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Overview

Brief Summary

The Leopard (Panthera pardus) has an extremely broad distribution across Africa and South and Southeast Asia. Leopards are found throughout Africa where there is sufficient cover and from the Arabian Peninsula through Asia to Manchuria and Korea. In the African rainforests and Sri Lanka, the Leopard is the only large predator. The black spot seen in the center of each rosette on a Jaguar's coat is typically lacking in Leopards. Melanistic Leopards ("black panthers"; melanistic Jaguars may also be referred to by this name) sometimes occur in several parts of Africa, but are more common in Thailand, Malaysia, and Java (Indonesia).

Leopards are found in an extraordinary range of habitats. In sub-Saharan Africa, they may be found in any habitats with annual rainfall greater than 50 mm, as well as along rivers penetrating true deserts. Leopards in the Kalahari Desert can reportedly go 10 days without drinking. In deserts with temperatures reaching 70 C, Leopards can survive by seeking shelter during the day in caves, animal burrows, and dense vegetation. In Central and West Africa, Leopards occur in rainforests receiving more than 1500 mm annual rainfall. Leopards are common throughout the Indian subcontinent in savannahs, acacia grasslands, deciduous and evergreen forests, and scrub woodlands. They may occur to 5200 m elevation in the mountains of Pakistan and Kashmir. In Southeast Asia, they occur in dense primary rainforest, among other habitats. In the Russian Far East, they may be found in forested mountainous regions where the snow is less than 15 cm deep. Leopards are capable of persisting in close proximity to humans.

The diet of the Leopard is highly varied, including both large and small prey. It often consists mainly of small and medium-sized mammals (5 to 45 kg), but may range from large beetles to ungulates (hoofed mammals) several times their size. Most hunting occurs at night. Sunquist and Sunquist (2009) review the diet of Leopards as reported from different portions of their range.

Like other felids (i.e., members of the cat family), Leopards commonly kill their prey with a bite to the throat, although smaller prey may be dispatched with a bite to the nape or back of the head. Large prey items may be dragged up into a tree and cached there, especially in Africa, where carcasses may otherwise be taken over by hyenas or lions. In Sri Lanka, where the Leopard is the only large carnivore, Leopards are reportedly often seen in open areas during the day.

In the wild, mating associations last just a day or two. The gestation period is around 96 days and young are born at 400 to 600 g. Litter size is typically one to three young (usually two, maximum six). Young travel with their mother starting at three to six months (when they weigh around three or four kg) and begin to eat meat. Permanent canines are emerged at around seven to eight months and the young are typically independent by 12 to 18 months (athough sometimes significantly later). Sexual maturity is reached at two to three years of age.

In some parts of their range, Leopards are endangered, whereas in other places they are considered pests.

(Estes 1991; Sunquist and Sunquist 2009)

  • Estes, R.D. 1991. The Behavior Guide to African Mammals. University of California Press, Berkeley.
  • Sunquist, M.E. and F.C. Sunquist. 2009. Family Felidae (Cats). Pp. 54-168 in: Wilson, D.E. and R.A. Mittermeier (eds.). Handbook of the Mammals of the World. Vol. 1. Carnivores. Lynx Edicions, Barcelona.
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MammalMAP: Leopard

The leopard is the smallest of four cats belonging to the Genus Panthera.  Its appearance is similar to a jaguar but it is slightly smaller, stockier in build and its rosettes are smaller, more densely packed and usually do not have central spots. Ambiguously, both melanistic forms of leopards and jaguars are called black panthers.

These beautiful animals are phenomenally adapted to a wide range of habitats.  It can run up to 58 km/hr, climb trees while carrying a carcass and is renowned for its stealth capabilities. An opportunistic hunter, it will feed on any animal it can catch from arthropods to large antelopes the size of an eland.

Depending on region, leopards may mate all year round.  A female typically gives birth to a litter of 2 – 4 cubs in a den in cave, boulder crevice or hollow tree.  After 3 months, cubs will follow their mother on hunts.  Mortality rate in cubs during their first year is at approximately 50%.

Leopards may have a wide range but their population is declining due to habitat loss and fragmentation.  They are also hunted for trade and pest control.  The IUCNclassifies leopards as a Near Threatened species that may soon slip into the Vulnerable category.

For more information on MammalMAP, visit the MammalMAP virtual museum or blog.

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Comprehensive Description

Miscellaneous Details

"Large leopards, or panthers as Jerdon calls them, often kill cattle, ponies, donkeys, and large deer such as sambar, but the smaller varieties have to content themselves with inferior prey. The leopard, however, is absolutely without prejudice in the matter of food all beasts, birds, and, I believe, reptiles that are not too large to kill or too small to catch are the same to him ; he will strike down an ox or bound upon a sparrow. If he has a predilection, it is probably for dogs and jackals. He is a terrible foe to monkeys, and kills many of the hanumans or langurs who inhabit the rocky hills in which he delights. Leopards, like tigers, sometimes kill their prey by breaking the neck ; but I am disposed to believe that they frequently either tear open the throat or hold it in Iheir jaws and strangle their victim. However, I have not had many opportunities of seeing animals killed by them. They carry away the body like tigers, and hide what they do not eat, very often in a tree. (Blanford, 1888)."
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Summary

"The Leopard is the smallest of the four """"Big Cats"""". It is well known for its stealth and its adaptibility to different habitats, which make it more successful in the wild than other big cats."
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Leopards are large spotted cats with long muscular bodies, but a sizable portion of their population is melanistic (solid black), especially in forest habitats. Whether spotted or black, this species’ camouflage is so effective that biologists report being unable to see them, even when only a few yards away.

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Distribution

Range Description

The leopard occurs across most of sub-Saharan Africa, as remnant populations in North Africa, and then in the Arabian peninsula and Sinai/Judean Desert (Egypt/Israel/Jordan), south-western and eastern Turkey, and through Southwest Asia and the Caucasus into the Himalayan foothills, India, China and the Russian Far East, as well as on the islands of Java and Sri Lanka (Nowell and Jackson 1996; Sunquist and Sunquist 2002; Hunter et al. in press).

In sub-Saharan Africa, leopards remain widely, albeit now patchily, distributed within historical limits (see Hunter et al. in press, and references therein). Ray et al. (2005) estimated that leopards have disappeared from at least 36.7% of their historical range in Africa. The most marked range loss has been in the Sahel belt, as well as in Nigeria and South Africa. They have been locally extirpated from areas densely populated with people or where habitat conversion is extreme (Hunter et al. in press). They are likely extinct on Zanzibar, where there have been no confirmed records since 1996 (Hunter et al. in press).

In North Africa, a tiny relict population persists in the Atlas Mountains of Morocco (Cuzin 2003), and there was a probable observation on the Morocco-Algerian border in Figuig in 2007 (F. Cuzin pers. comm.), while a population was recently found in the Ahaggar of south-eastern Algeria, a region from which they had not previously been recorded (Busby et al. 2006). Leopard are likely extinct in Egypt, although they may occur in the Eastern Desert (Hunter et al. in press).

