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Asiatic lion
Male Asiatic lion in Gir National Park
Male Asiatic lion in Gir National Park
Female with cub
Female with cub
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species: P. leo
Subspecies: P. l. leo
Population: Asiatic lion
Current range of the Asiatic lion
Current range of the Asiatic lion

The Asiatic lion is a lion population of the subspecies Panthera leo leo. Until the 19th century, it occurred in Saudi Arabia, eastern Turkey, Iran, Mesopotamia, and from east of the Indus River in Pakistan to the Bengal region and the Narmada River in Central India. Since the turn of the 20th century, its range has been restricted to Gir National Park and the surrounding areas in the Indian state of Gujarat. The first scientific description of the Asiatic lion was published in 1826 by the Austrian zoologist Johann N. Meyer, who named it Felis leo persicus.

The population has steadily increased since 2010. In 2015, the 14th Asiatic Lion Census was conducted over an area of about 20,000 km2 (7,700 sq mi); the lion population was estimated at 523 individuals, and in 2017 at 650 individuals.

Taxonomy

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Felis leo persicus was the scientific name proposed by Johann N. Meyer in 1826 who described an Asiatic lion skin from Persia.[2] In the 19th century, several zoologists described lion zoological specimen from other parts of Asia that used to be considered synonyms of P. l. persica:[3]

In 2017, the Asiatic lion was subsumed to P. l. leo due to close morphological and molecular genetic similarities with Barbary lion specimens.[8][9] However, several scientists continue using P. l. persica for the Asiatic lion.[10][11][12][13][14][15] A standardised haplogroup phylogeny supports that the Asiatic lion is not a distinct subspecies, and that it represents a haplogroup of the northern P. l. leo.[16]

Evolution

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Main article: Lion § Evolution and phylogeny

Lions first left Africa at least 700,000 years ago, giving rise to the Eurasian Panthera fossilis which later evolved into Panthera spelaea (commonly known as the cave lion), which became extinct around 14,000 years ago. Genetic analysis of P. spelaea indicates that it represented a distinct species from the modern lion that diverged from them around 500,000 years ago and unrelated to modern Asian lions.[17] Pleistocene fossils assigned as belonging or probably belonging to the modern lion have been reported from several sites in the Middle East, such as Shishan Marsh in the Azraq Basin, Jordan, dating to around 250,000 years ago,[18] and Wezmeh Cave in the Zagros Mountains of western Iran, dating to around 70-10,000 years ago,[19] with other reports from Pleistocene deposits in Nadaouiyeh Ain Askar and Douara Cave, Syria.[18] In 1976, fossil lion remains were reported from Pleistocene deposits in West Bengal.[20] A fossil carnassial excavated from Batadomba Cave indicates that lions inhabited Sri Lanka during the Late Pleistocene. This population may have become extinct around 39,000 years ago, before the arrival of humans in Sri Lanka.[21]

Phylogeography

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Results of a phylogeographic analysis based on mtDNA sequences of lions from across the global range, including now extinct populations like Barbary lions, indicates that sub-Saharan African lions are phylogenetically basal to all modern lions. These findings support an African origin of modern lion evolution with a probable centre in East and Southern Africa. It is likely that lions migrated from there to West Africa, eastern North Africa and via the periphery of the Arabian Peninsula into Turkey, southern Europe and northern India during the last 20,000 years. The Sahara, Congolian rainforests and the Great Rift Valley are natural barriers to lion dispersal.[22]

Genetic markers of 357 samples from captive and wild lions from Africa and India were examined. Results indicate four lineages of lion populations: one in Central and North Africa to Asia, one in Kenya, one in Southern Africa, and one in Southern and East Africa; the first wave of lion expansion probably occurred about 118,000 years ago from East Africa into West Asia, and the second wave in the late Pleistocene or early Holocene periods from Southern Africa towards East Africa.[23] The Asiatic lion is genetically closer to North and West African lions than to the group comprising East and Southern African lions. The two groups probably diverged about 186,000–128,000 years ago. It is thought that the Asiatic lion remained connected to North and Central African lions until gene flow was interrupted due to extinction of lions in Western Eurasia and the Middle East during the Holocene.[24][25]

Asiatic lions are less genetically diverse than African lions, which may be the result of a founder effect in the recent history of the remnant population in the Gir Forest.[26]

