp. 143–165.
- Eaton, R.L. (1977). "Reproductive biology of the leopard". Zoologischer Garten. 47 (5): 329–351.
- "Leopard: The Spotted Cat". animals.about.com.
- Salisbury, Susan (2014-08-09) Roxanne, oldest spotted leopard in captivity, dies at Acreage preserve. The Palm Beach Post
- "Nie ma już lampartów cejlońskich w warszawskim zoo ...". 15 December 2010.
- "Co w zoo? – wtorkowy raport aktualności". 30 March 2010.
- "Benin: an African kingdom" (PDF). British Museum. Retrieved 2016-03-29.
- Strickland, Debra Higgs; Debra Hassig (1999). The Mark of the Beast: The Medieval Bestiary in Art, Life, and Literature. Taylor & Francis. ISBN 978-0-8153-2952-7.
- Pedersen, Christian Fagd (1971). The International Flag Book in Color. Morrow.
- Owen, James (November 3, 2005). "Medieval Lion Skulls Reveal Secrets of Tower of London 'Zoo'". National Geographic Magazine. National Geographic. Retrieved 2007-09-05.
- Wines, M. (2004). "In South Africa, It's All in the Game". The New York Times. Retrieved 6 February 2015.
- Weerasinghe, U.M.I.R.K., Kariyawasm, D. and M. De Zoysa (2003). Ruhuna (Yala) National Park in Sri Lanka: Visitors, Visitation, and Eco-Tourism. Contribution to the XII World Forestry Congress, Quebec.
- Kunzru, Hari (8 May 2015). "Life Among the Leopards" – via NYTimes.com.
- Capstick, Peter Hathaway (1978). Death in the Long Grass. St. Martin's Press. ISBN 978-0-312-18613-5.
- Anderson, Kenneth (1954). Nine Man-Eaters and one Rogue. Allen & Unwin.
- Allsen, Thomas T. (2007). "Natural History and Cultural History: The Circulation of Hunting Leopards in Eurasia, Seventh-Seventeenth Centuries". In Mair, Victor H. Contact and Exchange in the Ancient World. Honolulu: University of Hawai'i Press. pp. 116–135. ISBN 978-0-8248-2884-4.
- Khalaf-von Jaffa; Norman Ali Bassam Ali Taher (June 2005). "The Arabian Leopard (Panthera pardus nimr)". Gazelle: the Palestinian Biological Bulletin (in German) (42): 1–8.
- Khalaf-Sakerfalke von Jaffa; Norman Ali Bassam Ali Taher (December 2006). "The Chinese leopard (Panthera pardus japonensis, Gray 1862) in Neunkirchen Zoo, Neunkirchen, Saarland, Germany". Gazelle: the Palestinian Biological Bulletin (60): 1–10.
- Nowell, K; Jackson, P., eds. (1996). "Wild Cats. Status Survey and Conservation Action Plan" (PDF). Gland, Switzerland: IUCN/SSC Cat Specialist Group.
- Schaller, George B. (1972). The Serengeti Lion. Chicago: University of Chicago Press. ISBN 978-0-226-73639-6.
- DeRuiter, D. J.; Berger, L. R. (2000). "Leopards as Taphonomic Agents in dolomitic Caves—Implications for bone Accumulations in the Hominid-bearing Deposits of South Africa". Journal of Archaeological Science. 27 (8): 665–684. doi:10.1006/jasc.1999.0470.
- Sanei, A. (2007). Analysis of leopard (Panthera pardus) status in Iran (in Persian). Tehran: Sepehr Publication Center. ISBN 978-964-6123-74-8.
- Sanei, A.; Zakaria, M.; Yusof, E.; Roslan, M. (2011). "Estimation of leopard population size in a secondary forest within Malaysia's capital agglomeration using unsupervised classification of pugmarks" (PDF). Tropical Ecology. 52 (1): 209–217.
- P., Taylor; Barrientos, Stephanie; Dolan, Catherine (2005). Beyond Conservation: A Wildland Strategy. Earthscan. ISBN 978-1-84407-197-5.
- Zakaria, M.; Sanei, A. (2011). "Conservation and management prospects of the Persian and Malayan leopards". Asia Life Sciences. Supplement 7: 1–5.
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