A 2006 Arabian Fauna Conservation Workshop estimated there were fewer than 200 leopards remaining on the Arabian peninsula, in three confirmed separate subpopulations: the Negev desert, the Wada'a mountains of Yemen, and the Dhofar mountains of Oman. Presence in Saudi Arabia is uncertain (Breitenmoser 2006, Spalton and Al Hikmani 2006).

I. Khorozyan (pers. comm. 2008) compiled detailed country information on the distribution of the Endangered Persian leopard as follows:

Russian North Caucasus: mountain ridges in the headwaters of the Avarskoe Koisu and Andiiskoe Koisu rivers (Republic of Dagestan). Possibly exists in the Chegem River canyon (Kabardino-Balkarian Republic); Erzi Reserve, Assa River valley (Republic of Ingushetia); Armkhi River basin (Republic of North Osetia-Alania), headwaters of the Sharoargun and Argun rivers (Chechen Republic) (Akkiev and Mokaev, 2006; Khorozyan and Abramov, 2007; Lukarevsky et al., 2007).

Georgia: Vashlovani Reserve in the south-east; Arkhoti River canyon in the upper part of the Assa River basin and the headwaters of the Andiiskoe Koisu River in the north-east (Lukarevsky et al., 2007). Some anecdotal records from south-western Georgia are either unreliable or can be attributed to individuals coming from north-eastern Turkey (Arabuli, 2006; Khorozyan and Abramov, 2007).

Armenia: south-western and southern parts of the country from Khosrov Reserve to the Armenian-Iranian state border throughout the Geghama, Zangezur, Aiotsdzor, Bargushat and Meghri ridges. The range boundaries are the Azat River in the north-west; Vardenis Ridge in the north; semi-desert of the Ararat Valley in the west; state border with Azerbaijan and the alpine meadow/nival belt transition zone in the south-west and east; Arax River basin along the Armenian-Iranian border in the south. Until the early 1970s it lived also in north-eastern parts of Armenia (Khorozyan et al., 2005; Khorozyan and Abramov, 2007).

Azerbaijan: Talysh Mts. in the extreme south-east, Akhar-Bakhar Ridge of the Iori-Mingechaur Highland in the north-west and the Zangezur Ridge in the Nakhichevan Republic along the state border with Armenia in the west (Lukarevsky et al., 2007).

Nagorno-Karabakh Republic: distribution in Shushi, Mardakert and Hadrut districts and in the adjoining Kelbajar district was recorded in 1941-1967 (Alekperov, 1966; Sludsky, 1973). Up-to-date information on leopard status is impossible to obtain for the political tension between Armenia and Azerbaijan over this republic, even though it plays a vital role as a corridor between the southern (Armenia, Azerbaijan's Nakhichevan Republic), central (Iori-Mingechaur Highland in Azerbaijan and Vashlovani Reserve in Georgia) and northern (Russian North Caucasus) parts of the Caucasus (Khorozyan and Abramov, 2007).

Iran: virtually all country, except the vast deserts of Desht-e-Kevir and Desht-e-Lut in central and eastern parts. Particularly common in the Alborz Mts. along the southern fringe of the Caspian Sea. Quite common in protected areas (e.g., Tandooreh, Sarigol, Bafgh, Golestan, Kolah'ghazy, Touran, Kavir, Khojir, Khabr and Bamu national parks; Kiamaki and Naybandan wildlife refuges; Jahan Nama, Central Alborz, Varjin, Arasbaran, Dena and Bahram'gur protected areas) and some unprotected lands (Chapur-Ghoymeh, Safee Abad-Dozain or Minoo Dasht, Ramsar, Khaeez and Darestan-Rudbar) (Joslin and Shoemaker 1988; Kiabi et al., 2002; Farhadinia et al. 2007; Abdoli et al. 2008; M Farhadinia and A. Ghoddousi pers. comms., 2008).

Turkey: north-east (around the Artvin city), east (vicinities of Mt. Ararat or Agri) and south-east (Bitlis Ridge). Possibly exists in the mountains of the Black Sea coast and south-westwards to the Taurus Mts. (H. Diker pers. comm., 2008). It is unclear whether leopards still survive in western Turkey.

Turkmenistan: western Kopetdag Ridge, central Kopetdag Ridge, eastern Kopetdag Ridge, Badkhyz Reserve and Giaz-Gyadyk Ridge (Lukarevsky, 2001).

Afghanistan: central (Hindu Kush, Kohe Baba, Kohe Paghman and Safed Koh ranges of the central highlands), north-eastern (Wakhan corridor) and northern (Darkad peninsula of Badakshan) parts of the country (Habibi, 2004).

In the Central Asian republics, leopard distribution is poorly known. Historically, leopards had a wider distribution in Turkmenistan, and were found in parts of Uzbekistan and Tajikistan. An old male leopard was killed by a local hunter in January 2000 in Kazakhstan, the first record of the species in this country, in a location over 600 km from possible occurrences in Uzbekistan and Tajikistan, and over 1,200 km from known occurrences in Turkmenistan. It is aplso possible that the leopard travelled along the foothills of the Pamirs, then proceeded via the Ugam and Pskem ranges into the Talas river valley. Habitat appears to be suitable, but the existence of any leopard subpopulation in any of these three countries is uncertain (Shakula 2004).

In Pakistan, the leopard is thinly distributed in montane areas, and there have only been a handful of confirmed records in recent years (Ahmed 2001).

Leopards occur widely in the forests of the Indian sub-continent, through Southeast Asia and into China, although they are becoming increasingly rare outside protected areas. They are not found on the islands of Borneo or Sumatra (Nowell and Jackson 1996).
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There are nine subspecies of Panthera pardus, which are distributed as follows: Panthera pardus pardus is in Africa; Panthera pardus nimr, Arabia; Panthera pardus saxicolor, Central Asia; Panthera pardus melas, Java; Panthera pardus kotiya, Sri Lanka; Panthera pardus fusca, the Indian sub-continent; Panthera pardus delacourii, southeast Asia into southern China; Panthera pardus japonensis, northern China; and Panthera pardus orientalis, far east Russia, on the Korean peninsula and in north-eastern China.

Biogeographic Regions: palearctic (Native ); oriental (Native ); ethiopian (Native )

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Distribution in Egypt

Localized (South Sinai).

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Historic Range:
Africa, Asia

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Physical Description

Morphology

Body size and color patterns of leopards varies geographically and probably reflects adaptations to particular habitats. Leopards have short legs relative to their long body. They have a broad head, and their massive skull allows for powerful jaw muscles. The leopard's scapula has specialized attachment sites for climbing muscles. They have small round ears, long whiskers extending from dark spots on the upper lip, and long whiskers in their eyebrows that protect their eyes while moving through dense vegetation. Their coat ranges from tawny or light yellow in warm, dry habitats to reddish-orange in dense forests. Subspecies are distinguished according to unique pelage characteristics. Their body is covered with black rosettes, which are circular in East Africa and square in South Africa. They have solid black spots on their chest, feet, and face and rings on their tail. Cubs have a smoky gray coat and their rosettes are not yet distinct. Each individual has a unique coat, which can be used for identification. Black panthers, which are most populous in humid forests, are leopards with recessive melanistic genes. Savannah and woodland leopards tend to be relatively large while mountain and desert leopards tend to be relatively small. Leopards are sexually dimorphic as males tend to be larger than females. Females range in body mass from 17 to 58 kg and in length from 1.7 to 1.9 m. Males range in mass from 31 to 65 kg and in length from 1.6 to 2.3 m.

