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Temporal range: Early Pleistocene tae recent
Lion waiting in Namibia.jpg
Okonjima Lioness.jpg
Female (lioness)
Conservation status
Scienteefic clessification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Family: Felidae
Genus: Panthera
Species: P. leo
Binomial name
Panthera leo
(Linnaeus, 1758)[3]
Lion distribution.png
Map Guj Nat Parks Sanctuary.png
Distribution o lions in Indie: The Gir Wid, in Gujarat, is the last naitural range o aboot 400 wild Asiatic lions. There are plans tae reintroduce some lions tae Kuno Wildlife Sanctuary in neighbouring Madhya Pradesh.
Felis leo
Linnaeus, 1758[3]

The lion (Panthera leo) is ane o the five big cat species in the genus Panthera an a memmer o the faimily Felidae. Wi some males ootdingin 250 kg (550 lb) in wicht,[4] it is the seicont-mucklest livin cat efter the tiger. Thir days, thare wild lions in sub-Saharan Africae an in Asie, wi an endangered remnant population in Gir Wid Naitional Pairk in Indie, haein disappeared frae North Africae an Soothwast Asie in heestoric times. Till the late Pleistocene, aboot 10,000 years ago, the lion wis the maist widespread muckle laund mammal efter humans. Thay wur foond in maist o Africae, athort Eurasie frae wastren Europe tae Indie, an in the Americas frae the Yukon tae Peru.[5] The lion is a vulnerable species, haein seen a major population decline o 30–50% ower the past seicont hauf o the 20t century.[2] Lion populations are untenable ootwi designatit reserves an naitional pairks. Awbesit the cause o the decline isnae fully unnerstuid, habitat loss an conflicts wi humans are the greatest causes o concern thir days. Athin Africae, the Wast African lion population is pairticularly endangered.

Lions live for 10–14 years in the wild, awbesit in captivity thay can live mair nor 20 years. In the wild, males seldom live langer than 10 years, as injuries sustained frae continual fechtin wi rival males greatly reduce thair longevity.[6] Thay teepically inhabit savanna an grassland, awbesit thay mey tak tae bush an wid. Lions are unuisually social compared tae ither cats. A pride o lions consists o relatit females an affspring an a smaw nummer o adult males. Groups o female lions teepically hunt thegither, preying maistly on lairge ungulates. Lions are apex an keystone predators, awbesit thay are an aa expert scavengers obtainin ower 50 percent o thair fuid bi scavengin as opportunity allows. While lions dae nae teepically hunt humans, some hae been kent tae dae sae. Sleeping mainly during the day, lions are primarily nocturnal, awbesit bordering on crepuscular in naitur.[7][8]

Heichly distinctive, the male lion is easy recognised bi its mane, an its face is ane o the widest recognised ainimal seembols in human cultur. Depictions hae existit frae the Upper Paleolithic period, wi carvins an pentins frae the Lascaux an Chauvet Caves, throu virtually aw auncient an medieval culturs whaur thay ance occurred. It haes been extensively depicted in sculpturs, in pentins, on naitional banners, an in contemporary films an leeteratur. Lions hae been kept in menageries syne the time o the Roman Empire, an hae been a key species sought for exhibeetion in zoos ower the warld syne the late aichteent century. Zoos are cooperatin warldwide in breedin programs for the endangered Asiatic subspecies.