Characteristics

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Asiatic lion cub
An Asiatic lioness

The Asiatic lion's fur ranges in colour from ruddy-tawny, heavily speckled with black, to sandy or buffish grey, sometimes with a silvery sheen in certain lighting. Males have only moderate mane growth at the top of the head, so that their ears are always visible. The mane is scanty on the cheeks and throat, where it is only 10 cm (4 in) long. About half of Asiatic lions' skulls from the Gir forest have divided infraorbital foramina, whereas African lions have only one foramen on either side. The sagittal crest is more strongly developed, and the post-orbital area is shorter than in African lions. Skull length in adult males ranges from 330–340 mm (13–13+1⁄2 in), and in females, from 292–302 mm (11+1⁄2–11+7⁄8 in). It differs from the African lion by a larger tail tuft and less inflated auditory bullae.[3] The most striking morphological character of the Asiatic lion is a longitudinal fold of skin running along its belly.[27]

Males have a shoulder height of up to 107–120 cm (42–47 in), and females of 80–107 cm (31+1⁄2–42 in).[28] Two lions in Gir Forest measured 1.98 m (6 ft 6 in) from head to body with a 0.79–0.89 m (31–35 in) long tail of and total lengths of 2.82–2.87 m (9 ft 3 in – 9 ft 5 in). The Gir lion is similar in size to the Central African lion,[3] and smaller than large African lions.[29] An adult male Asiatic lion weighs 160.1 kg (353 lb) on average with the limit being 190 kg (420 lb); a wild female weighs 100 to 130 kg (220 to 285 lb).[30][31][1]

Manes

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Asiatic (above) and southern African (below) lions. Note the larger tail tuft, sparser mane on the head and prominent fold of skin on the abdomen of the former.

Colour and development of manes in male lions varies between regions, among populations and with age of lions.[32] In general, the Asiatic lion differs from the African lion by a less developed mane.[3] The manes of most lions in ancient Greece and Asia Minor were also less developed and did not extend to below the belly, sides or ulnas. Lions with such smaller manes were also known in the Syrian region, Arabian Peninsula and Egypt.[33][34]

Exceptionally sized lions

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The confirmed record total length of a male Asiatic lion is 2.92 m (9 ft 7 in), including the tail.[35] Emperor Jahangir allegedly speared a lion in the 1620s that measured 3.10 m (10 ft 2 in) and weighed 306 kg (675 lb).[36] In 1841, English traveller Austen Henry Layard accompanied hunters in Khuzestan, Iran, and sighted a lion which "had done much damage in the plain of Ram Hormuz," before one of his companions killed it. He described it as being "unusually large and of very dark brown colour", with some parts of its body being almost black.[37] In 1935, a British admiral claimed to have sighted a maneless lion near Quetta in Pakistan. He wrote "It was a large lion, very stocky, light tawny in colour, and I may say that no one of us three had the slightest doubt of what we had seen until, on our arrival at Quetta, many officers expressed doubts as to its identity, or to the possibility of there being a lion in the district."[38]

Distribution and habitat

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Habitat in Gir Forest

In Saurashtra's Gir Forest, an area of 1,412.1 km2 (545.2 sq mi) was declared as a sanctuary for Asiatic lion conservation in 1965. This sanctuary and the surrounding areas are the only habitats supporting the Asiatic lion.[39] After 1965, a national park was established covering an area of 258.71 km2 (99.89 sq mi) where human activity is not allowed. In the surrounding sanctuary only Maldharis have the right to take their livestock for grazing.[40]

Lions inhabit remnant forest habitats in the two hill systems of Gir and Girnar that comprise Gujarat's largest tracts of tropical and subtropical dry broadleaf forests, thorny forest and savanna, and provide valuable habitat for a diverse flora and fauna. Five protected areas currently exist to protect the Asiatic lion: Gir Sanctuary, Gir National Park, Pania Sanctuary, Mitiyala Sanctuary, and Girnar Sanctuary. The first three protected areas form the Gir Conservation Area, a 1,452 km2 (561 sq mi) large forest block that represents the core habitat of the lion population. The other two sanctuaries Mitiyala and Girnar protect satellite areas within dispersal distance of the Gir Conservation Area. An additional sanctuary is being established in the nearby Barda Wildlife Sanctuary to serve as an alternative home for lions.[39] The drier eastern part is vegetated with acacia thorn savanna and receives about 650 mm (26 in) annual rainfall; rainfall in the west is higher at about 1,000 mm (39 in) per year.[30]