Range mass: 17 to 65 kg.

Range length: 1.6 to 2.3 m.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

  • 1997. "Thinkquest: Library" (On-line). Wild Habitat: Leopard (Panthera pardus). Accessed March 18, 2009 at http://library.thinkquest.org/11234/leopard_any.html.
  • Hunter, L., G. Hinde. 2005. Cats of Africa: Behavior, Ecology, and Conservation. United Kingdom: New Holland Publisher.
  • Nowell, K., P. Jackson. 1996. Wild Cats: status survey and conservation action plan. Gland, Switzerland: IUCN.
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"Size considerably less than that of the tiger or lion, the length of the head and body seldom exceeding 4 ft., with the tail considerably over half the length of the head and body, the hair on the cheeks never growing into a definite fringe longer than that of the body and, although the hair on the nape typically forms a median crest from the junction of convergent streams from the sides of the neck, as in lions, it never, even in the male. forms anything approaching a definite mane. The coat varies greatly in length, thickness, and texture, according to the season, in some races ; the ground colour above is also exceedingly variable, from greyish or whitish-buff, with sometimes hardly a trace of buff, to olivaceous with a huffish tinge, or to bright reddish-oehreous ; below, from the front of the upper lip and chin backwards, including the lower cheek, the lower side of the tail and the inner surfaces of the limbs, it is generally wholly white, and sharply contrasted with the tint of the upper side and outside of the limbs, though the paws may be paler than the rest of the limbs ; there is a white patch on the otherwise black back of the ear, but there is no white patch over the eye as in the tiger. The pattern is very different from that of the tiger, but not unlike the pattern of some lion-cubs. Itconsists of solid black spots on the head, sometimes for a short distance behind it, on the outer side of the limbs and on the belly, which is typically heavily spotted, but elsewhere on the body the pattern mostly consists of definite """" rosettes """" which appear to result from the coalescence of some four or five smallish solid spots to form definite but irregularly shaped rings surrounding an area of darker tint as a rule, especially low down on the sides, than the brighter tint of the interspaces between the spots. The rosettes vary considerably in size and spacing, and on the spine, especially over the hind back and loins, they are usually more elongated and show a tendency to run in longitudinal stripe-like lines. On the tail the pattern is less regular and less rosette-like, the individual rosettes being liable to coalesce more or less towards the end above, whereas the underside here may be almost wholly unspotted, so that. when upturned it looks white. The normal pattern of leopards, above described, with its suggestion here and there of longitudinally lineal arrangement, is much more primitive than the highly specialized transverse stripes of the tiger. On account of its extensive distribution and capacity for adaptation to widely different environments the leopard varies locally in coloration much more than the lion or tiger, and many geographical races or subspecies of it have been described. Those admitted in the Indian fauna are considered below. But the species is also susceptible to an unusual amount of individual variation in colour and pattern in the same locality. These variations are termed """" varieties """" or """" spots."""" The """" black """" variety is the commonest of them. In this type the blackness is due to the deposit of black or dark brown pigment in the hairs that are normally yellow or white, so that the spots are obscured although probably always visible in certain lights and generally at least better defined on the underside because the normally white hair is browner than the normally yellow hair of the upper side. Leopards sometimes also exhibit the opposite phenomenon, namely, failure to develop pigment in the normally pigmented areas. This may result in complete albinism, when the pattern as well as the interspaces are white."
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Type Information

Type for Panthera pardus
Catalog Number: USNM 181600
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Mammals
Sex/Stage: Male; Adult
Preparation: Skin; Skull
Collector(s): Aggate & E. Heller
Year Collected: 1911
Locality: Loita Plains, Aggate's Farm, Rift Valley, Kenya, Africa
  • Type: Heller, E. 1913 Nov 08. Smithsonian Miscellaneous Collections. 61 (19): 5.
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Type for Panthera pardus
Catalog Number: USNM 164764
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Mammals
Sex/Stage: Male; Adult
Preparation: Skin; Skull
Collector(s): E. Heller
Year Collected: 1910
Locality: Gondokoro, 7 mi E, Central Equatoria, Sudan, Africa
Microhabitat: lake
  • Type: Heller, E. 1913 Nov 08. Smithsonian Miscellaneous Collections. 61 (19): 6.
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Ecology

Habitat

Leopards inhabit a variety of terrain. They are most populous in mesic woodlands, grassland savannas, and forests. They also occupy mountainous, scrub, and desert habitats. They favor trees throughout their entire geographic distribution, and have been recorded at 5638 meters on Mt. Kilimanjaro.

Range elevation: 5638 m (high) m.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: desert or dune ; savanna or grassland ; forest ; rainforest ; scrub forest ; mountains

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Habitat and Ecology

Habitat and Ecology
The leopard has the widest habitat tolerance of any Old World felid, ranging from rainforest to desert. In Africa, they are most successful in woodland, grassland savanna and forest but also occur widely in mountain habitats, coastal scrub, swampy areas, shrubland, semi-desert and desert. They range from sea level to as much as 4,600 m on Mt Kenya (Hunter et al. in press). In Southwest and Central Asia, leopards formerly occupied a range of habitats, but now are confined chiefly to the more remote montane and rugged foothill areas. Through India and Southeast Asia, Leopard are found in all forest types, from tropical rainforest to the temperate deciduous and alpine coniferous (up to 5,200 m in the Himalaya), and also occur in dry scrub and grasslands (Nowell and Jackson 1996).

Leopards have extremely catholic diets including more than 90 species in sub-Saharan Africa, ranging from arthropods to large antelope up to the size of adult male Eland Tragelaphus oryx (Hunter et al. in press). Densities vary with habitat, prey availability, and degree of threat, from fewer than one per 100 km² to over 30 per 100 km², with highest densities obtained in protected East and southern African mesic woodland savannas (Hunter et al. in press).

A study in Thailand found a home range of 8.8 km² for a radio-collared female, and 17.3-18 km² for two adult males (Grassman 1998). Important prey species were hog badger Arctonyx collaris (45.9%), muntjac Muntiacus muntjak (20.9%) and wild pig Sus scrofa (6.3%).

Systems
  • Terrestrial
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Leopards could, at one time, be found throughout the Old World from Europe and Africa to Asia. Although fragmented Asian populations exist, Africa is their remaining stronghold. The versatile leopard inhabits dense lowland forests, rugged mountains, savanna, bush and even semi-deserts.

