The leopard (), ''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. The leopard was once distributed across eastern and southern Asia and Africa, from Siberia to South Africa, but its 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 Indonesia, 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.
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 , its unequaled ability to climb trees even when carrying a heavy carcass, and its notorious ability for stealth. The leopard consumes virtually any animal that it can hunt down and catch. Its habitat ranges from rainforest to desert terrains.
Physical characteristics
Leopards are agile and stealthy predators. Although smaller than other members of the ''Panthera'' genus, they are able to take large prey due to their massive skulls that facilitate powerful jaw muscles. Head and body length is between , and the tail reaches . Shoulder height is . The muscles attached to the scapula are exceptionally strong, which enhance their ability to climb trees. They are very diverse in size. Males are about 30% larger than females, weighing compared to for females. Large males of up to have been documented in Kruger National Park in South Africa; however, males in the South Africa's coastal mountains average . 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. The largest verified leopard weighed , with larger sizes reported but considered unreliable. The leopard's body is comparatively long, and its legs are short.
Leopards show a great diversity in coat color and rosette patterns. Their rosettes are circular in East Africa but tend to be squarer in southern Africa and larger in Asian populations. Their yellow coat tends to 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.
Leopards may sometimes be confused with two other large spotted cats, the cheetah, with which it may co-exist in Africa, and the jaguar, a neotropical species that it does not naturally co-exist with. However, the patterns of spots in each are different: the cheetah has simple black spots, evenly spread; the jaguar has small spots inside the polygonal rosettes; while the leopard normally has rounder, smaller rosettes than those of the jaguar. The cheetah has longer legs and a thinner build that makes it look more streamlined and taller but less powerfully built than the leopard. The jaguar is more similar in build to the leopard but is generally larger in size and has a more muscular, bulky appearance.
Variant coloration
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. (They are commonly called
black panthers, although 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 that up to half of all leopards are black, but a 2007 camera-trap study in Taman Negara National Park found that all specimens were melanistic. The benefits of melanism are difficult to interpret, but it may serve as camouflage in the rainforest habitat. It is also possible that the color variation is a relic adaptation to an epidemic; genes causing melanism can also affect the immune system. Genetic research has found four independent origins for melanism in cats, suggesting that there may be an adaptive advantage.
In Africa, black leopards are much less common, as melanism is not an adaptive advantage in 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.
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 color. 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.
Etymology
In
antiquity, a leopard was believed to be a
hybrid of a lion and a
panther, as is reflected in its name, which is a Greek
compound of ''leōn'' (lion) and ''pardos'' (male panther). The Greek word is related to
Sanskrit (snake, tiger, panther), and probably is derived from a Mediterranean language, such as
Egyptian.
A ''panther'' can be any of several species of large felids: the term can refer to cougars and jaguars in the American continents; and everywhere else, to leopards.
The generic component of its modern scientific designation, ''Panthera pardus'', is derived from Latin via Greek πάνθηρ (''pánthēr''). Folk etymology held that it was a compound of παν (''pan'', all) and θηρ (beast). However, it is believed instead to be derived from an Indo-Iranian word meaning "white-yellow, pale"; in Sanskrit, this word's reflex was पाण्डर ''pāṇḍara'', which was derived from पुण्डरीक ''puṇḍárīka'' (''tiger'', among other things), then borrowed into Greek.
Taxonomy and evolution
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.
Carl Linnaeus placed leopards under the genus ''Felis'' as the binominal ''Felis pardus''. In the 18th and 19th centuries, most naturalists and taxonomists followed his example. In 1816, Lorenz Oken proposed a definition of the genus ''Panthera'', with a subgenus Panthera using Linnaeus’ ''Felis pardus'' as a type specimen. But most disagreed with his definition, and until the beginning of the 20th century continued using ''Felis'' or ''Leopardus'' when describing leopard subspecies. In 1916, Reginald Innes Pocock accorded ''Panthera'' generic rank defining ''Panthera pardus'' as species.
It is believed that the basal divergence amongst the Felidae family occurred about 11 million years ago. The last common ancestor of the lion, tiger, leopard, jaguar, snow leopard, and clouded leopard is believed to have occurred about 6.37 million years ago. ''Panthera'' is believed to have emerged in Asia, with ancestors of the leopard and other cats subsequently migrating into Africa. The researchers suggest that the snow leopard is most closely aligned with the tiger, whereas the leopard possibly has diverged from the ''Panthera'' lineage subsequent to these two species, but before the lion and jaguar. 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.