The lion population recovered from the brink of extinction to 411 individuals by 2010. In that year, approximately 105 lions lived outside the Gir forest, representing a quarter of the entire lion population. Dispersing sub-adults established new territories outside their natal prides, and as a result the satellite lion population has been increasing since 1995.[39] By 2015, the total population had grown to an estimated 523 individuals, inhabiting an area of 7,000 km2 (2,700 sq mi) in the Saurashtra region., comprising 109 adult males, 201 adult females and 213 cubs.[41][10][42] The Asiatic Lion Census conducted in 2017 revealed about 650 individuals.[43]

By 2020, at least six satellite populations had spread to eight districts in Gujarat and live in human-dominated areas outside the protected area network.[44] 104 lived near the coastline. Lions living along the coast, as well as those between the coastline and the Gir forest, have larger individual ranges.[45]

Former range

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Men with a chained lion in Iran, c. 1880.[46] Photograph by Antoin Sevruguin exhibited in the National Museum of Ethnology (Netherlands)
The Lion Hunt of Ashurbanipal, a sequence of Assyrian palace reliefs, Nineveh, Mesopotamia, 7th century BC

The Asiatic lion used to occur in Arabia, the Levant, Mesopotamia and Baluchistan.[3] In South Caucasia, it was known since the Holocene and became extinct in the 10th century. Until the middle of the 19th century, it survived in regions adjoining Mesopotamia and Syria, and was still sighted in the upper reaches of the Euphrates River in the early 1870s.[33][47] By the late 19th century, it had become extinct in Saudi Arabia and Turkey.[48][49] The last known lion in Iraq was killed on the lower Tigris in 1918.[50]

Historical records in Iran indicate that it ranged from the Khuzestan Plain to Fars Province at elevations below 2,000 m (6,600 ft) in steppe vegetation and pistachio-almond woodlands.[51] It was widespread in the country, but in the 1870s, it was sighted only on the western slopes of the Zagros Mountains, and in the forest regions south of Shiraz.[33] It served as the national emblem and appeared on the country's flag. Some of the country's last lions were sighted in 1941 between Shiraz and Jahrom in Fars Province, and in 1942, a lion was spotted about 65 km (40 mi) northwest of Dezful.[52] In 1944, the corpse of a lioness was found on the banks of the Karun River in Iran's Khuzestan Province.[53][54]

In India, the Asiatic lion occurred in Sind, Bahawalpur, Punjab, Gujarat, Rajasthan, Haryana, Bihar and eastward as far as Palamau and Rewa, Madhya Pradesh in the early 19th century.[55][37] It once ranged to Bangladesh in the east and up to Narmada River in the south.[37] Because of the lion's restricted distribution in India, Reginald Innes Pocock assumed that it arrived from Europe, entering southwestern Asia through Balochistan only recently, before humans started limiting its dispersal in the country. The advent and increasing availability of firearms led to its local extirpation over large areas.[3] Heavy hunting by British colonial officers and Indian rulers caused a steady and marked decline of lion numbers in the country.[40] Lions were exterminated in Palamau by 1814, in Baroda State, Hariana and Ahmedabad district in the 1830s, in Kot Diji and Damoh district in the 1840s. During the Indian Rebellion of 1857, a British officer shot 300 lions. The last lions of Gwalior and Rewah were shot in the 1860s. One lion was killed near Allahabad in 1866.[55] The last lion of Mount Abu in Rajasthan was spotted in 1872.[56] By the late 1870s, lions were extinct in Rajasthan.[37] By 1880, no lion survived in Guna, Deesa and Palanpur districts, and only about a dozen lions were left in Junagadh district. By the turn of the century, the Gir Forest held the only Asiatic lion population in India, which was protected by the Nawab of Junagarh in his private hunting grounds.[3][37]

Ecology and behaviour

[edit]
A male lion scent marking his territory in Gir Forest
Female and male lion
Young male