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Trophic Strategy

Leopards are ambush predators, pouncing on their prey before it chance to react. They approach potential prey by crouching low to the ground, getting as close as 3 to 10 m to prey before pouncing. Leopards are not likely to chase prey after the first pounce. Once a prey item is captured, they immediately break the prey's neck, causing paralysis. After breaking the prey's neck, leopards asphyxiate them and carry the carcass to a secluded feeding location, typically in a nearby tree. They may also cover prey carcasses in leaves and soil. Their tremendous strength allows them to tackle prey up to 10 times their own weight.

Leopards generally prey upon mid-sized ungulates, which includes small antelopes (Bovidae), gazelles (Gazella), deer (Cervidae), pigs (Sus), primates (Primates) and domestic livestock. They are opportunistic carnivores and eat birds (Aves), reptiles (Reptilia), rodents (Rodentia), arthropods (Arthropoda), and carrion when available. Leopards prefer prey that weigh between 10 and 40 kg. They are also known to scavenge from cheetahs (Acinonyx jubatus), solitary hyenas (Hyaenidae), and smaller carnivores as well. They are known to cache food and may continue hunting despite having multiple carcasses already cached.

Animal Foods: birds; mammals; reptiles; fish; carrion ; insects

Foraging Behavior: stores or caches food

Primary Diet: carnivore (Eats terrestrial vertebrates)

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Associations

Threatened Vertebrate Associates in the Hindu Kush Alpine Meadow Ecoregion

The Hindu Kush alpine meadow has an expanse of some 10,900 square miles, situated in northeastern Afghanistan and northern Pakistan. Most of the lands lie within the Hindu Kush Mountain Range in  the altitude bracket between 3000 to 4000 meters, and correspondingly most of the precipitation is in the form of snow. This ecoregion is classified within the Montane Grasslands and Shrublands biome.

This ecoregion manifests a low rate of vertebrate endemism; however there are ten special status mammals found here, ranging from the status of Endangered to Near Threatened. The Hindu Kush alpine meadow ecoregion consists of higher elevation terrain of moderate to severe slopes. Vegetation is often sparse or almost lacking, with resulting pastoral usage of low intensity grazing of goats and sheep in some areas. Soils are largely leptosols, but many areas are covered by large expanses of rock outcrop or rocky scree. In the limited areas of arable soils, wheat is sometimes farmed, although growing of opium poppies is the only cash crop. Most of the water available for plant and animal life is supplied by snowmelt. The Helmand River, Afghanistan's largest watercourse, represents the chief catchment within the ecoregion, with headwaters rising in the Hindu Kush Range, and eventual discharge to the endorheic Sistan Basin.

Special status mammals found in the Hindu Kush alpine meadow are: the Near Threatened argali (Ovis ammon), the Vulnerable Asiatic black bear (Ursus thibetanus), the Near Threatened European otter (Lutra lutra), the Near Threatened leopard (Panthera pardus), the Endangered markhor (Capra falconeri), the Near Threatened mountain weasel (Mustela altaica), the Near Threatened Schreiber's long-fingered bat (Miniopteris schreibersi), the Endangered snow leopard (Uncia uncia), the Near Threatened striped hyena (Hyaena hyaena) and the Endangered Moschus leucogaster. Special status birds in the Hindu Kush alpine meadow are represented by the Endangered Egyptian vulture (Neophron percnopteris).

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Threatened Vertebrate Associates in the Hindu Kush Alpine Meadow Ecoregion

The Hindu Kush alpine meadow has an expanse of some 10,900 square miles, situated in northeastern Afghanistan and northern Pakistan. Most of the lands lie within the Hindu Kush Mountain Range in  the altitude bracket between 3000 to 4000 meters, and correspondingly most of the precipitation is in the form of snow. This ecoregion is classified within the Montane Grasslands and Shrublands biome.

This ecoregion manifests a low rate of vertebrate endemism; however there are ten special status mammals found here, ranging from the status of Endangered to Near Threatened. The Hindu Kush alpine meadow ecoregion consists of higher elevation terrain of moderate to severe slopes. Vegetation is often sparse or almost lacking, with resulting pastoral usage of low intensity grazing of goats and sheep in some areas. Soils are largely leptosols, but many areas are covered by large expanses of rock outcrop or rocky scree. In the limited areas of arable soils, wheat is sometimes farmed, although growing of opium poppies is the only cash crop. Most of the water available for plant and animal life is supplied by snowmelt. The Helmand River, Afghanistan's largest watercourse, represents the chief catchment within the ecoregion, with headwaters rising in the Hindu Kush Range, and eventual discharge to the endorheic Sistan Basin.

Special status mammals found in the Hindu Kush alpine meadow are: the Near Threatened argali (Ovis ammon), the Vulnerable Asiatic black bear (Ursus thibetanus), the Near Threatened European otter (Lutra lutra), the Near Threatened leopard (Panthera pardus), the Endangered markhor (Capra falconeri), the Near Threatened mountain weasel (Mustela altaica), the Near Threatened Schreiber's long-fingered bat (Miniopteris schreibersi), the Endangered snow leopard (Uncia uncia), the Near Threatened striped hyena (Hyaena hyaena) and the Endangered Moschus leucogaster. Special status birds in the Hindu Kush alpine meadow are represented by the Endangered Egyptian vulture (Neophron percnopteris).

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Leopards compete for food with lions (Panthera leo), tigers (Panthera tigris), spotted hyenas (Crocuta crocuta), and African wild dogs (Lycaon pictus). To avoid attacks from potential predators, leopards tend to hunt at different times of the day and avoid areas where potential predators are most populous. When competition for larger prey items is high, leopards prey on smaller animals, which reduces interspecific competition. Leopards are host to many common felid parasites, including lung flukes (Paragominus westermani), flat worms (Pseudophyllidea), spirurian nematodes (Spiruroidea), hookworms (Ancylostomatidae), lung worms (Aelurostrongylus), intestinal and hepatic parasites (Capillaria), and parasitic protozoa (Sarcocystis).

Commensal/Parasitic Species:

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Humans are the primary predator of leopards. Leopards are hunted as trophy animals for their fur, and retaliatory killings by farmers protecting their livestock are not uncommon. Lions (Panthera leo), tigers (Panthera tigris), spotted hyenas (Crocuta crocuta), and African wild dogs (Lycaon pictus) prey upon leopard cubs and are capable of killing adult leopards. Typically, when an adult is killed it is due to a territorial confrontation. Many of the characteristics that make leopards great predators also serve as excellent predator defense mechanisms. For example, a leopard's spots allows them to travel inconspicuously and avoid detection.

Known Predators:

Anti-predator Adaptations: cryptic

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Known predators

Panthera pardus is prey of:
Crocodylidae
Canis lupus
Panthera leo
Panthera tigris

This list may not be complete but is based on published studies.
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Known prey organisms

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Predator: prey includes wildebeest, impalas, reed-bucks, Thomson's gazelles, jackals, baboons and storks.

