Asiatic lion

Asiatic lion
Male
Female
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Felidae
Genus: Panthera
Species: P. leo
Subspecies: P. l. persica
Trinomial name
Panthera leo persica
Meyer, 1826
Current distribution of the Asiatic lion in the wild
Synonyms

P. l. asiaticus, P. l. bengalensis, P. l. indica, P. l. goojratensis[2]

The Asiatic lion (Panthera leo persica), also known as the Indian lion or Persian lion,[3] is a lion subspecies that exists as a single population in India's Gujarat state. It is listed as Endangered by IUCN due its small population size.[1] Since 2010, the lion population in the Gir Forest National Park has steadily increased.[4]

In May 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, comprising 109 adult males, 201 adult females and 213 cubs.[5][6]

The Asiatic lion was first described by the Austrian zoologist Johann N. Meyer under the trinomen Felis leo persicus.[7]

The Asiatic lion is one of five big cat species found in India, along with Bengal tiger, Indian leopard, snow leopard and clouded leopard.[8] It formerly occurred in Southeastern Europe, Black Sea Basin, Caucasus, Persia, Canaan, Mesopotamia, Baluchistan, from Sindh in the west to Bengal in the east, and from Rampur and Rohilkund in the north to Nerbudda in the south. It differs from the African lion by less inflated auditory bullae, a larger tail tuft and a less developed mane.[9]

Characteristics

Adult male Asiatic lion at the Gir Forest.

The most striking morphological character, which is always seen in Asiatic lions, and rarely in African lions, is a longitudinal fold of skin running along its belly.[10] Asiatic lions are slightly smaller than African lions. Adult males weigh 160 to 190 kg (350 to 420 lb), while females weigh 110 to 120 kg (240 to 260 lb).[11] The height at the shoulders is about 3.5 ft (110 cm).[12] The record total length of a male Asiatic lion is 2.92 m (115 in) including the tail.[13]

The 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 with where it is only 4 in (10 cm) long. About half of Asiatic lion 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).[9]

Compared to African lion populations, the Asiatic lion revealed a diminished amount of genetic variation, which may result from a founder effect in the recent history of the remnant population in the Gir Forest.[14]

Distribution and habitat

The Gir National Park and Wildlife Sanctuary in Western Gujarat is the only habitat for the Asiatic lion where an area of 1,412.1 km2 (545.2 sq mi) was declared as a sanctuary for their conservation in 1965. Later, 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.[15]

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.[4] 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.[11]

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.[4]

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.[5]

Former range

The Asiatic lions used to live in West, Southwest, South and Central regions of Asia and in Eastern Europe in historic times. Now the population of the lions currently exists in Western India's Gir Forest National Park. The type specimen of the Asiatic lion was first described from Persia in 1826, followed by descriptions of specimens from Hariana and Basra. Asiatic lions formerly occurred in Persia, Arabia, Palestine, Mesopotamia and Baluchistan.[9]

Panthera leo persica. Sketch by A. M. Komarov[16]

They survived in regions adjoining Mesopotamia and Syria until the middle of the 19th century, and were still sighted in the upper reaches of the Euphrates River in the early 1870s. They were widespread in Iran, but in the 1870s were sighted only on the western slopes of the Zagros Mountains and in the forest regions south of Shiraz.[16]

The advent of firearms led to their extinction over large areas. By the late 19th century, Asiatic lions had been eradicated in Turkey.[17] In Iran, lions 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. In 1963, the last pride of five Persian lions was hunted in the Dasht-i Arzhan of Fars Province. The national newspapers and media "celebrated" the killing of these lions with pictures and fanfare. The diminished pride consisted of a female with four cubs that inhabited a cave. The male had been shot already. The female was shot on the spot, and the cubs were taken as trophies. No subsequent sightings have been reported from Iran.[18]

In India, Asiatic lions once ranged to the state of Bengal, but declined under heavy hunting pressure.[9] In the early 19th century, they were found in north-western and central India in Hariana, Khandesh (in modern-day Maharashtra), Rajasthan, Sindh, and eastward as far as Palamu and Rewa, Madhya Pradesh.[19] Severe hunting by Indian royalties and colonial personnel led to a steady and marked decline of lion numbers in the country.[15] Asiatic lions were exterminated in Palamau by 1814, in Baroda, Haryana and Ahmedabad 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. Until 1880, no lion survived in Guna, Deesa and Palanpur, and only about a dozen lions were left in the Junagadh district. By the turn of the century, they were confined to the Gir Forest and protected by the Nawab of Junagadh in his private hunting grounds.[9]

One lion was killed near Allahabad in 1866.[19] The last lion of Mount Abu was spotted in 1872.