Male Asiatic lions are solitary, or associate with up to three males, forming a loose pride. Pairs of males rest, hunt and feed together, and display marking behaviour at the same sites. Females associate with up to twelve other females, forming a stronger pride together with their cubs. They share large carcasses among each other, but seldom with males. Female and male lions usually associate only for a few days when mating, but rarely live and feed together.[57][58]

Results of a radio telemetry study indicate that annual home ranges of male lions vary from 144 to 230 km2 (56 to 89 sq mi) in dry and wet seasons. Home ranges of females are smaller, varying between 67 and 85 km2 (26 and 33 sq mi).[59] During hot and dry seasons, they favour densely vegetated and shady riverine habitats, where prey species also congregate.[60][61]

Coalitions of males defend home ranges containing one or more female prides.[62] Together, they hold a territory for a longer time than single lions. Males in coalitions of three to four individuals exhibit a pronounced hierarchy with one male dominating the others.[63]

The lions in Gir National Park are active at twilight and by night, showing a high temporal overlap with sambar (Rusa unicolor), wild boar (Sus scrofa) and nilgai (Boselaphus tragocamelus).[64]

Feeding ecology

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In general, lions prefer large prey species within a weight range of 190 to 550 kg (420 to 1,210 lb), irrespective of their availability.[65] Domestic cattle have historically been a major component of the Asiatic lions' diet in the Gir Forest.[3] Inside Gir Forest National Park, lions predominantly kill chital (Axis axis), sambar deer, nilgai, cattle (Bos taurus), domestic water buffalo (Bubalus bubalis), and less frequently wild boar.[59] They most commonly kill chital, which weighs only around 50 kg (110 lb).[62] They prey on sambar deer when the latter descend from the hills during summer. Outside the protected area where wild prey species do not occur, lions prey on water buffalo and cattle, and rarely on dromedary (Camelus dromedarius). They generally kill most prey less than 100 m (330 ft) away from water bodies, charge prey from close range and drag carcasses into dense cover.[59] They regularly visit specific sites within the protected area to scavenge on dead livestock dumped by Maldhari livestock herders.[66] During dry, hot months, they also prey on mugger crocodiles (Crocodylus palustris) on the banks of Kamleshwar Dam.[54]: 148 

In 1974, the Forest Department estimated the wild ungulate population at 9,650 individuals. In the following decades, the wild ungulate population has grown consistently to 31,490 in 1990 and 64,850 in 2010, including 52,490 chital, 4,440 wild boar, 4,000 sambar, 2,890 nilgai, 740 chinkara (Gazella bennetti), and 290 four-horned antelope (Tetracerus quadricornis). In contrast, populations of domestic buffalo and cattle declined following resettlement, largely due to direct removal of resident livestock from the Gir Conservation Area. The population of 24,250 domestic livestock in the 1970s declined to 12,500 by the mid-1980s, but increased to 23,440 animals in 2010. Following changes in both predator and prey communities, Asiatic lions shifted their predation patterns. Today, very few livestock kills occur within the sanctuary, and instead most occur in peripheral villages. Depredation records indicate that in and around the Gir Forest, lions killed on average 2,023 livestock annually between 2005 and 2009, and an additional 696 individuals in satellite areas.[39]

Dominant males consume about 47% more from kills than their coalition partners. Aggression between partners increases when coalitions are large, but kills are small.[63]

Reproduction

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Lions mating in Gir Forest
A pride of adult lions and their cubs
Asiatic lions mating

Asiatic lions mate foremost between October and November.[67] Mating lasts three to six days. During these days, they usually do not hunt, but only drink water. Gestation lasts about 110 days. Litters comprise one to four cubs.[68] The average interval between births is 24 months, unless cubs die due to infanticide by adult males or because of diseases and injuries. Cubs become independent at the age of about two years. Subadult males leave their natal pride latest at the age of three years and become nomads until they establish their own territory.[58] Dominant males mate more frequently than their coalition partners. During a study carried out between December 2012 and December 2016, three females were observed switching mating partners in favour of the dominant male.[63] Monitoring of more than 70 mating events showed that females mated with males of several rivaling prides that shared their home ranges, and that these males were tolerant toward the same cubs. Only new males that entered the female territories killed unfamiliar cubs. Young females mated foremost with males within their home ranges. Older females selected males at the periphery of their home ranges.[69]

