Asiatic lion

Taxonomic history

Following Meyer's first description of an Asiatic lion skin from Persia, other naturalists and zoologists also described lions from other parts of Asia that today are all considered synonyms of P. l. persica:

Evolution

Fossil remains found in the Cromer Stage suggest that the lion or a lion-like animal that entered Europe was of a gigantic size. Frequently encountered lion bones in cave deposits from Eemian times suggest that the late Pleistocene Eurasian cave lion Panthera leo spelaea survived in the Balkans and Asia Minor. There was probably a continuous population extending into India. Cave lions appeared about 600,000 years ago and were distributed throughout Europe, across Siberia and into western Alaska. The gradual formation of dense forest likely caused the decline in geographic range of lions near the end of the late Pleistocene. Phylogenetic analysis of cave lion DNA samples showed that they were highly distinct from their living relatives, and represent lineages that were isolated from lions in Africa and Asia ever since their dispersal over Europe in prehistoric times, and became extinct without mitochondrial descendants on other continents.

Fossil remains of lions were found in Pleistocene deposits in West Bengal. A fossil carnassial found in the Batadomba Cave indicates that Panthera leo sinhaleyus inhabited Sri Lanka during the late Pleistocene, and is thought to have become extinct around 39,000 years ago. This subspecies was described by Deraniyagala in 1939. It is distinct from the extant Asiatic lion.

Modern lions

A phylogeographic analysis based on mtDNA sequences of lions from across their entire range 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 eastern–Southern Africa, from where lions migrated 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. Natural barriers to lion dispersal comprise the Sahara Desert, equatorial rainforests and the Great Rift Valley.

In a study about lion evolution, Genetic markers of 357 samples from captive and wild lions from Africa and India were examined. Results suggest four lineages of lion populations: one from Kenya, one from Southern Africa, one from Central and Northern Africa to Asia, and one from Southern and Eastern Africa. The authors conclude that the first wave of lion expansion occurred about 118,000 years ago from Eastern Africa into West Asia, and the second wave at the transition of Pleistocene and Holocene periods from Southern Africa towards East Africa. Lions in Southern and East Africa are genetically different from lions in West and Central Africa, which are more closely related to the Asiatic lion.

Characteristics

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 lights. 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 (3.9 in) long. About half of Asiatic lions' skulls from the Gir forest have divided infraorbital foramina, whereas in African lions, there is only one foramen on either side. The sagittal crest is more strongly developed, and the post-orbital area is shorter than in African lion. Skull length in adult males ranges from 330 to 340 mm (13 to 13 in), and in females from 292 to 302 mm (11.5 to 11.9 in). The most striking morphological character of the Asiatic lion is a longitudinal fold of skin running along its belly.

After the tiger, the Asiatic lion is the second biggest extant cat in the wilderness of India and Eurasia. Adult males weigh 160 to 190 kg (350 to 420 lb), while females weigh 110 to 120 kg (240 to 260 lb). Shoulder height is about 1.10 m (43 in).

Recorded flesh measurements of two lions in Gir Forest were head-and-body measurements of 1.98 m (78 in) each, with tail-lengths of 0.79–0.89 m (31–35 in) and total lengths of 2.82–2.87 m (111–113 in), respectively. The Gir lion is similar in size to the Central African lion, and smaller than large African lions.

Compared to populations of African lions, the Asiatic lion has less genetic variation, which may result from a founder effect in the recent history of the remnant population in the Gir Forest.

Exceptionally sized lions

The record total length of a male Indian lion is 2.92 m (115 in), including the tail.

During the year 1841, in Khuzestan, Iran, Austen Henry Layard, accompanied by hunters from Luristan, saw 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 it body being almost black.

Though the last lion of what is now Pakistan was thought to have been killed near Kot Diji in Sindh Province in 1810, a British Admiral, while traveling on a train accompanied by two others, reportedly saw a maneless lion eating a goat near Quetta in 1935. 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."

