Present and historical distribution of the leopard[2]
The leopard (Panthera pardus) is one of the five extant
species in the
genusPanthera. It has a pale yellowish to dark golden fur with dark spots grouped in
rosettes. Its body is slender and muscular reaching a length of 92–183 cm (36–72 in) with a 66–102 cm (26–40 in) long tail and a shoulder height of 60–70 cm (24–28 in). Males typically weigh 30.9–72 kg (68–159 lb), and females 20.5–43 kg (45–95 lb).
The leopard was first
described in 1758, and several
subspecies were proposed in the 19th and 20th centuries. Today, eight subspecies are recognised in its wide range in
Africa and
Asia. It initially evolved in Africa during the
Early Pleistocene, before migrating into Eurasia around the Early–
Middle Pleistocene transition. Leopards were formerly present across Europe, but became extinct in the region at around the end of the
Late Pleistocene-early
Holocene.
The leopard is adapted to a variety of habitats ranging from
rainforest to
steppe, including
arid and montane areas. It is an
opportunistic predator, hunting mostly
ungulates and
primates. It relies on its spotted pattern for
camouflage as it stalks and ambushes its prey, which it sometimes drags up a tree. It is a
solitary animal outside the mating season and when raising cubs. Females usually give birth to a litter of 2–4 cubs once in 15–24 months. Both male and female leopards typically reach
sexual maturity at the age 2–2.5 years.
Listed as
Vulnerable on the
IUCN Red List, leopard populations are currently threatened by
habitat loss and fragmentation, and are declining in large parts of the global range. Leopards have had cultural roles in
Ancient Greece,
West Africa and modern Western culture. Leopard skins are popular in fashion.
Etymology
The English name "leopard" comes from
Old Frenchleupart or
Middle Frenchliepart, that derives from
Latinleopardus and
ancient Greekλέοπάρδος (leopardos). Leopardos could be a
compound of λέων (leōn), meaning 'lion', and πάρδος (pardos), meaning 'spotted'.[3][4][5] The word λέοπάρδος originally referred to a
cheetah (Acinonyx jubatus).[6]
"Panther" is another common name, derived from Latin panther and ancient Greek πάνθηρ (pánthēr);[3] The
generic namePanthera originates in Latin panthera, a hunting net for catching wild beasts to be used by the
Romans in combats.[7]Pardus is the
masculine singular form.[8]
Following Linnaeus' first description, 27 leopard
subspecies were proposed by naturalists between 1794 and 1956. Since 1996, only eight subspecies have been considered
valid on the basis of
mitochondrial analysis.[15] Later analysis revealed a ninth valid subspecies, the
Arabian leopard.[16]
In 2017, the Cat Classification Task Force of the Cat Specialist Group recognized the following eight subspecies as valid taxa:[17]
The Balochistan leopard population in the south of Iran, Afghanistan and Pakistan is separated from the northern population by the
Dasht-e Kavir and
Dasht-e Lut deserts.[28]
Results of an
analysis of molecular variance and pairwise
fixation index of 182 African leopard museum specimens showed that some African leopards exhibit higher genetic differences than Asian leopard subspecies.[35]
Evolution
Results of
phylogenetic studies based on
nuclear DNA and
mitochondrial DNA analysis showed that the last
common ancestor of the Panthera and Neofelis genera is thought to have lived about 6.37 million years ago. Neofelisdiverged about 8.66 million years ago from the Pantheralineage. The tiger diverged about 6.55 million years ago, followed by the snow leopard about 4.63 million years ago and the leopard about 4.35 million years ago. The leopard is a
sister taxon to a
clade within Panthera, consisting of the lion and the jaguar.[36][37]
Results of a phylogenetic analysis of chemical
secretions amongst cats indicated that the leopard is closely related to the lion.[40]
The geographic origin of the Panthera is most likely northern Central Asia. The leopard-lion clade was distributed in the Asian and African
Palearctic since at least the early
Pliocene.[41] The leopard-lion clade diverged 3.1–1.95 million years ago.[38][39] Additionally, a 2016 study revealed that the
mitochondrial genomes of the leopard, lion and
snow leopard are more similar to each other than their
nuclear genomes, indicating that their ancestors
hybridized with the snow leopard at some point in their
evolution.[42]
The oldest unambiguous
fossils of the leopard are from Eastern Africa, dating to around 2 million years ago.[43]
Leopard-like fossil bones and teeth possibly dating to the
Pliocene were excavated in
Perrier in
France, northeast of
London, and in
Valdarno,
Italy. Until 1940, similar fossils dating back to the
Pleistocene were excavated mostly in
loess and caves at 40 sites in Europe, including
Furninha Cave near
Lisbon,
Genista Caves in
Gibraltar, and
Santander Province in northern
Spain to several sites across 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.[44]