Ecology and behaviour

Male Asiatic lion resting under tree cover.
Comparative illustration of typical Asiatic lion mane types.
Male Asiatic lion after a fight.

Asiatic lions live in prides. Mean pride size, measured by the number of adult females, tends to be smaller than for African lions: most Gir prides contain just two adult females, with the largest having five.[20] 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).[21]

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.[22] Domestic cattle have historically been a major component of the Gir lions’ diet.[9]

In 1974, the Forest Department estimated the wild ungulate population to be 9,650 individuals. This population grew consistently in subsequent surveys, reaching 31,490 in 1990 and 64,850 in 2010, consisting of 52,490 spotted deer, 4,440 wild boar, 4,000 sambar, 2,890 blue bull, 740 chinkara, and 290 four-horned antelope. Thus, in the past four decades, the population of wild ungulates increased by over ten times. 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 resident animals 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 2023 livestock annually between 2005 and 2009, and an additional 696 individuals in satellite areas.[4]

In 2012, an Asiatic lion dragged a man from his house and killed him near the Gir forest, Amreli, India. This was the second attack in the area after a man was attacked and killed in Dhodadar.[23]

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.[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 Maldhari resettlement during the last four decades.[4] Farmers on the periphery of the Gir Forest frequently use crude and illegal electrical fences by powering them with high voltage overhead power lines. These are usually intended to protect their crops from nilgai, but lions and other wildlife are also killed. 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.

Inbreeding

The wild population of Asiatic lions is said to have derived from just a dozen individuals that had survived in the 1880s. After the Nawab of Junagadh established a sanctuary for their protection, the population increased and was roughly estimated at 70 to 200 individuals by 1930.[9] During the first census conducted in 1936 on the basis of pugmarks, the population was estimated at 287 lions in the Junagarh state.[24]

The population was thus thought to be highly inbred, and especially vulnerable to disease.[25] Semen and blood samples collected from 28 wild-caught and captive-bred lions from the Gir forest showed a high incidence (79%) of morphologically abnormal spermatozoa compared to free-ranging African lions, which is nearly always associated with infertility. The Gir lion population may have suffered a drastic population bottleneck or series of bottlenecks followed by inbreeding in their recent history.[10]

In the course of a later study, semen and blood samples were collected from seven lions in three Indian zoos. These samples showed high percentage of motile spermatozoa and low incidence of abnormal spermatozoa, thus implying that inbreeding depression had not affected these animals. The low genetic variability may be a feature of the species and not a result of inbreeding in recent times.[26] However, the RAPD techniques used in this population genetics research have been criticized as being imprecise and having major technical and analytical drawbacks.[27]

Conservation

Panthera leo persica is included on CITES Appendix I, and is fully protected in India.[20]

Reintroduction

The Asiatic Lions at Sanjay Gandhi National Park, Borivali, Mumbai Lion Safari, India

In the 1950s, biologists advised the government to re-establish at least one wild population in the Asiatic lion's former range in order 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 Uttar Pradesh government 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.[28]

The initiative 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.[29] 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).[28]

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.[30] 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

When kept in zoos in colder climates, lions usually develop stronger manes as shown by this male at Chester Zoo, UK.
A captive Asiatic lioness resting and yawning.

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."[31][32]

DNA fingerprinting studies of Asiatic lions have helped in identifying individuals with high genetic variability, which can be used for conservation breeding programs.[33]

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".[32][34] 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).[35] 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.[10]

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.[35]

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.[36]

Evolution

Asiatic lion depicted on a hunting scene (7th century BC, Nineveh)

Fossil remains found in the Cromer Stage suggest that the lion entered Europe with a gigantic form. Frequently encountered lion bones in cave deposits from Eemian times suggest that the late Pleistocene European cave lion, Panthera leo spelaea, survived in the Balkans and Asia Minor. There was probably a continuous population extending into India.[37] 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.[14]

African (above) and Asiatic (below) lions, as illustrated in Johnsons Book of Nature

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. They went extinct without mitochondrial descendants on other continents.[38]

Fossil remains of lions were found in Pleistocene deposits in West Bengal.[39] 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.[40]