Distribution and habitat

In the 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 in Saurashtra, Western India, are the only wild habitats supporting the Asiatic lion. After 1965, a national park covering an area of 258.71 km² (99.89 sq mi) was established where no human activity is allowed. In the surrounding sanctuary only Maldharis have the right to graze their livestock.

The population recovered from the brink of extinction to 411 individuals in 2010. Lions occupy remnant forest habitats in the two hill systems of Gir and Girnar that comprise Gujarat's largest tracts of dry deciduous forest, 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) forest block that represents the core habitat of the Asiatic lions. 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 forest to serve as an alternative home for Gir lions. 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.

As of 2010, approximately 105 lions, comprising 35 males, 35 females, 19 subadults, and 16 cubs existed outside the Gir forest, representing a full quarter of the entire lion population. The increase in satellite lion populations may represent the saturation of the lion population in the Gir forest and subsequent dispersal by sub-adults compelled to search for new territories outside their natal pride. Over the past two decades, these satellite areas became established, self-sustaining populations as evidenced by the presence of cubs since 1995.

As of May 2015, the lion population was estimated at 523 individuals, comprising 268 individuals in the Junagadh district, 44 in the Gir Somnath District, 174 in the Amreli District and 37 in the Bhavnagar District.

Former range

The Asiatic lion used to live in Eastern Europe, West, Central and South Asia in historic times. The type specimen of the Asiatic lion was first described from Persia in 1826, followed by descriptions of specimens from Hariana and Basra. It also occurred in Arabia, Palestine, Mesopotamia and Baluchistan.

It inhabited the southern part of the Balkan peninsula up to Macedonia and probably the Danube River, but disappeared in Greece around the first century. 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. It was widespread in Iran, but in the 1870s, it was sighted only on the western slopes of the Zagros Mountains and in the forest regions south of Shiraz.

Reginald Innes Pocock suggested that the restricted distribution of the Asiatic lion in India, compared to that of the tiger, indicated that it was a comparatively recent immigrant that arrived in the country through Persia and Baluchistan, before humans limited its dispersal. In the early 19th century, the Asiatic lion occurred in Sind, Bahawalpur, Punjab, Gujarat, Rajastan, Hariana, Bihar and eastward as far as Palamau and Rewa, Madhya Pradesh. It once ranged to Bengal in the east and up to the Narmada River in the south, but declined under heavy hunting pressure. The advent and increasing availability of firearms led to its extinction over large areas.

Heavy hunting by British colonial officers and Indian rulers led to a steady and marked decline of lion numbers in the country. Lions were exterminated in Palamau by 1814, in Baroda, Hariana and Ahmedabad district in the 1830s, in Kot Diji and Damoh 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. The last lion of Mount Abu, in what is now Rajasthan, was spotted in 1872. By the late 1870s, lions were extinct in Rajastan. By 1880, no lion survived in Guna, Deesa and Palanpur, and only about a dozen lions were left in Junagadh district. By the turn of the century, the Asiatic lion was confined to the Gir Forest and protected by the Nawab of Junagadh in his private hunting grounds.

By the late 19th century, the Asiatic lion had become extinct in modern-day Turkey. In Iran, the lion served as the national emblem and appeared on the country's flag. Some of the last lions were sighted in 1941 between Shiraz and Jahrom in the Fars Province. In 1944, the corpse of a lioness was found on the banks of the Karun River in Iran's Khuzestan Province.

Ecology and behaviour

Asiatic lions live in prides. Mean pride size, measured by the number of adult females, tends to be smaller than for those in Africa lions, apart perhaps from those in Western and Central Africa. Most Gir prides contain just two adult females, with the largest having five. Coalitions of males defend home ranges containing one or more groups of females; but, unlike African lions, Gir males generally associate with their pride females only when mating or on a large kill. A lesser degree of sociability in the Gir lions may be a function of the smaller prey available to them: the most commonly taken species (45% of known kills), the chital, weighs only around 50 kg (110 lb).