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Life History and Behavior

Behavior

Although leopards are silent most of the time, they may give a hoarse, rasping cough at repeated intervals to advertise their presence to conspecifics. Males use this unique call to announce territorial boundaries. If another leopard is in the vicinity, it may answer with a similar vocalization and continue vocalizing as it exits the area. Males also grunt at each other and females call to potential mates when in estrous. Some leopards may purr while feeding.

Communication Channels: tactile ; acoustic ; chemical

Other Communication Modes: pheromones ; scent marks

Perception Channels: visual ; tactile ; acoustic ; chemical

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Behaviour

"It does not confine itself to the forests, but prowls among villages, carrying off sheep, goats, dogs, and sometimes commits great depredations. It is very fearless, frequently seizing a dog in the middle of a village, whilst the inhabitants are still stirring. It always seizes its prey by the back of the neck or the throat. It is a particularly silent creature, very stealthy, and will contrive to dodge and hide itself in places where it would appear impossible that a creature of its size could find concealment. They are popularly said to be much in the habit of climbing trees. They are fearful of water, and will not readily swim, and are therefore rarely found on small islands. Like the Tiger, the Leopard will, if hungry, eat any dead carcass he can find. Leopards occasionally take to man-eating and, owing to their boldness, become even a more fearful scourge than tigers."
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Leopards are probably the most accomplished stalkers and climbers of the big cats. Their varied diet includes wildebeest, impalas, reed-bucks, Thomson's gazelles, jackals, baboons and storks. They routinely drag carcasses bigger than themselves into trees to avoid losing prey to other carnivores.

Leopards are most active at night and generally solitary. Females den in caves, crevices, thickets, or hollow trees and give birth to two to three cubs, of which only one or two will survive. Cubs hunt with their mother at about three months of age and remain with her for another two years.

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Life Expectancy

In captivity, leopards can live to be 21 to 23 years old, with the oldest known individual being 27 years old. Wild leopards may live to be 10 to 12 years old, with the oldest known individual being 17 years old. Survival rates for cubs range from 41% to 50%.

Range lifespan

Status: wild:
17 (high) years.

Range lifespan

Status: captivity:
27 (high) years.

Typical lifespan

Status: wild:
10 to 12 years.

Typical lifespan

Status: captivity:
21 to 23 years.

  • Guggisberg, C. 1975. Wild Cats of the World. New York: Taplinger Publishing Company.
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Lifespan, longevity, and ageing

Maximum longevity: 27.3 years (captivity) Observations: One specimen lived 27.3 years at Madrid Zoo. A hybrid between a leopard and a lion lived for 24 years (Richard Weigl 2005).
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Reproduction

Leopards are promiscuous, as both males and females have multiple mates. Females attract potential mates by excreting pheromones in their urine. Females initiate mating by walking back and forth in front of a male and brushing up against him or swatting him with her tail. The male then mounts the female while frequently biting her nape. Copulation last an average of three seconds with six minute intervals between each copulation bout. A single breeding pair may copulate up to 100 times per day for several days, during which time they share food resources.

Mating System: polygynandrous (promiscuous)

The reproductive season is year-round but peaks during the rainy season in May. In China and southern Siberia, leopards mainly breed in January and February. Females are in estrus for 7 days and have a 46 day long cycle. Gestation last 96 days and females usually give birth once every 15 to 24 months. Typically, females stop reproducing around 8.5 years old.

Breeding interval: Leopards breed every 15 to 24 months

Breeding season: Leopards breed year-round, with a peak during the rainy season

Range number of offspring: 2 to 3.

Average gestation period: 96 days.

Average weaning age: 3 months.

Range time to independence: 13 to 18 months.

Average age at sexual or reproductive maturity (female): 2.5 years.

Average age at sexual or reproductive maturity (male): 2 years.

Key Reproductive Features: iteroparous ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous

Average birth mass: 550 g.

Average gestation period: 97 days.

Average number of offspring: 2.

Average age at sexual or reproductive maturity (male)

Sex: male:
771 days.

Average age at sexual or reproductive maturity (female)

Sex: female:
937 days.

Leopard cubs weigh less than 1 kg at birth, and their eyes remain closed for the first week. Mothers leave their cubs in the protection of dense bush, rock clefts, or hollow tree trunks for up to 36 hours while hunting and feeding. They move den sites frequently, which helps prevent cubs from falling prey to lions and other predators. Cubs learn to walk at 2 weeks of age and regularly leave the den at 6 to 8 weeks old, around which time they begin to eat solid food. Mothers share less than a third of their food with their cubs. Cubs are completely weaned by 3 months old and independent at just under 20 months old. Often, siblings maintain contact during the early years of independence. Territories are flexible and young may linger in their natal area.

Parental Investment: altricial ; female parental care ; pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female); extended period of juvenile learning

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Molecular Biology and Genetics

Molecular Biology

Barcode data: Panthera pardus

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 3 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

AACCGCTGACTATTTTCAACCAATCACAAAGATATTGGAACTCTTTACCTTCTATTTGGTGCCTGGGCTGGCATGGTGGGGACTGCTCTC---AGTCTCTTAATCCGAGCCGAACTGGGTCAACCTGGCACACTGCTAGGGGAC---GACCAAATTTATAATGTAGTCGTTACCGCCCATGCTTTTGTAATAATCTTCTTTATAGTAATGCCCATCATGATTGGAGGATTCGGAAACTGATTGGTCCCATTAATA---ATTGGAGCCCCCGATATAGCATTCCCTCGAATGAATAATATGAGCTTTTGACTCCTTCCCCCATCTTTCCTACTTTTGCTCGCATCATCTATGGTAGAGGCTGGGGCAGGAACTGGATGAACAGTATACCCACCCCTAGCCGGCAACCTAGCCCATGCAGGGGCATCCGTAGATTTA---ACTATTTTTTCACTACACCTGGCAGGTGTCTCCTCAATCTTAGGCGCTATTAATTTTATTACTACTATTATTAATATAAAACCCCCTGCTATATCCCAATACCAAACACCTCTATTCGTCTGATCGGTCTTAATCACTGCTGTATTGCTACTCCTATCACTGCCAGTTTTAGCAGCA---GGCATCACTATGCTACTGACAGATCGAAATCTGAACACCACATTCTTTGACCCTGCCGGAGGGGGGGATCCTATCTTATACCAGCACCTATTCTGATTTTTTGGTCACCCAGAAGTTTATATTTTAATTTTACCCGGGTTCGGAATAATTTCACACATTGTCACCTATTACTCAGGTAAAAAA---GAGCCTTTTGGCTATATGGGAATAGTTTGAGCTATAATATCGATTGGCTTCCTGGGCTTTATCGTGTGAGCCCACCACATGTTTACTGTAGGAATAGATGTGGACACACGAGCATACTTTACATCAGCTACTATAATTATTGCTATTCCCACTGGAGTAAAAGTATTTAGCTGACTA---GCCACT
-- end --

Download FASTA File

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Statistics of barcoding coverage: Panthera pardus

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 2
Specimens with Barcodes: 19
Species With Barcodes: 1
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Conservation

Conservation Status

Leopards are declining in parts of their geographic range due to habitat loss and fragmentation, and hunting for trade and pest control. As a result, leopards are listed as "near threatened" on the IUCN Red List of Threatened Species. Leopards appear to show some resistance to minor habitat disturbances and are relatively tolerant of humans. Currently, leopards are protected throughout most of their range in west Asia; however, populations in this part of their range are too small to maintain stable growth. Although habitat reserves and national parks exist throughout their geographic range in Africa, a majority of leopards live outside these protected areas. Although leopards are the most populous of the "great cats", 5 of 9 subspecies are listed as endangered or critically endangered.