Results of phylogenetic analyses of chemical secretions amongst cats has suggested that the leopard is closely related to the lion.
Results of a mitochondrial DNA study carried out later suggest that the leopard is closely related to the snow leopard, which is placed as a fifth ''Panthera'' species, ''Panthera uncia''.
Distribution and habitat
Leopards have the largest distribution of any wild cat, occurring widely in eastern and central
Africa, 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. But 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; but 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 live mainly in grasslands, woodlands, and riverine forests. They are usually associated with savanna and rainforest, but leopards are exceptionally adaptable: in the Russian Far East, they inhabit temperate forests where winter temperatures reach a low of .
Distribution of subspecies
Since
Carl Linnaeus published his description of leopards in the
10th edition of Systema Naturae in 1758, as many as 27 leopard subspecies were subsequently described by naturalists from 1794 to 1956. In 1996, according to DNA analysis carried out in the 1990s, only eight subspecies are considered valid. Later analysis revealed a ninth valid subspecies, the
Arabian leopard (''P. p. nimr''). Because of limited sampling of
African leopards, this number might be an underestimation.
The nine subspecies recognised by IUCN are:
African leopard (''P. p. pardus''), (Linnaeus, 1758) — inhabits sub-Saharan Africa;
Indian leopard (''P. p. fusca''), (Meyer, 1794) — inhabits the Indian Subcontinent;
Javan leopard (''P. p. melas''), (Cuvier, 1809) — inhabits Java, Indonesia.
Arabian leopard (''P. p. nimr''), (Hemprich and Ehrenberg, 1833) — inhabits the Arabian Peninsula;
Amur leopard (''P. p. orientalis''), (Schlegel, 1857) — inhabits the Russian Far East, Korean Peninsula and Northeast China;
North Chinese leopard (''P. p. japonensis''), (Gray, 1862) — inhabits northern China;
Caucasian leopard (''P. p. ciscaucasica''), (
Satunin, 1914), later described as
Persian leopard (''P. p. saxicolor''), (
Pocock, 1927) — inhabits central Asia: the
Caucasus,
Turkmenistan and northern
Iran;
Indo-Chinese leopard (''P. p. delacouri''), (Pocock, 1930) — inhabits mainland Southeast Asia;
Sri Lankan leopard (''P. p. kotiya''), (Deraniyagala, 1956) — inhabits Sri Lanka.
A morphological analysis of characters of leopard skulls implies the validity of two more subspecies:
Anatolian leopard (''P. p. tulliana''), (Valenciennes, 1856) — inhabits Western Turkey;
Balochistan leopard (''P. p. sindica''), (Pocock, 1930) — inhabits Pakistan, and possibly also parts of Afghanistan and Iran.
Ecology and behavior
Leopards are elusive, solitary and largely
nocturnal. They have primarily been studied in open savanna habitats, which may have biased common descriptions. Activity level varies depending on the habitat and the type of prey that they hunt. Radio-tracking and scat analysis in West Africa showed that rainforest leopards are more likely to be
diurnal and
crepuscular. Forest leopards are also more specialized in prey selection and exhibit seasonal differences in activity patterns.
Leopards are known for their ability in climbing, and have been observed resting on tree branches during the day, dragging their kills up trees and hanging them there, and descending from trees headfirst. They are powerful swimmers, although not as strong as some other big cats, such as the tiger. They are very agile, and can run at over , leap over horizontally, and jump up to vertically. They produce a number of vocalizations, including grunts, roars, growls, meows, and "sawing" sounds.
Social structure and home range
Home ranges of male leopards vary between and , and of females between . Virtually all sources suggest that males do have larger home ranges. There seems to be little or no overlap in territory among males, although overlap exists between the sexes; one radio-collar analysis in the
Ivory Coast found a female home range completely enclosed within a male's.
Research in a conservation area in Kenya showed similar territory sizes and sex differential: average ranges for males, and for females.
In Nepal, somewhat larger male ranges have been found at about , while female ranges at ; female home ranges decreased to when young cubs were present, while the sexual difference in range size seemed to be in positive proportion to overall increase.