Threats

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A painting of Mughal Emperor Shah Jahan hunting lions in the area of Burhanpur, present-day Madhya Pradesh, from 1630

The Asiatic lion currently exists as a single subpopulation, and is thus vulnerable to extinction from unpredictable events, such as an epidemic or large forest fire. There are indications of poaching incidents in recent years, as well as reports that organized poacher gangs have switched attention from local Bengal tigers to the Gujarat lions. There have also been a number of drowning incidents, after lions fell into wells.[1]

Prior to the resettlement of Maldharis, the Gir forest was heavily degraded and used by livestock, which competed with and restricted the population sizes of native ungulates. Various studies reveal tremendous habitat recovery and increases in wild ungulate populations following the resettlement of Maldharis since the 1970s.[39]

Nearly 25 lions in the vicinity of Gir Forest were found dead in October 2018. Four of them had died because of canine distemper virus, the same virus that had also killed several lions in the Serengeti.[70][71]

Conflicts with humans

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Since the mid 1990s, the Asiatic lion population has increased to an extent that by 2015, about a third resided outside the protected area. Hence, conflict between local residents and wildlife also increased. Local people protect their crops from nilgai, wild boar, and other herbivores by using electrical fences that are powered with high voltage. Some consider the presence of predators a benefit, as they keep the herbivore population in check. But some also fear the lions, and killed several in retaliation for attacks on livestock.[72]

In July 2012, a lion dragged a man from the veranda of his house and killed him about 50–60 km (31–37 miles) from Gir Forest National Park. This was the second attack by a lion in this area, six months after a 25-year-old man was attacked and killed in Dhodadar.[73]

Conservation

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Panthera leo persica was included on CITES Appendix I, and is fully protected in India,[38] where it is considered endangered.[74]

Reintroduction

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Proposed reintroduction sites in India. Pink spots indicate former populations, blue spots indicate proposed sites.

India

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Main article: Asiatic Lion Reintroduction Project

In the 1950s, biologists advised the Indian government to re-establish at least one wild population in the Asiatic lion's former range to ensure the population's reproductive health and to prevent it from being affected by an outbreak of an epidemic. In 1956, the Indian Board for Wildlife accepted a proposal by the Government of Uttar Pradesh to establish a new sanctuary for the envisaged reintroduction, Chandra Prabha Wildlife Sanctuary, covering 96 km2 (37 sq mi) in eastern Uttar Pradesh, where climate, terrain and vegetation is similar to the conditions in the Gir Forest. In 1957, one male and two female wild-caught Asiatic lions were set free in the sanctuary. This population comprised 11 animals in 1965, which all disappeared thereafter.[75]

The Asiatic Lion Reintroduction Project to find an alternative habitat for reintroducing Asiatic lions was pursued in the early 1990s. Biologists from the Wildlife Institute of India assessed several potential translocation sites for their suitability regarding existing prey population and habitat conditions. The Palpur-Kuno Wildlife Sanctuary in northern Madhya Pradesh was ranked as the most promising location, followed by Sita Mata Wildlife Sanctuary and Darrah National Park.[76] Until 2000, 1,100 families from 16 villages had been resettled from the Palpur-Kuno Wildlife Sanctuary, and another 500 families from eight villages were expected to be resettled. With this resettlement scheme the protected area was expanded by 345 km2 (133 sq mi).[75][77]

Gujarat state officials resisted the relocation, since it would make the Gir Sanctuary lose its status as the world's only home of the Asiatic lion. Gujarat raised a number of objections to the proposal, and thus the matter went before the Indian Supreme Court. In April 2013, the Indian Supreme Court ordered the Gujarat state to send some of their Gir lions to Madhya Pradesh to establish a second population there.[78] The court had given wildlife authorities six months to complete the transfer. The number of lions and which ones to be transported will be decided at a later date. As of now, the plan to shift lions to Kuno is in jeopardy, with Madhya Pradesh having apparently given up on acquiring lions from Gujarat.[79][80]

Iran

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See also: Iran's Project Lion

In 1977, Iran attempted to restore its lion population by transporting Gir lions to Arzhan National Park, but the project met resistance from the local population, and thus it was not implemented.[47][52] However, this did not stop Iran from seeking to bring back the lion.[81][82] In February 2019, Tehran Zoological Garden obtained a male Asiatic lion from Bristol Zoo in the United Kingdom,[83] followed in June by a female from Dublin Zoo. There are hopes for them to successfully reproduce.[84]