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. Yet they predominately take prey substantially smaller than this, reflecting their opportunistic hunting behaviour. Within this range, they prefer species that weigh 350 kg (770 lb), which is much larger than the largest recorded weight of lion. The group hunting strategy of lions enables exceptionally large prey items to be taken. Hunting success in lions is influenced by hunting-group size and composition, the hunting method used and by environmental factors such as grass and shrub cover, time of day, moon presence and terrain. Domestic cattle have historically been a major component of the Gir lions' diet.

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 spotted deer, 4,440 wild boar, 4,000 sambar, 2,890 blue bull, 740 chinkara, and 290 four-horned antelope. 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 in 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. In and around the Gir forest, depredation records indicate that lions killed on average 2,023 livestock annually between 2005 and 2009, and an additional 696 individuals in satellite areas.

Sympatric carnivores

The Gir forest and surrounding landscapes are home for the Indian leopard and the striped hyena. The presence of jungle cat, Asiatic wildcat and rusty-spotted cat has also been reported. The golden jackal scavenges on carcasses of large herbivores, and preys on chital fawn and Indian hares.

Asiatic lion versus tiger

In the 19th century, clashes between Asiatic lions and tigers in the wilderness of India were reported. Just as the lion co-existed with the Bengal tiger in parts of India, it occurred in areas inhabited by the Caspian tiger, like northern Iraq, northern Persia and the Trans-Caucasus, before humans extirpated either lions or tigers in these regions.

Today, the Asiatic lion does not share its range with the tiger, but both inhabit the ecoregion of Kathiawar-Gir dry deciduous forests. Though the Asiatic lion's population has grown to the extent that it is no longer confined to the Gir Forest, it is still restricted to the Kathiawar Peninsula in Saurashtra, and the nearest place with the presence of Bengal tigers is the border-triangle of Gujarat, Maharashtra and Madhya Pradesh. If the project to move some lions to Palpur-Kuno Wildlife Sanctuary is successful, then the lion and tiger could once again coexist in the wilderness of South Asia, due to there being tigers in that sanctuary.

Threats

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. There are reports that organized gangs have switched attention from tigers to these lions. There have also been a number of drowning incidents after lions fell into wells.

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 Maldhari resettlement during the last four decades.

Conflicts with humans

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 pigs and other herbivores by using electrical fences that are powered with high voltage. Some consider the presence of predators a benefit, as latter keep the herbivore population in check. But some people also fear the lions and killed several in retaliation for attacks on livestock.

Nearly 20,000 open wells dug by farmers in the area for irrigation have also acted as traps, which led to many lions drowning. To counteract the problem, suggestions for walls around the wells, as well as the use of "drilled tube wells" have been made.

In July 2012, a lion dragged a man from the veranda of his house and killed him about 50–60 km (31–37 mi) from the 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.

Conservation

Panthera leo persica is included on CITES Appendix I, and is fully protected in India.

Reintroduction

In the 1950s, biologists advised the 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 : the Chandraprabha 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.

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 the Sita Mata Wildlife Sanctuary and the Darrah National Park. Until 2000, 1,100 families from 16 villages had been resettled from the Palpur-Kuno Wildlife Sanctuary, and another 500 families from eight villages envisaged to be resettled. With this resettlement scheme the protected area was expanded by 345 km² (133 sq mi).

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 has raised a number of objections to the proposal, and the matter is now 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. The court has 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.

In captivity

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."

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

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". Now only pure native Asiatic lions are bred in India.

The Asiatic lion International Studbook was initiated in 1977, followed in 1983 by the North American Species Survival Plan (SSP). 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.

In the early 1990s, three European zoos imported pure Asiatic lions from India: the London Zoo obtained two pairs; the Zürich Zoologischer Garten one pair; and the Helsinki Zoo 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.

There are now over 100 Asiatic lions in the EEP. The SSP did not yet resume; pure-bred Asiatic lions are needed to form a new founder population for breeding in American zoos.

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