Studies of leopard home range size have tended to focus on protected areas, which may have led to skewed data; as of the mid-1980s, only 13% of the leopard range actually fell within a protected area. However, significant variations in the size of home ranges have been suggested across the leopard's range. Research in Namibia that focused on spatial ecology in farmlands outside of protected areas revealed ranges that were consistently above with some more than . Admitting that their data were at odds with others, the researchers found little or no sexual variation in the size of territories.
Hunting and diet
Leopards are versatile, opportunistic hunters, and have a very broad diet. They feed on a greater diversity of prey than other members of the ''Panthera'' species, and will eat anything from
dung beetles to male
giant elands. Their diet consists mostly of
ungulates and
monkeys, but they also eat
rodents,
reptiles,
amphibians, insects, birds (like the
Vulturine Guineafowl), fish and sometimes smaller predators (such as
bat-eared foxes,
martens, and
jackals). In at least one instance, a leopard has predated a sub-adult
nile crocodile that was crossing over land. They stalk their prey silently, pounce on it at the last minute, and strangle its throat with a quick bite. In Africa, mid-sized
antelopes provide a majority of their prey, especially
impala and
Thomson's gazelles.
In the open savanna of Tsavo National Park, they kill more prey when hunting between sunset and sunrise. In Kruger National Park, males and females with cubs are more active at night. At least 92 prey species have been documented in their diet. They focus their hunting activity on locally abundant medium-sized ungulate species in the range, while opportunistically taking other prey. Analysis of leopard scats found that 67% contained ungulate remains, of which 60% were impala, the most abundant antelope, with adult weights of . Small mammal remains were found most often in scats of sub-adult leopards, especially females. Average daily consumption rates was estimated at for adult males and for females.
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 favored prey of the leopard were chitals. A study at the Wolong Reserve in China revealed how adaptable their hunting behaviour is. Over the course of seven years, the vegetative cover receded, and the animals opportunistically shifted from primarily consuming tufted deer to pursuing bamboo rats and other smaller prey.
They select their prey focusing on small herds, dense habitat, and low risk of injury, preferring prey weights of such as impala, chital, bushbuck and common duiker with an average body weight of .
In search of safety, leopards often stash their young or recent kills high up in a tree. There were observed hauling carcasses of young giraffe, estimated to weigh up to , 2–3 times the weight of the leopard, up to into trees.
Interspecific predatory relationships
Leopards must compete for food and shelter with other large predators such as
lions,
tigers,
spotted hyenas, and both
African and
Asiatic wild dogs. These animals may steal the leopard's kill or devour its young, though lions are most likely to kill and not eat the young if they're discovered. In some areas of Africa, troops of large
baboon species (potentially leopard prey themselves) will kill and sometimes eat leopard young if they discover them. Occasionally,
nile crocodiles may predate on leopards. Leopards co-exist alongside these other predators by hunting for different types of prey and by avoiding areas frequented by them. Lions are occasionally successful in climbing trees and fetching leopard kills. 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.
Resource partitioning occurs where leopards share their range with lions or tigers. Leopards tend to take smaller prey, usually less than , where the larger cats are present. In the Chitwan National Park in Nepal, leopards killed prey ranging from less than to in weight with most kills in the range; tigers killed more prey in the range. In the tropical forests of India’s Nagarhole National Park, tigers selected prey weighing more than , whereas leopards selected prey in the range. The average weights of leopard prey was , and of tiger prey was with a bias towards adult males of chital, sambar and wild pig, and young gaur. In tropical forest they 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. 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.
Aggressive encounters have been observed. Two of five males studied over a period of a year at a game reserve in South Africa died, both violently. One was initially wounded in a male–male territorial battle over a carcass; taken in by researchers, it was released after a successful convalescence only to be killed by a different male a few months later. A second was killed by another predator, possibly a spotted hyena. A third of the five was badly wounded in intraspecific fighting, but recovered.
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. But 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 color 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.
Leopards have been reported to reach 21 years of age in captivity.
Hybrids
Crossbreeding between leopards and other members of the genus ''Panthera'' has been documented, resulting in
hybrids. A cross between a lioness and a male leopard is known as a leopon (or a lipard if the sex of the parents is reversed). Leopons have been bred in captivity; a well-documented case occurred at the Koshien Hanshin Park in
Nishinomiya, Japan in the late 1950s. 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. 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 jaguars 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.