Etymology[eedit | eedit soorce]

The lion's name, seemilar in mony Romance leids, is derived frae the Laitin leo,[9] an the Ancient Greek λέων (leon).[10] The Hebrew wird לָבִיא (lavi) mey an aa be relatit.[11] It wis ane o the species oreeginally describit bi Linnaeus, wha gae it the name Felis leo, in his aichteent-century wirk, Systema Naturae.[3]

Characteristics[eedit | eedit soorce]

A skeletal moont o an African Lion attackin a Common Eland on display at The Museum o Osteology, Oklahoma City, Oklahoma

Ahind anerly the trrger, the lion is the seicont mucklemaist livin felid in lenth an wecht. Its skull is verra seemilar tae that o the teeger, awbesit the frontal region is uisually mair depressed an flattened, wi a slichtly shorter postorbital region. The lion's skull haes broader nasal openings than the tiger, housamiver, due tae the amoont o skull variation in the twa species, uisually, anly the structur o the lawer jaw can be uised as a reliable indicator o species.[12] Lion colouration varies frae licht buff tae yellowish, reiddish, or dark ochraceous broun. The unnerpairts are generally lichter an the tail tuft is black. Lion cubs are born wi broun rosettes (spots) on thair body, rather lik those o a leopard. awbesit thir spots fade as lions reach adulthuid, faint spots eften mey still be seen on the shanks an unnerpairts, parteecularly on lionesses.

Lions are the anerly members o the cat faimily tae display obvious sexual dimorphism – that is, males an females leuk distinctly different. Thay an aa hae specialised roles that ilka gender plays in the pride. For instance, the lioness, the hunter, lacks the male's thick mane. The colour o the male's mane varies frae blond tae black, generally acomin darker as the lion grows aulder. The maist distinctive characteristic shared bi baith females an males is that the tail ends in a hairy tuft. In some lions, the tuft conceals a haurd "spine" or "spur", approximately 5 mm lang, formed o the final sections o tail bane fused thegither. The lion is the anly felid tae hae a tufted tail – the function o the tuft an spine are unkent. Absent at birth, the tuft develops aroond 5½ months o age an is readily identifiable at 7 months.[13]

The size o adult lions varies athort thair range wi those frae the soothren African populations in Zimbabwe, the Kalahari an Kruger Pairk averagin aroond 189.6 kg (418 lb) an 126.9 kg (280 lb) in males an females respectively compared tae 174.9 kg (386 lb) an 119.5 kg (263 lb) o male an female lions frae East Africae.[14] Reportit body measurements in males are heid-body lenths rangin frae 170 tae 250 cm (5 ft 7 in tae 8 ft 2 in), tail lenths o 90–105 cm (2 ft 11 in–3 ft 5 in). In females reportit heid-body lenths range frae 140 tae 175 cm (4 ft 7 in tae 5 ft 9 in), tail lenths o 70–100 cm (2 ft 4 in–3 ft 3 in),[4] houiver, the aften citit maximum heid an body lenth o 250 cm (8 ft 2 in) fits rather tae extinct Pleistocene forms, lik the American lion, wi even lairge modren lions meisurin several centimetres less in lenth.[15] Record measurements frae huntin records are supposedly a tot lenth o naur 3.6 m (12 ft) for a male shot naur Mucsso, soothren Angolae in October 1973 an a wecht o 313 kg (690 lb) for a male shot ootside Hectorspruit in eastren Transvaal, Sooth Africae in 1936.[16] Anither notably outsized male lion, that wis shot naur Moont Kenyae, wiched in at 272 kg (600 lb).[17]

Mane[eedit | eedit soorce]

During agonistic confrontations wi ither lions, the mane maks the lion appear lairger.

The mane o the adult male lion, unique amang cats, is ane o the maist distinctive characteristics o the species. It mey provide an excellent intimidation display; aidin the lion during confrontations wi ither lions.[18] The presence, absence, colour, an size o the mane is associatit wi genetic precondeetion, sexual maturity, climate, an testosterone production; the rule o thumb is the darker an fuller the mane, the healthier the lion. Sexual selection o mates bi lionesses favors males wi the densest, darkest mane.[19] Research in Tanzanie an aa suggests mane lenth signals fechtin success in male–male relationships. Darker-maned individuals mey hae langer reproductive lives an higher affspring survival, awbesit thay suffer in the hettest months o the year.[20]

A maneless lion.