Leopards arrived in Eurasia during the late Early to Middle Pleistocene around 1.2[45] to 0.6 million years ago.[43]
Four European Pleistocene leopard subspecies were proposed. P. p. begoueni from the beginning of the
Early Pleistocene 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.[46]P. p. spelaea is the most recent subspecies that appeared at the beginning of the
Late Pleistocene and survived until about 11,000 years ago and possibly into the early
Holocene in the
Iberian Peninsula.[46][47]
Leopards depicted in cave paintings in
Chauvet Cave provide indirect evidence of leopard presence in Europe.[46]
Leopard fossils dating to the Late Pleistocene were found in Biśnik Cave in south-central
Poland.[48][45]
Fossil remains were also excavated in the Iberian[49][50] and
Italian Peninsula,[51] and in the
Balkans.[52][53]
Leopard fossils dating to the Pleistocene were also excavated in the
Japanese archipelago.[54] Leopard fossils were also found in
Taiwan.[55]
In 1953, a male leopard and a female lion were
crossbred in Hanshin Park in
Nishinomiya, Japan. Their offspring known as a
leopon was born in 1959 and 1961, all cubs were spotted and bigger than a juvenile leopard. Attempts to mate a leopon with a tigress proved unsuccessful.[56]
Characteristics
Mounted skeleton
Rosettes of a leopard
A melanistic leopard or black panther
The leopard's fur is generally soft and thick, notably softer on the belly than on the back.[57] Its skin colour varies between individuals from pale yellowish to dark golden with dark spots grouped in
rosettes. Its underbelly is white and its ringed tail is shorter than its body. Its pupils are round.[58] Leopards living in arid regions are pale cream, yellowish to
ochraceous and
rufous in colour; those living in forests and mountains are much darker and deep golden. Spots fade toward the white underbelly and the insides and lower parts of the legs.[59] Rosettes are circular in East African leopard populations, and tend to be squarish in Southern African and larger in Asian leopard populations. The fur tends to be grayish in colder climates, and dark golden in
rainforest habitats.[60] Rosette patterns are unique in each individual.[61][62] This pattern is thought to be an adaptation to dense vegetation with patchy shadows, where it serves as
camouflage.[63]
Its white-tipped tail is about 60–100 cm (23.6–39.4 in) long, white underneath and with spots that form incomplete bands toward the end of the tail.[64]
The guard hairs protecting the basal hairs are short, 3–4 mm (0.1–0.2 in) in face and head, and increase in length toward the flanks and the belly to about 25–30 mm (1.0–1.2 in). Juveniles have woolly fur that appear to be dark-coloured due to the densely arranged spots.[61][65]
Its fur tends to grow longer in colder climates.[66]
The leopard's rosettes differ from those of the
jaguar, which are darker and with smaller spots inside.[58] The leopard has a
diploidchromosome number of 38.[67]
The leopard is a slender and muscular cat, with relatively short limbs and a broad head. It is
sexually dimorphic with males larger and heavier than females.[64] 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 ranges between 92 and 183 cm (36 and 72 in) with a 66 to 102 cm (26 to 40 in) long tail. Sizes vary geographically. Males typically weigh 30.9–72 kg (68–159 lb), and females 20.5–43 kg (45–95 lb).[74] Occasionally, large males can grow up to 91 kg (201 lb). Leopards from the
Cape Province in South Africa are generally smaller, reaching only 20–45 kg (44–99 lb) in males.[65][66][75]
The maximum recorded weight of a wild leopard in Southern Africa was around 96 kg (212 lb), and it measured 262 cm (103 in).[76] In 2016, an Indian leopard killed in
Himachal Pradesh measured 261 cm (103 in) with an estimated weight of 78.5 kg (173 lb); it was perhaps the largest known wild leopard in India.[77][78]
The largest recorded skull of a leopard was found in India in 1920 and measured 28 cm (11 in) in
basal length, 20 cm (7.9 in) in breadth, and weighed 1 kg (2.2 lb). The skull of an African leopard measured 286 mm (11.3 in) in basal length, and 181 mm (7.1 in) in breadth, and weighed 790 g (28 oz).[79]
The leopard has the largest distribution of all wild cats, occurring widely in Africa, the
Caucasus and Asia, although populations are fragmented and declining. It is considered to be
locally extinct in
North Africa.[2] It inhabits foremost
savanna and
rainforest, and areas where
grasslands,
woodlands, and riverine forests remain largely undisturbed.[60] In
sub-Saharan Africa, it is still numerous and surviving in marginal habitats where other large cats have disappeared. There is considerable potential for human-leopard conflict due to leopards preying on livestock.[80]
Leopard populations in the
Arabian Peninsula are small and fragmented.[81][82][83] In southeastern