US Federal List: endangered

CITES: appendix i

IUCN Red List of Threatened Species: near threatened

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IUCN Red List Assessment


Red List Category
NT
Near Threatened

Red List Criteria

Version
3.1

Year Assessed
2008

Assessor/s
Henschel, P., Hunter, L., Breitenmoser, U., Purchase, N., Packer, C., Khorozyan, I., Bauer, H., Marker, L., Sogbohossou, E. & Breitenmoser-Wursten, C.

Reviewer/s
Nowell, K., Breitenmoser-Wursten, C., Breitenmoser, U. (Cat Red List Authority) & Hoffmann, M. (Global Mammal Assessment Team)

Contributor/s

Justification
Leopards have a wide range and are locally common in some parts of Africa and tropical Asia. However, they are declining in large parts of their range due to habitat loss and fragmentation, and hunting for trade and pest control. These threats may be significant enough that the species could soon qualify for Vulnerable under criterion A.

History
  • 2002
    Least Concern
  • 1990
    Threatened
    (IUCN 1990)
  • 1988
    Threatened
    (IUCN Conservation Monitoring Centre 1988)
  • 1986
    Vulnerable
    (IUCN Conservation Monitoring Centre 1986)
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Current Listing Status Summary

Status: Endangered
Date Listed: 06/02/1970
Lead Region: Foreign (Region 10) 
Where Listed: except where threatened

Status: Threatened
Date Listed: 01/28/1982
Lead Region: Foreign (Region 10) 
Where Listed: Gabon to Kenya & southward


Population detail:

Population location: In Africa, in the wild, south of, and including, the following countries: Gabon, Congo, Zaire, Uganda, Kenya
Listing status: T

Population location: Wherever found, except where it is listed as Threatened as set forth below
Listing status: E

For most current information and documents related to the conservation status and management of Panthera pardus , see its USFWS Species Profile

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Red List Category & Criteria: Near Threatened ver 3.1 Year Published: 2008
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Status in Egypt

Native, extinct?

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IUCN Red List Status: NEAR THREATENED

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Population

Population
The leopard is an adaptable, widespread species that nonetheless has many threatened subpopulations. While still numerous and even thriving in some marginal habitats from which other big cats have disappeared in many parts of sub-Saharan Africa, in North Africa leopards are on the verge of extinction.

There are no reliable continent-wide estimates of population size in Africa, and the most commonly cited estimate of over 700,000 leopards in Africa (Martin and de Meulenaar 1988) is flawed. In India, based on pugmark censuses (a methodology which has been criticized as inaccurate), 9,844 leopards were estimated in 2001. Many populations are believed to be increasing (Singh 2005), and there are high levels of human-leopard conflict (Singh et al. 2008).

Several Asian subspecies are included on the Red List, with population information as follows:

Amur leopard P.p. orientalis CR C2a(ii),D: 14-20 (Anon. 2007)
Arabian leopard P.p. nimr CR C2a(I): <200 (Breitenmoser 2006, Spalton and Al Hikmani 2006)
Javan leopard P.p. melas CR C2a(i): 323-525, with <250 mature breeding adults (A. Ario pers. comm. 2007)
Sri Lankan leopard P.p. kotiya EN C2a(i): 700-950 (Kittle and Watson 2007)
Persian leopard P.p. saxicolor EN C2a(i): 871-1290 (Khorozyan et al., 2005; Lukarevsky et al. 2007)

Population Trend
Decreasing
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Threats

Major Threats
Throughout Africa, the major threats to Leopard are habitat conversion and intense persecution, especially in retribution for real and perceived livestock loss (Ray et al. 2005). In intact rainforest, the chief threat to Leopards is probably competition with human hunters for prey; the tremendous volume of wild meat harvests denudes forests of prey and may drive localized extinctions. Nonetheless, Leopard are somewhat tolerant of habitat conversion, and may persist close to large human populations provided they have suitable cover and prey (Hunter et al. in press).

Leopard come into conflict with people across their range. A rapidly increasing threat to Leopards is the poisoning of carcasses targeting carnivores, either as a means of predator control or incidentally.

The impact of trophy hunting on populations is unclear, but may have impacts at the demographic and population level, especially when females are shot. In Tanzania, which allows only males to be hunted, females comprised 28.6% of 77 trophies shot between 1995 and 1998 (Spong et al. 2000).

Skins and canines are still widely traded domestically in some central and West African countries where parts are used in traditional rituals and sold openly in villages and cities (Hunter et al. in press). Djibouti is an important conduit for Leopard skins from East Africa that are bought mainly by French military personnel and carried illegally to Europe.

In West Asia, small leopard subpopulations are threatened primarily by habitat fragmentation, killing in defence of livestock, and poaching for trade (Habibi 2004, Breitenmoser et al. 2006, Breitenmoser et al. 2007).

In Indo-Malaya, leopards are threatened primarily by habitat loss (deforestation) as well as poaching for illegal trade (Nowell and Jackson 1996). In India, leopards are feared for their attacks on people (Singh 2005).
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"Persecution, poaching, conflict."
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Leopards are highly adaptable but they still face many threats. Growing human populations are increasingly encroaching on leopard habitat and killing these cats in fear or for trophies. Illegal hunting of leopards became so common in the 1960s that as many as 50,000 skins were taken annually.

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Management

Conservation Actions

Conservation Actions
Included on CITES Appendix I. Legal international traffic is limited largely to exports of skins and hunting trophies under a CITES Appendix I quota system by 13 African countries (2005 CITES quota is 2,590). Leopards are protected under national legislation throughout most of their range (Nowell and Jackson 1996). In Africa, although Leopards occur in numerous protected areas across their range, the majority of the population occurs outside of protected areas, necessitating a need for improved conflict mitigation measures (including livestock management, conflict resolution) (Hunter et al. in press). In West Asia, leopards are essentially restricted to protected areas, many of which are too small to support viable populations, and need expansion through buffer zones and connectivity through corridors (Breitenmoser et al. 2006, 2007). In Indo-Malaya and China, leopards need better protection from illegal trade in skins and bones (Nowell 2007). Leopards are protected in Afghanistan having recently been placed on the country's Protected Species List (2009), prohibiting all hunting and trading of the species within Afghanistan.
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Relevance to Humans and Ecosystems

Benefits

When natural prey abundances are low, leopards have been known to kill livestock. Injured or sickly leopards have been known to hunt humans as easy prey.