Scientists ance believed that the distinct status o some subspecies could be juistified bi morphology, includin the size o the mane. Morphology wis uised tae identify subspecies lik the Barbary lion an Cape lion. Research haes suggestit, houiver, that environmental factors influence the colour an size o a lion's mane, lik the ambient temperatur.[20] The cuiler ambient temperatur in European an North American zoos, for example, mey result in a hivier mane. Thus the mane is nae an appropriate merker for identifyin subspecies.[21][22] The males o the Asiatic subspecies, houiver, are characterised bi sparser manes than average African lions.[23]

In the Pendjari Naitional Pairk aurie awmaist aw males are maneless or hae verra weak manes.[24] Maneless male lions hae an aa been reportit frae Senegal, frae Sudan (Dinder Naitional Pairk), an frae Tsavo East Naitional Pairk in Kenyae, an the oreeginal male white lion frae Timbavati an aa wis maneless. The testosterone hormone haes been linked tae mane growth, therefore castratit lions eften hae minimal tae no mane, as the removal o the gonads inhibits testosterone production.[25]

Cave pentins o extinct European cave lions awmaist exclusively shaw ainimals wi no manes, suggestin that thay war aither athoot manes,[26] or that the pentins depict lionesses as seen huntin in a group.

White lions[eedit | eedit soorce]

White lions owe thair colouring tae a recessive allele; thay are rare forms o the subspecies Panthera leo krugeri

The white lion is nae a distinct subspecies, but a special morph wi a genetic condeetion, leucism,[27]> that causes paler colouration akin tae that o the white teeger; the condeetion is seemilar tae melanism, that causes black panthers. Thay are nae albinos, haein normal pigmentation in the ees an skin. White Transvaal lion (Panthera leo krugeri) individuals occasionally hae been encountered in an aroond Kruger Naitional Pairk an the adjacent Timbavati Private Game Reserve in eastren Sooth Africae, but are mair commonly foond in captivity, whaur breeders deliberately select them. The unuisual cream colour o thair coats is due tae a recessive allele.[28] Reportedly, thay hae been bred in camps in South Africae for uise as trophies tae be killt during canned hunts.[29]

Behaviour[eedit | eedit soorce]

Adult male lion stretching.

Lions spend much o thair time restin an are inactive for aboot 20 oors per day.[30] awbesit lions can be active at ony time, thair activity generally peaks efter dusk wi a period o socialisin, gruimin, an defecatin. Intermittent bursts o activity follae throu the nicht oors till dawn, whan huntin maist eften taks place. Thay spend an average o twa oors a day walkin an 50 minutes eatin.[31]

Group organization[eedit | eedit soorce]

Twa lionesses an a matur male o a pride.
Twa males an a female lion.

Lions are the maist socially inclined o aw wild felids, maist o thaim remainin quite solitar in naitur. The lion is a predatory carnivore wi twa teeps o social organisation. Some lions are residents, livin in groups centerin aroond relatit lionesses, cried prides.[32] Females form the stable social unit in a pride an dae nae tolerate ootside females.[33] Membership anly chynges wi the births an daiths o lionesses,[34] awbesit some females dae leave an acome nomadic.[35] awbesit extremely lairge prides, consistin o up tae 30 individuals, hae been observed, the average pride consists o five or sax females, thair cubs o baith sexes, an ane or twa males (kent as a coaleetion if mair nor ane) wha mate wi the adult females. The nummer o adult males in a coaleetion is uisually twa but mey increase tae as mony as fower afore decreasin again ower time.[35] The sole exception tae this pattern is the Tsavo lion pride that aye haes juist ane adult male.[36] Male cubs are excluded frae thair maternal pride whan thay reach maturity at aroond 2–3 years o age.[35] The seicont organisational behaviour is labeled nomads, wha range widely an muive aboot sporadically, either singularly or in pairs.[32] Pairs are mair frequent amang relatit males wha hae been excludit frae thair birth pride. Note that a lion mey switch lifestyles; nomads mey acome residents an vice versa. Males, as a rule, live at least some portion o thair lives as nomads, an some can never jyne anither pride. A female wha becomes a nomad haes much greater difficulty joining a new pride, as the females in a pride are relatit, an thay reject maist attempts bi an unrelated female tae join thair faimily group.