Lions 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 the Trans-Caucasus, they were known since the Holocene and became extinct in the 10th century.[16] Pocock suggested that their restricted distribution in India indicated that they were comparatively recent immigrants that came to India through Persia and Baluchistan.[9]

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 center 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.[41]

Taxonomic history

Indian lion male (Panthera leo persica) at Gir Forest National Park

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:[9]

In mythology, religion and art

Hindu Goddess Durga has an Asiatic lion as her vahanam or divine mount
A page from Kelileh o Demneh dated 1429, from Herat, a Persian translation of the ancient Indian Panchatantra.
Emblem of the Hoysala Empire in Ancient India, depicting Sala fighting the Lion.
Dirham coin of Kaykhusraw II, Sivas, AH 638/AD 1240-1
For more details on this topic, see Cultural depictions of lions.

See also

References

  1. 1 2 3 Breitenmoser, U., Mallon, D. P., Ahmad Khan, J. and Driscoll, C. (2008). "Panthera leo ssp. persica". IUCN Red List of Threatened Species. Version 2014.3. International Union for Conservation of Nature.
  2. ↑ Wozencraft, W.C. (2005). "Order Carnivora". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 546. ISBN 978-0-8018-8221-0. OCLC 62265494.
  3. ↑ Humphreys, P., Kahrom, E. (1999). Lion and Gazelle: The Mammals and Birds of Iran. Images Publishing, Avon.
  4. 1 2 3 4 5 Singh, H. S., Gibson, L. (2011). "A conservation success story in the otherwise dire megafauna extinction crisis: The Asiatic lion (Panthera leo persica) of Gir forest" (PDF). Biological Conservation 144 (5): 1753–1757. doi:10.1016/j.biocon.2011.02.009.
  5. 1 2 DeshGujarat (2015). "Asiatic Lion population up from 411 to 523 in five years". Retrieved 10 May 2015.
  6. ↑ Anonymous (2015). "Asiatic lion population in Gujarat rises to 523". Deccan Herald.
  7. ↑ Meyer, J. N. (1826). Dissertatio inauguralis anatomico-medica de genere felium. Doctoral thesis, University of Vienna.
  8. ↑ Pandit, M. W., Shivaji, S., Singh, L. (2007). You Deserve, We Conserve: A Biotechnological Approach to Wildlife Conservation. I. K. International Publishing House Pvt. Ltd., New Delhi. ISBN 9788189866242.
  9. 1 2 3 4 5 6 7 8 9 Pocock, R. I. (1939). The Fauna of British India, including Ceylon and Burma. Mammalia. – Volume 1. Taylor and Francis Ltd., London. Pp. 212–222.
  10. 1 2 3 O’Brien, S. J., Joslin, P., Smith, G. L. III, Wolfe, R., Schaffer, N., Heath, E., Ott-Joslin, J., Rawal, P. P., Bhattacharjee, K. K., and Martenson, J. S. (1987). "Evidence for African origins of founders of the Asiatic lion Species Survival Plan" (PDF). Zoo Biology 6 (2): 99–116. doi:10.1002/zoo.1430060202.
  11. 1 2 Chellam, R. and A. J. T. Johnsingh. (1993). Management of Asiatic lions in the Gir Forest, India. In N. Dunstone and M. L. Gorman (eds.) Mammals as predators: the proceedings of a symposium held by the Zoological Society of London and the Mammal Society, London. Volume 65 of Symposia of the Zoological Society of London. Zoological Society of London, London. Pp. 409–423.
  12. ↑ Sterndale, R. A. (1884). Natural History of the Mammalia of India and Ceylon. Thacker, Spink and Co., Calcutta.
  13. ↑ Sinha, S. P. (1987). Ecology of wildlife with special reference to the lion (Panthera leo persica) in Gir Wildlife Sanctuary, Saurashtra, Gujurat. Ph.D. thesis, Saurashtra University, Rajkot ISBN 3844305459.
  14. 1 2 O’Brien, S. J., Martenson, J. S., Packer, C., Herbst, L., de Vos, V., Joslin, P., Ott-Joslin, J., Wildt, D. E. and Bush, M. (1987). "Biochemical genetic variation in geographic isolates of African and Asiatic lions" (PDF). National Geographic Research 3 (1): 114–124.