Egypt, a leopard killed in 2017 was the first sighting of the species in this area in 65 years.[84]
In the
Indian subcontinent, the leopard is still relatively abundant, with greater numbers than those of other Panthera species.[2] As of 2020, the leopard population within forested habitats in
India's tiger range landscapes was estimated at 12,172 to 13,535 individuals. Surveyed landscapes included elevations below 2,600 m (8,500 ft) in the
Shivalik Hills and
Gangetic plains,
Central India and
Eastern Ghats,
Western Ghats, the
Brahmaputra River basin and hills in
Northeast India.[85] Some leopard populations in the country live quite close to human settlements and even in semi-developed areas. Although 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 stealth, people often remain unaware that it lives in nearby areas.[86]
The leopard is a solitary and
territorial animal. It is typically shy and alert when crossing roadways and encountering oncoming vehicles, but may be emboldened to attack people or other animals when threatened. Adults associate only in the mating season. Females continue to interact with their offspring even after weaning and have been observed sharing kills with their offspring when they can not obtain any prey. They produce a number of vocalizations, including growls, snarls, meows, and purrs.[65] The
roaring sequence in leopards consists mainly of grunts,[98] also called "sawing", as it resembles the sound of sawing wood. Cubs call their mother with an urr-urr sound.[65]
The whitish spots on the back of its ears are thought to play a role in communication.[99]
It has been hypothesized that the white tips of their tails may function as a 'follow-me' signal in
intraspecific communication. However, no significant association were found between a conspicuous colour of tail patches and behavioural variables in carnivores.[100][101]
Leopards are mainly active from dusk till dawn and will rest for most of the day and some hours at night in thickets, among rocks or over tree branches. Leopards have been observed walking 1–25 km (0.62–15.53 mi) across their range at night; wandering up to 75 km (47 mi) if disturbed.[65][75] In some regions, they are
nocturnal.[102][103] In western African forests, they have been observed to be largely
diurnal and hunting during twilight, when their prey animals are active; activity patterns vary between seasons.[104]
Leopards can climb trees quite skillfully, often resting on tree branches and descending headfirst.[60]
They can run at over 58 km/h (36 mph; 16 m/s), leap over 6 m (20 ft) horizontally, and jump up to 3 m (9.8 ft) vertically.[98]
A female leopard with white markings on the backs of her ears.
In
Kruger National Park, most leopards tend to keep 1 km (0.62 mi) apart.[106] Males occasionally interact with their partners and cubs, and exceptionally this can extend beyond to two generations.[74][107] Aggressive encounters are rare, typically limited to defending territories from intruders.[66] In a South African reserve, a male was wounded in a male–male territorial battle over a carcass.[102]
Males occupy
home ranges that often overlap with a few smaller female home ranges, probably as a strategy to enhance access to females. In the
Ivory Coast, the home range of a female was completely enclosed within a male's.[108] Females live with their cubs in home ranges that overlap extensively, probably due to the association between mothers and their offspring. There may be a few other fluctuating home ranges belonging to young individuals. It is not clear if male home ranges overlap as much as those of females do. Individuals try to drive away intruders of the same sex.[65][75]
A study of leopards in the Namibian farmlands showed that the size of home ranges was not significantly affected by sex, rainfall patterns or season; the higher the prey availability in an area, the greater the leopard population density and the smaller the size of home ranges, but they tend to expand if there is human interference.[109]
Sizes of home ranges vary geographically and depending on habitat and availability of prey. In the
Serengeti, males have home ranges of 33–38 km2 (13–15 sq mi) and females of 14–16 km2 (5.4–6.2 sq mi);[110][111] but males in northeastern Namibia of 451 km2 (174 sq mi) and females of 188 km2 (73 sq mi).[112] They are even larger in arid and montane areas.[66] In Nepal's
Bardia National Park, male home ranges of 48 km2 (19 sq mi) and female ones of 5–7 km2 (1.9–2.7 sq mi) are smaller than those generally observed in Africa.[113]
The leopard is a
carnivore that prefers medium-sized prey with a body mass ranging from 10–40 kg (22–88 lb). Prey species in this weight range tend to occur in dense habitat and to form small herds. Species that prefer open areas and have well-developed anti-predator strategies are less preferred. More than 100 prey species have been recorded. The most preferred species are
ungulates, such as
impala,
bushbuck,
common duiker and
chital.