In captivity

[edit]
Lion in Nawab Wajid Ali Shah Zoological Garden
Asiatic lion in Tierpark Berlin

Until the late 1990s, captive Asiatic lions in Indian zoos were haphazardly interbred with African lions confiscated from circuses, leading to genetic pollution in the captive Asiatic lion stock. Once discovered, this led to the complete shutdown of the European and American endangered species breeding programs for Asiatic lions, as its founder animals were captive-bred Asiatic lions originally imported from India and were ascertained to be intraspecific hybrids of African and Asian lions. In North American zoos, several Indian-African lion crosses were inadvertently bred, and researchers noted that "the fecundity, reproductive success, and spermatozoal development improved dramatically."[85][86]

DNA fingerprinting studies of Asiatic lions have helped in identifying individuals with high genetic variability, which can be used for conservation breeding programs.[87]

In 2006, the Central Zoo Authority of India stopped breeding Indian-African cross lions stating that "hybrid lions have no conservation value and it is not worth to spend resources on them".[85][88] Now only pure native Asiatic lions are bred in India.

In 1972 the Sakkarbaug Zoo sold a pair of young pure-stock lions to the Fauna Preservation Society; which decided they would be accommodated at the Jersey Wildlife Trust where it was hoped to begin a captive breeding programme.[89]

The Asiatic lion International Studbook was initiated in 1977, followed in 1983 by the North American Species Survival Plan (SSP).[90] The North American population of captive Asiatic lions was composed of descendants of five founder lions, three of which were pure Asian and two were African or African-Asian hybrids. The lions kept in the framework of the SSP consisted of animals with high inbreeding coefficients.[27]

In the early 1990s, three European zoos imported pure Asiatic lions from India: London Zoo obtained two pairs; the Zürich Zoologischer Garten one pair; and the Korkeasaari Zoo in Helsinki one male and two females. In 1994, the European Endangered Species Programme (EEP) for Asiatic lions was initiated. The European Association of Zoos and Aquaria (EAZA) published the first European Studbook in 1999. By 2005, there were 80 Asiatic lions kept in the EEP — the only captive population outside of India.[90] As of 2009, more than 100 Asiatic lions were kept within the EEP. The SSP had not resumed; pure-bred Asiatic lions are needed to form a new founder population for breeding in American zoos.[91]

In culture

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The original sandstone sculpted Lion Capital of Ashoka preserved at Sarnath Museum, which was originally erected around 250 BC atop an Ashoka Pillar at Sarnath
Lion depicted in a 15th century Persian manuscript kept at the Topkapi Palace Museum
A painting showing a lion hunt in India, ca. 1810
Further information: Lion (heraldry) and Cultural depictions of lions

South and East Asia

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See also: Bharat Mata and Kangla Sha

Neolithic cave paintings of lions were found in Bhimbetka rock shelters in central India, which are at least 30,000 years old.[92]

The Sanskrit word for 'lion' is 'सिंह' siṃhaḥ, which is also a name of Shiva and signifies the Leo of the Zodiac.[93] The Sanskrit name of Sri Lanka is Sinhala meaning 'Abode of Lions'.[94] Singapore derives its name from the Malay words singa 'lion' and pura 'city', which in turn is from the Sanskrit 'सिंह' siṃhaḥ and पुर pur, latter also meaning 'fortified town'.[93][95]

In Hindu mythology, the half man half lion avatar Narasimha is the fourth incarnation of Vishnu.[96] Simhamukha is a lion-faced protector and dakini in Tibetan Buddhism.[97]

In the 18th book of the Mahabharata, Bharata deprives lions of their prowess.[98] The lion plays a prominent role in The Fables of Pilpay that were translated into Persian, Greek and Hebrew languages between the 8th and 12th centuries.[99] The lion is the symbol of Mahavira, the 24th and last Tirthankara in Jainism.[100][101]

West Asia and Europe

[edit]

Lions are depicted on vases dating to about 2600 BCE that were excavated near Lake Urmia in Iran.[106] The lion was an important symbol in Ancient Iraq and is depicted in a stone relief at Nineveh in the Mesopotamian Plain.[107][108]

See also

[edit]

References

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Further reading

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