  15. 1 2 Varma, K. (2009). "The Asiatic Lion and the Maldharis of Gir Forest: An Assessment of Indian Eco-Development" (PDF). The Journal of Environment Development 18 (2): 154–176. doi:10.1177/1070496508329352.
  16. 1 2 3 Geptner, V. G., Sludskij, A. A. (1972). Mlekopitajuščie Sovetskogo Soiuza. Vysšaia Škola, Moskva. (In Russian; English translation: Heptner, V.G., Sludskii, A. A., Komarov, A., Komorov, N.; Hoffmann, R. S. (1992). Mammals of the Soviet Union. Vol III: Carnivores (Feloidea). Smithsonian Institution and the National Science Foundation, Washington DC).
  17. ↑ Üstay, A. H. (1990). Hunting in Turkey. Istanbul: BBA.
  18. ↑ Guggisberg, C. A. W. (1961). Simba: The Life of the Lion. Cape Town: Howard Timmins.
  19. 1 2 Blanford, W. T. (1889). The Fauna of British India, including Ceylon and Burma. Mammalia. Taylor and Francis, London.
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  21. ↑ Johnsingh, A.J.T. and R. Chellam. (1991). Asiatic lions. pp. 92–93 in: Seidensticker, J., Lumpkin, S. and F. Knight. (eds.) Great Cats. London, Merehurst.
  22. ↑ Hayward, M. W. and G. I. H. Kerley (2005). "Prey preferences of the lion (Panthera leo)" (PDF). Journal of Zoology 267 (3): 309–322. doi:10.1017/S0952836905007508.
  23. ↑ "Man-eater lion kills 50-year-old in Amreli, preys on him". dna. 20 July 2012. Retrieved March 28, 2015.
  24. ↑ Mitra, S. (2005). Gir Forest and the Saga of the Asiatic Lion. Indus Publishing Company, New Delhi.
  25. ↑ Klum, M. (2001). "Extinction stalks the Asiatic lion, a regal subspecies now crowded into a single sanctuary in India’s Gir Forest". National Geographic Society.
  26. ↑ Shivaji, S., Jayaprakash, D. and Patil, S. B. (1998). "Assessment of inbreeding depression in big cats: Testosterone levels and semen analysis" (PDF). Current Science 75 (9): 923–930. html version
  27. ↑ Sunnucks, P., Meglécz, E. (2000). "Efficient genetic markers for population biology". Trends in Ecology and Evolution 15 (9): 376–377. doi:10.1016/S0169-5347(00)01825-5. PMID 10931678.
  28. 1 2 Johnsingh, A.J.T. (2006). "Kuno Wildlife Sanctuary ready to play second home to Asiatic lions?". Field Days: A Naturalist's Journey Through South and Southeast Asia. Hyderabad: Universities Press. pp. 126–138. ISBN 8173715521.
  29. ↑ Walker, S. (1994). Executive summary of the Asiatic lion PHVA. First draft report. Zoo’s Print: 2–22.
  30. ↑ Anand, U. (2013). Supreme Court gives Madhya Pradesh lions' share from Gujarat's Gir. The Indian Express Ltd., 17 April 2013.
  31. ↑ Avise, J. C.; Hamrick, J. L. (1996). Conservation Genetics. Springer Science & Business Media. p. 67. ISBN 9780412055812. Furthermore, when Asiatic lions were inadvertently bred to African lion subspecies in North America, the fecundity, reproductive success, and spermatozoal development improved dramatically.
  32. 1 2 Tudge, C. (2011). Engineer In The Garden. Random House. p. 42. ISBN 9781446466988. But when Asian lions and African lions are brought together in zoos, the resulting hybrids are indeed 'fully viable', and in fact, as we will see again in Chapter 8, the first serious attempts to conserve Asian lions in zoos were spoiled because Africans were allowed to cross-breed with them (and the breeding programme had to begin all over again).
  33. ↑ Shankaranarayanan, P., Banerjee, M., Kacker, R. K., Aggarwal, R. K. and Singh, L. (1997). Genetic variation in Asiatic lions and Indian tigers. Electrophoresis 18 (9): 1693–1700. doi:10.1002/elps.1150180938