Whether born to a female puma mated to a male leopard, or to a male puma mated to a female leopard, pumapards inherit a form of dwarfism. Those reported grew to only half the size of the parents. They have a puma-like long body (proportional to the limbs, but nevertheless shorter than either parent), but short legs. The coat is variously described as sandy, tawny or greyish with brown, chestnut or faded rosettes.
Leopards and humans
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.
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.
Heraldry
The lion passant guardant or leopard is a frequently used charge in heraldry, most commonly appearing in groups of three. The heraldric leopard lacks spots and sports a mane, making it visually almost identical to the heraldric lion, and the two are often used interchangeably. These traditional 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.
Tourism
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. 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.
Man-eating
Most leopards avoid people, but humans may occasionally be targeted as prey. Most healthy leopards prefer wild prey to humans, but injured, sickly, or struggling cats 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 more than 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. Both were killed by hunter
Jim Corbett. Man-eating leopards are considered bold 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:
See also
List of solitary animals
References
Further reading
Nowell, K.; Jackson, P. eds. (1996). ''Wild Cats. Status Survey and Conservation Action Plan.'' IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland.
Asian Leopard Specialists' Newsletter
External links
The Animal Files: Leopard
Leopards: wild-cat.org — Anthology about research and conservation of leopards in Asia
Asian Leopard Specialist Society — Research, conservation and management of Asian leopard subspecies
Category:Leopards
Category:Mammals of Africa
Category:Mammals of Asia
Category:Megafauna of Africa
Category:Megafauna of Eurasia
Category:National symbols of Benin
Category:National symbols of the Democratic Republic of the Congo
Category:National symbols of Malawi
Category:National symbols of Somalia
Category:Big cats of India
Category:Animals described in 1758
af:Luiperd
am:ግሥላ
ang:Lafor
ar:نمر
az:Bəbir
bn:চিতাবাঘ
zh-min-nan:Hoe-pà
be:Леапард
be-x-old:Леапард
bo:གཟིག
bg:Леопард
ca:Lleopard
cs:Levhart skvrnitý
tum:Nyalubwe
co:Leupardu
da:Leopard
de:Leopard
nv:Łizhiní Bikéyahdę́ę́ʼ náshdóítsoh łikizhígíí
et:Leopard
el:Λεοπάρδαλη
es:Panthera pardus
eo:Leopardo
eu:Lehoinabar
fa:پلنگ
fo:Leopardur
fr:Léopard (félin)
ff:Cirgu
ga:Liopard
gag:Leopard
gd:Leopard
gu:દિપડો
hak:Pau-é
ko:표범
ha:Damisa
hy:Ընձառյուծ
hi:तेन्दुआ
hr:Leopard
io:Leopardo
ig:Ágwu
id:Macan tutul
ia:Leopardo
is:Hlébarði
it:Panthera pardus
he:נמר
ka:ჯიქი
kk:Леопард
sw:Chui
ht:Lewopa
ku:Pars(Leopar)
lbe:Паланг
ltg:Leopards
la:Pardus
lv:Leopards
lt:Leopardas
ln:Nkɔi
hu:Leopárd
mk:Леопард
ml:പുള്ളിപ്പുലി
mr:बिबट्या
arz:فهد
ms:Harimau Bintang
mn:Цоохор ирвэс
my:ကျားသစ်
nl:Luipaard
ja:ヒョウ
no:Leopard
nn:Leopard
oc:Panthera pardus
pa:ਲੈਪਰਡ
pnb:تیندوہ
pl:Lampart
pt:Leopardo
kbd:Щомыщ
ro:Leopard
qu:Misti hawar
ru:Леопард
sg:Zé
st:Nkwe
sq:Leopardi
scn:Liupardu
si:කොටියා
simple:Leopard
sk:Leopard škvrnitý
cu:Лєопардъ
sl:Leopard
sr:Леопард
sh:Leopard
su:Maung tutul
fi:Leopardi
sv:Leopard
tl:Leopardo
th:เสือดาว
chr:ᏢᏓᏥ ᎤᏅᏣᏗ
ve:Nngwe
tr:Pars
tk:Alajagaplaň
uk:Пантера плямиста
ur:تیندوا
ug:قاپلان
vec:Lionpardo
vi:Báo hoa mai
zh-yue:豹
zh:豹