Primates preyed upon include
white-eyelid mangabeys,
guenons and
gray langurs. Leopards also kill smaller carnivores like
black-backed jackal,
bat-eared fox,
genet and cheetah.[114]
The largest prey killed by a leopard was reportedly a male
eland weighing 900 kg (2,000 lb).[98] A study in
Wolong National Nature Reserve in southern China demonstrated variation in the leopard's diet over time; over the course of seven years, the vegetative cover receded, and leopards opportunistically shifted from primarily consuming
tufted deer to pursuing
bamboo rats and other smaller prey.[115]
The leopard depends mainly on its acute senses of hearing and vision for hunting.[116] It primarily hunts at night in most areas.[65] In western African forests and Tsavo National Park, they have also been observed hunting by day.[117] They usually hunt on the ground. In the Serengeti, they have been seen to ambush prey by descending on it from trees.[118]
It stalks its prey and tries to approach as closely as possible, typically within 5 m (16 ft) of the target, and, finally, pounces on it and kills it by suffocation. It kills small prey with a bite to the back of the neck, but holds larger animals
by the throat and strangles them.[65] It caches kills up to 2 km (1.2 mi) apart.[74] It is able to take large prey due to its powerful jaw muscles, and is therefore strong enough to drag carcasses heavier than itself up into trees; an individual was seen to haul a young giraffe weighing nearly 125 kg (276 lb) up 5.7 m (18 ft 8 in) into a tree.[117] It eats small prey immediately, but drags larger carcasses over several hundred meters and caches it safely in trees, bushes or even caves; this behaviour allows the leopard to store its prey away from rivals, and offers it an advantage over them. The way it stores the kill depends on local topography and individual preferences, varying from trees in Kruger National Park to bushes in the plain terrain of the Kalahari.[66][119]
Average daily consumption rates of 3.5 kg (7 lb 11 oz) were estimated for males and of 2.8 kg (6 lb 3 oz) for females.[106] In the southern
Kalahari Desert, leopards meet their water requirements by the bodily fluids of prey and
succulent plants; they drink water every two to three days and feed infrequently on moisture-rich plants such as
gemsbok cucumbers,
watermelon and Kalahari
sour grass.[120]
Enemies and competitors
In parts of its range, the leopard is
sympatric with other large predators such as the
tiger,
lion,
cheetah,
spotted hyena,
striped hyena,
brown hyena,
African wild dog,
dhole,
wolf and up to five
bear species. Some of these species steal its kills, kill its cubs and even kill adult leopards. Leopards retreat up a tree in the face of direct aggression, and were observed when killing or preying on smaller competitors such as black-backed jackal,
African civet,
caracal, and
African wildcat.[60][121] Leopards generally seem to avoid encounters with adult bears, killing vulnerable bear cubs instead. In
Sri Lanka, a few recorded fights between leopards and
sloth bears apparently result in both animals winding up either dead or grievously injured.[122][123] Leopards generally avoid large packs of African wild dogs and dholes, and flee up a tree at the sight of them.[98]
While interspecies killing of full-grown leopards is generally rare, given the opportunity, both the tiger and lion readily kill and consume both young and adult leopards.[118][121][124][125] In the
Kalahari Desert, leopards frequently lose kills to brown hyenas, if they are unable to move the kill up a tree. Single brown hyenas have been observed charging at and displacing male leopards from kills.[126][127] Lions occasionally fetch leopard kills from trees.[119]
Resource partitioning occurs where leopards share their range with tigers. Leopards tend to kill smaller prey, usually less than 75 kg (165 lb), where tigers are present.[60]
In areas where leopards and tigers are sympatric, coexistence is reportedly not the general rule, with leopards being few where tigers are numerous.[124] Tigers appear to inhabit the deep parts of the forest while leopards are pushed closer to the fringes.[128] In tropical forests, leopards do not always avoid the larger cats by hunting at different times. With relatively abundant prey and differences in the size of the selected prey, tigers and leopards seem to successfully coexist without competitive exclusion or interspecies dominance hierarchies.[129] In Africa, leopards do not appear to be motivated by an avoidance of lions, as they use heavy vegetation regardless of whether lions are present in an area and both cats are active around the same time of day. In addition, there is no evidence that lions effect leopard abundance.[130]
In some areas, leopards mate all year round. In
Manchuria and
Siberia, they mate during January and February. On average, females begin to breed between the ages of 2½ and three, and males between the ages of two and three.[60] The female's