Negative Impacts: injures humans (bites or stings)

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Leopards can be seen in National Parks throughout Asia and Africa. They help control baboon populations and disperse seeds that stick to their fur. Chiefs and warriors from tribal cultures throughout the leopard's geographic range wear their fur as a symbol of honor and courage. Tribal medicine men and women suggest leopard skins as a remedy for bad omens. Leopards are often captured for pet trade and are targeted by trophy hunters as well.

Positive Impacts: pet trade ; body parts are source of valuable material; ecotourism

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Wikipedia

Leopard

The leopard /ˈlɛpərd/ (Panthera pardus) is one of the five "big cats" in the genus Panthera. It is a member of the Felidae family with a wide range in some parts of sub-Saharan Africa, West Asia, the Middle East, South and Southeast Asia to Siberia.

Compared to other members of the Felidae, 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 are known as black panthers.

The species' success in the wild is in part due to its opportunistic hunting behavior, its adaptability to habitats, its ability to run at speeds approaching 58 kilometres per hour (36 mph), its unequaled ability to climb trees even when carrying a heavy carcass,[2] 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.

It is listed as Near Threatened on the IUCN Red List because it is declining in large parts of its range due to habitat loss and fragmentation, and hunting for trade and pest control. It is regionally extinct in Hong Kong, Singapore, Kuwait, Syria, Libya and Tunisia.[1]

Characteristics[edit]

A one-eyed African leopard in Kruger National Park, South Africa

Leopards show a great diversity in coat color and rosettes patterns. In general, the coat color varies from pale yellow to deep gold or tawny, and is patterned with black rosettes. The head, lower limbs and belly are spotted with solid black. Coat color and patterning are broadly associated with habitat type. 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 be 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 coloured and of a softer, downy type. Solid black spots in place of open rosettes are generally seen along the face, limbs and underbelly.[3]

Leopards are agile and stealthy predators. Although they are smaller than most 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 usually between 90 and 165 cm (35 and 65 in). The tail reaches 60 to 110 cm (24 to 43 in) long, around the same length as the tiger's tail and proportionately long for the genus (though snow leopards and the much smaller marbled cats have relatively longer tails).[4][5] Shoulder height is from 45 to 80 cm (18 to 31 in). 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 30 to 91 kg (66 to 201 lb) compared to 23 to 60 kg (51 to 132 lb) for females. Large males of up to 91 kg (201 lb) have been documented in Kruger National Park in South Africa; however, males in South Africa's coastal mountains average 31 kg (68 lb) and the females from the desert-edge in Somalia average 23 to 27 kg (51 to 60 lb). This wide variation in size is thought to result from the quality and availability of prey found in each habitat. The most diminutive leopard subspecies overall is the Arabian leopard (P. p. nimr), from deserts of the Middle East, with adult females of this race weighing as little as 17 kg (37 lb).[6]

Other large subspecies, in which males weigh up to 91 kg (201 lb), are the Sri Lankan leopard (P. p. kotiya) and the Anatolian leopard (P. p. tulliana). Such larger leopards tend to be found in areas which lack tigers and lions, thus putting the leopard at the top of the food chain with no competitive restriction from large prey items.[7] The largest verified leopards weighed 96.5 kg (213 lb) and can reach 190 cm (75 in) in head-and-body length. Larger sizes have been reported but are generally considered unreliable.[5][8] The leopard's body is comparatively long, and its legs are short.[9]

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.[10]

Variant coloration[edit]

Black Panther at the Bronx Zoo
Persian leopard with atypical coat pattern

Melanistic leopards are commonly called black panthers, a term that also applies to melanistic jaguars. Pseudomelanism (abundism) also occurs in leopards.[11] Melanism in leopards is inherited as a Mendelian, monogenic recessive trait relative to the spotted form. Pairings of black animals inter se have a significantly smaller litter size than other possible pairings.[12] The black color is caused by recessive gene loci.[13]

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.[14] 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 445 photographs of melanistic leopards taken, 410 came from study sites south of the Isthmus of Kra, where the non-melanistic morph was never photographed. These data suggest the near fixation of the dark allele in the region. The expected time to fixation of this recessive allele due to genetic drift alone ranged from about 1,100 years to about 100,000 years.[15][16]

Melanism in leopards has been hypothesized to be causally associated with a selective advantage for ambush.[17]

A rare "Strawberry" leopard has been confirmed to exist at South Africa's Madikwe Game Reserve.[18] It is thought the leopard has erythrism, a little-understood genetic condition that's thought to cause either an overproduction of red pigments or an underproduction of dark pigments

Etymology[edit]

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 λέων leōn (lion) and πάρδος pardos (male panther). The Greek word is related to Sanskrit पृदाकु pṛdāku (snake, tiger, panther), and probably derives from a Mediterranean language, such as Egyptian.[19][20]

A panther can be any of several species of large felids: the term can refer to cougars and jaguars in the American continents but (given the European origins of the word) it is largely thought to define the leopard at its source. Black panther refers to leopards with melanistic genes, which are not uncommon in rainforest habitats.[21]

The generic component of its modern scientific designation, Panthera pardus, derives from Latin via Greek πάνθηρ (pánthēr).[22] Folk etymology saw the name as a compound of παν (pan, all) and θηρ (beast).[23] 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.[20][22]

Taxonomy and evolution[edit]

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.

The leopard was among the first animals named under the modern system of biological classification, since it was described by Carl Linnaeus in 1758 in the 10th edition of Systema Naturae. Linnaeus placed the leopard under the genus Felis as the binominal Felis pardus.[24] 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 species. But most disagreed with his definition, and until the beginning of the 20th century continued using Felis or Leopardus when describing leopard subspecies.[25] In 1916, Reginald Innes Pocock accorded Panthera generic rank defining Panthera pardus as species.[26]

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.[27]

Results of phylogenetic analyses of chemical secretions amongst cats has suggested that the leopard is closely related to the lion.[28] 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.[29]

Fossil records[edit]

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.[30]

In Europe, the leopard is known at least since the Pleistocene. Fossil leopard bones and teeth dating from the Pliocene were found in Perrier in France, northeast of London, and in Valdarno in Italy. At 40 sites in Europe fossil bones and dental remains of leopards dating from the Pleistocene were excavated mostly in loess and caves. The sites of these fossil records range 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.[31] The Pleistocene leopards of Europe can be divided into four subsequent subspecies. The first European leopard subspecies P. p. begoueni is known since the beginning of the early Pleistocene and was replaced about 600,000 years ago by P. p. sickenbergi, which in turn was replaced by P. p. antiqua at around 300,000 years ago. The last 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 large parts of Europe.[32]

Distribution and habitat[edit]

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.[3] 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.[1][33]

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).[30] 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.[33] In India, leopard populations sometimes live quite close to human settlements and even in semi-developed areas.[34] 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.[33][34] Due to the leopard's superlative stealthiness, people often remain unaware that big cats live in nearby areas.[34]

Distribution of subspecies[edit]