  34. ↑ "Hybrid lions at Chhatbir Zoo in danger". The Times of India. 18 September 2006. Retrieved 30 September 2014. Hybrid lions at Chhatbir Zoo 23 in all may not have a rip-roaring time in days ahead; they face a bleak future. Authorities have stopped their breeding since "hybrid lions have no conservation value and it is not worth to spend resources on them". The zoo, probably the only one in north India to offer a lion safari, has only these lions who have been crossbred from Asiatic and African lions. These hybrid lions will be replaced by pure Asiatic lions. According to zoo officials, the Central Zoo Authority is focusing on the conservation of pure Asiatic lions.
  35. 1 2 Zingg, R. (2007). Asiatic Lion Studbooks: a short history. Zoos' Print Journal XXII (6): 4.
  36. ↑ "The Asiatic lion captive breeding programme". Archived from the original on 2009-02-06.
  37. ↑ Kurtén, B. (1968). Pleistocene Mammals of Europe. Transaction Publishers, 2007. p. 317. ISBN 0202309533.
  38. ↑ Burger, J., Rosendahl, F., Loreille, O., Hemmer, H., Eriksson, T., Götherström, A., Hiller, J., Collins, M. J., Wess, T., Alt, K. W. (2004). "Molecular phylogeny of the extinct cave lion Panthera leo spelaea". Molecular Phylogenetics and Evolution 30 (3): 841–849. doi:10.1016/j.ympev.2003.07.020. PMID 15012963.
  39. ↑ Dutta, A. K. (1976). "Occurrence of fossil lion and spotted hyena from Pleistocene deposits of Susunia, Bankura District, West Bengal". Journal of the Geological Society of India 17 (3): 386–391.
  40. ↑ Manamendra-Arachchi, K.; Pethiyagoda, R.; Dissanayake, R.; Meegaskumbura, M. (2005). "A second extinct big cat from the late Quaternary of Sri Lanka" (PDF). The Raffles Bulletin of Zoology (Supplement 12): 423–434.
  41. ↑ Barnett, R.; Yamaguchi, N.; Barnes, I.; Cooper, A. (2006). "The origin, current diversity and future conservation of the modern lion (Panthera leo)". Proceedings of the Royal Society B 273 (1598): 2119–2125. doi:10.1098/rspb.2006.3555. PMC 1635511. PMID 16901830.
  42. ↑ Bennett, E. T. (1829). The Tower Menagerie, Comprising the Natural History of the Animals Contained in That Establishment; With Anecdotes of Their Characters and History. Printed for Robert Jennings, London.
  43. ↑ Smee, W. (1833). Felis leo, Linn., Var. goojratensis. Proceedings of the Zoological Society of London, Part I (December 1833): 140.
  44. ↑ Jardine, W. (1834). The Lion. In: Natural History of the Felinae. Series: Naturalist's library. H. G. Bohn, London.
  45. ↑ Blainville, H. M. D. (1843). Felis. Plate VI. in: Ostéographie, ou Description iconographique comparée du squelette et du système dentaire des mammifères récents et fossiles pour servir de base à la zoologie et à la géologie. J.B. Ballière et fils, Paris.
  46. ↑ Apte, V. S. (1957–1959). सिंहः siṃhḥ. In: Revised and enlarged edition of Prin. V. S. Apte's The practical Sanskrit-English dictionary. Prasad Prakashan, Poona.
  47. ↑ "Simhamukha". Himalayanart.org. Archived from the original on 25 December 2010. Retrieved 2010-12-14.
  48. ↑ Turner, R. L. (Ralph Lilley), Sir. A comparative dictionary of Indo-Aryan languages. London: Oxford University Press, 1962-1966. Includes three supplements, published 1969-1985.
  49. ↑ McCleod, W. H. (1989). The Sikhs: History, Religion, and Society. Columbia University Press, New York
  50. ↑ Singh, K. (1963). A History of the Sikhs. Volume I. Princeton University Press, Princeton, New Jersey
  51. ↑ The Wisdom Library: The Mahavamsa. The Great Chronicle of Sri Lanka. Chapter 34 − The Eleven Kings
  52. ↑ The American Heritage Dictionary of the English Language (2000). "Singapore" (Fourth ed.). Houghton Mifflin Company.
  53. ↑ "Early History". Ministry of Information, Communications and the Arts, Singapore. Archived from the original on 27 April 2006. Retrieved 2006-04-14.
  54. ↑ Goswamy, B. N. (2002). "Where does the Lion come from in ancient Chinese culture? Celebrating with the Lion Dance". The Tribune Newspaper, Chandigarh, India. Retrieved 2010-12-14.

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