estrous cycle lasts about 46 days, and she is usually in heat for 6–7 days.[134]Gestation lasts for 90 to 105 days.[135] Cubs are usually born in a
litter of 2–4 cubs.[136] The mortality rate of cubs is estimated at 41–50% during the first year.[106] Lions and spotted hyenas are the biggest cause for leopard cub mortality during their first year. Male leopards are known to cause
infanticide, in order to bring the female back into heat.[74] Intervals between births average 15 to 24 months, but can be shorter, depending on the survival of the cubs.[60]
Females give birth in a cave, crevice among boulders, hollow tree or thicket. Newborn cubs weigh 280–1,000 g (9.9–35.3 oz), and are born with closed eyes, which open four to nine days after birth.[98][66] 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. They begin to eat meat at around nine weeks.[74] Around three months of age, the young begin to follow the mother on hunts. At one year of age, cubs can probably fend for themselves, but will remain with the mother for 18–24 months.[137] After separating from their mother, sibling cubs may travel together for months.[60] Both male and female leopards typically reach sexual maturity at 2–2⅓ years.[74]
The
generation length of the leopard is 9.3 years.[138]
The average life span of a leopard is 12–17 years.[98]
The oldest leopard was a captive female that died at the age of 24 years, 2 months and 13 days.[139]
The leopard is primarily threatened by
habitat fragmentation and conversion of forest to agriculturally used land, which lead to a declining natural prey base,
human–wildlife conflict with livestock herders and high leopard mortality rates. It is also threatened by
trophy hunting and
poaching.[2] Contemporary records suggest that the leopard occurs in only 25% of its historical range.[141][142]
Between 2002 and 2012, at least four leopards were estimated to have been poached per week in India for the
illegal wildlife trade of its skins and bones.[143]
In spring 2013, 37 leopard skins were found during a 7-week long market survey in major Moroccan cities.[144] In 2014, 43 leopard skins were detected during two surveys in Morocco. Vendors admitted to have imported skins from sub-Saharan Africa.[145]
Surveys in the Central African Republic's
Chinko area revealed that the leopard population decreased from 97 individuals in 2012 to 50 individuals in 2017. In this period,
transhumantpastoralists from the border area with Sudan moved in the area with their livestock. Rangers confiscated large amounts of
poison in the camps of livestock herders who were accompanied by armed merchants. They engaged in poaching large herbivores, sale of
bushmeat and trading leopard skins in
Am Dafok.[146]
In Java, the leopard is threatened by illegal hunting and trade. Between 2011 and 2019, body parts of 51 Javan leopards were seized including six live individuals, 12 skins, 13 skulls, 20 canines and 22 claws.[147]
Leopards have been featured in art, mythology and folklore of many countries. In
Greek mythology, it was a symbol of the god
Dionysus, who was depicted wearing leopard skin and using leopards as means of transportation. In one myth, the god was captured by pirates but two leopards rescued him.[148] Numerous Roman mosaics from North African sites depict fauna now found only in
tropical Africa.[149] 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.[150] The
Ashanti people also used the leopard as a symbol of leadership, and only the king was permitted to have a ceremonial leopard stool. Some African cultures considered the leopard to be a smarter, better hunter than the lion and harder to kill.[148]
In
Rudyard Kipling's "How the Leopard Got His Spots", one of his Just So Stories, a leopard with no spots in the
Highveld lives with his hunting partner, the Ethiopian. When they set off to the forest, the Ethiopian changed his brown skin, and the leopard painted spots on his skin.[151] A leopard played an important role in the 1938 Hollywood film Bringing Up Baby. African chiefs, European queens, Hollywood actors and
burlesque dancers wore coats made of leopard skins.[148]
The
Leopard of Rudraprayag killed more than 125 people; the Panar Leopard was thought to have killed over 400 people. Both were shot by British hunter
Jim Corbett.[154] The spotted devil of Gummalapur killed about 42 people in Karnataka, India.[155]
In captivity
The
ancient Romans kept leopards in captivity to be slaughtered in
hunts as well as
execute criminals.[148] In Benin, leopards were kept and paraded as mascots, totems and sacrifices to deities.[150] Several leopards were kept in a menagerie originally established by King
John of England 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.[156] In modern times, leopards have been
trained and
tamed in
circuses.[148]
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