Since Carl Linnaeus published his description of leopards in the 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.[35] 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.[30]

The nine subspecies recognised by IUCN are:[1][30][36]

A morphological analysis of characters of leopard skulls implies the validity of two more subspecies:[36]

Persian leopard kept in Hanover's Adventure Zoo, Germany 
Indian leopard at Nagarhole National Park 
Northern Chinese leopard kept in the Jardin des Plantes, Paris 
Sri Lankan Leopard 
Himalayan Leopard (Darjeeling Zoo) 

Ecology and behavior[edit]

Leopard resting on a tree
Leopard in a tree

Leopards are elusive, solitary and largely nocturnal.[37][38] 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.[39]

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.[40] 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.[41] They produce a number of vocalizations, including grunts, roars, growls, meows, and purrs.[42]

Social structure and home range[edit]

Female leopard in the Sabi Sands area of South Africa. Note the white spot on its tail, used for communicating with cubs while hunting or in long grass.[2]

Home ranges of male leopards vary between 30 km2 (12 sq mi) and 78 km2 (30 sq mi), and of females between 15 to 16 km2 (5.8 to 6.2 sq mi).[3] 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.[43]

Research in a conservation area in Kenya showed similar territory sizes and sex differential: 32.8 km2 (12.7 sq mi) average ranges for males, and 14 km2 (5.4 sq mi) for females.[44]

In Nepal, somewhat larger male ranges have been found at about 48 km2 (19 sq mi), while female ranges at 17 km2 (6.6 sq mi); female home ranges decreased to 5 to 7 km2 (1.9 to 2.7 sq mi) when young cubs were present, while the sexual difference in range size seemed to be in positive proportion to overall increase.[45]

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 100 km2 (39 sq mi) with some more than 300 km2 (120 sq mi). Admitting that their data were at odds with others, the researchers found little or no sexual variation in the size of territories.[46]

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.[37]

Hunting and diet[edit]

Leopard stalking in Okavango Delta, Botswana
Leopard with kill on a tree in Ngala Game Reserve, South Africa

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 genus, and are reported to eat anything from dung beetles to common elands, though medium-sized prey species in the 20–80 kg (44–176 lb) range are usually taken.[3] The largest prey reported killed by a leopard was a 900 kg (2,000 lb) male eland.[47] although leopards generally do not prey on such large animals.[48] Their diet consists mostly of ungulates, followed by primates, primarily monkeys of various species, including the Vervet monkey. However, they will also opportunistically eat rodents, reptiles, amphibians, insects, birds (especially ground-based types like the Vulturine Guineafowl), fish and sometimes smaller predators (such as foxes, jackals, martens and smaller felid species). In at least one instance, a leopard has predated a sub-adult Nile crocodile that was crossing over land.[49] Leopards are the only natural predators of adult chimpanzees and gorillas, although the cat may sometimes choose to avoid these as they are potentially hazardous prey, especially large male silverback gorillas.[50][51] 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.[52]

In the open savanna of Tsavo National Park, they kill most of their prey while hunting between sunset and sunrise.[53] 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 20 to 80 kg (44 to 176 lb) 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 40 to 60 kg (88 to 132 lb). Small mammal remains were found most often in scats of sub-adult leopards, especially females. Average daily consumption rates was estimated at 3.5 kg (7.7 lb) for adult males and 2.8 kg (6.2 lb) for females.[54]

Leopard killing young bushbuck

In Asia, the leopard primarily 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.[55] 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.[56]

They select their prey focusing on small herds, dense habitat, and low risk of injury, preferring prey weights of 10 to 40 kg (22 to 88 lb) such as impala, chital, bushbuck and common duiker with an average body weight of 25 kg (55 lb).[57]

In search of safety, leopards often stash their young or recent kills high up in a tree, which can be a great feat of strength considering that they may be carrying prey heavier than themselves in their mouth while they climb vertically.[52] One leopard was seen to haul a young giraffe, estimated to weigh up to 125 kg (276 lb), more than twice the weight of the cat, up 5.7 m (19 ft) into a tree.[53]

Interspecific predatory relationships[edit]

Video of a leopard in the wild

Leopards must compete for food and shelter with other large predators such as tigers, lions, spotted hyenas, striped hyenas, up to 5 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, African wild cat and the cubs of lions, cheetahs, hyenas, and wild dogs.[3]

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.[3] 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.[58] Mean leopard density decreased significantly (from 9.76 animals/100km2 to 2.07 animals/100km2) 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.[59] 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%).[59] In the Primore region of the Russian Far East, Amur leopards were absent or very rarely encountered at places where Siberian tigers reside.[60] 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.[58] 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.[58] There were also differences in the microhabitat preferences of the individual tiger and leopard followed over a 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 [58] Usually when a tiger began to kill baits at sites formerly frequented by leopards, the leopards would no longer come and kill there.[58] 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.[61] 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.[62] 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.[58]

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.[63] Occasionally, Nile crocodiles may predate on leopards of any age.[64] Lions are occasionally successful in climbing trees and fetching leopard kills,[52] but leopards are also known to kill or prey on cubs of large lions.[3] 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.[65][66] Burmese Pythons have been known to prey on leopards, with an adult cat having been recovered from the stomach of an 18-foot specimen.[67]

Reproduction and life cycle[edit]

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.[68] Gestation lasts for 90 to 105 days.[69] Cubs are usually born in a litter of 2–4 cubs.[70] But mortality of cubs is estimated at 41–50% during the first year.[54]

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.[47] 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.[40]

The average typical life span of a leopard is between 12 and 17 years.[71] 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. [72] 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.[73] 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.[74]

Hybrids[edit]

Pumapard, 1904
Main article: Panthera hybrid

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.[75] Although lions and leopards may come into contact in sub-Saharan Africa, they are not widely believed to interbreed naturally. However, there have been anecdotal reports of small lions with exceptionally pronounced spotting, known as "marozis" and various other names, in several African countries, for which there has been cryptozoological speculation that they may be naturally occurring lion-leopard hybrids.[76]

Crossbreeding between jaguars and leopards in captivity has also been documented.[77][78] A cross between a female leopard and a male jaguar is referred to as a jagupard, the reverse is known a leguar; however, 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.[citation needed] There have also been a few claims of crosses between tigers and leopards.[79]

Main article: Pumapard

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.[80]

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.[80]

Leopards and humans[edit]

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 used to symbolize the power of the king or oba.[81]

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.[82]

Heraldry[edit]

Three leopards (loggerheads) on the flag of Shropshire, England.
Main article: Leopard (heraldry)

The lion passant guardant or leopard is a frequently used charge in heraldry, most commonly appearing in groups of three.[83] 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.[84]

Tourism[edit]

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 typically make leopard sightings rare.[85] 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.[86]

Man-eating[edit]

Main article: Leopard attack

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 Jim Corbett.[87] 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.[88] 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""

In July 2012, two people were killed by leopards in separate attacks in distant parts of India.[89]

In popular culture[edit]

See also[edit]

References[edit]

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