The aurie a pride occupies is cried a pride aurie, whauras that bi a nomad is a range.[32] The males associated wi a pride tend tae stay on the fringes, patrollin thair territory. Why sociality – the maist pronoonced in ony cat species – haes developed in lionesses is the subject o much debate. Increased huntin success appears an obvious reason, but this is less nor sure upon examination: coordinated huntin daes allou for mair successfu predation but an aa ensurs that non-huntin members reduce per capita calorific intake; houiver, some tak a role raisin cubs, wha mey be left alane for extendit periods o time. Members o the pride regularly tend tae play the same role in hunts an hone thair skills. The heal o the hunters is the primary need for the survival o the pride, an thay are the first tae consume the prey at the site it is taken. Ither benefits include possible kin selection (better tae share fuid wi a relatit lion than wi a stranger), pertection o the young, maintenance o territory, an individual insurance against injury an hunger.[17]

Video o a wild lioness

Lionesses dae maist o the huntin for thair pride. Thay are mair effective hunters, as thay are smawer, swifter, an mair agile than the males an unencumbered bi the hivy an conspicuous mane, that causes owerheatin during exertion. Thay act as a coordinatit group wi members wha perform the same role consistently in order tae stalk an bring doun the prey successfully. Smawer prey is eaten at the location o the hunt, tharebi bein shared amang the hunters; whan the kill is lairger it eften is dragged tae the pride aurie. Thare is mair sharin o lairger kills,[37] awbesit pride members aften behave aggressively toward ane anither as ilkane ettles at consumin sae muckle a hantle o fuid as possible. Naur the conclusion o the hunt, males hae a tendency tae dominate the kill ance the lionesses hae succeeded. Thay are mair lik tae share this wi the cubs as wi the lionesses, but males rarely share fuid thay hae killt bi thairsels.

Baith males an females can defend the pride against intruders, but the male lion is better-suitit for this purpose due tae its stockier, mair powerful big.[38] Some individuals consistently lead the defence against intruders, while ithers lag behind.[39] Lions tend tae assume speceefic roles in the pride. Those laggin behind mey provide ither valuable services tae the group.[40] An alternative hypothesis is that thare is some reward associated wi bein a leader wha fends aff intruders, an the rank o lionesses in the pride is reflectit in these responses.[41] The male or males associatit wi the pride must defend thair relationship tae the pride frae ootside males wha attempt tae tak ower thair relationship wi the pride.

Huntin an diet[eedit | eedit soorce]

Lion teeth are teepical o a carnivore.

Lions prefer tae scavenge whan the opportunity presents itsel[42] wi carrion providin mair nor 50% o thair diet.[43] Thay scavenge ainimals either deid frae naitural causes (disease) or killt bi ither predators, an keep a constant lookout for circlin vultures, bein keenly aware that thay indicate an ainimal dead or in distress.[42] In fact, maist deid prey on whilk baith hyenas an lions feed upon are killt bi the hyenas insteid o the lions.[4]

The lionesses dae maist o the huntin for the pride. The male lion associatit wi the pride uisually stays an watches ower young cubs till the lionesses return frae the hunt. Teepically, several wirk thegither an encircle the herd frae different pynts. Ance thay hae closed in on the herd, thay uisually target the ainimal closest tae them. The attack is short an pouerfu; thay attempt tae catch the victim wi a fast rush an final leap. The prey uisually is killt bi strangulation,[44] that can cause cerebral ischemia or asphyxia (that results in hypoxemic, or "general", hypoxia). The prey an aa mey be killt bi the lion enclosin the ainimal's mooth an nostrils in its jaws (that wad result in asphyxia an aa).[4]

Lions uisually hunt in coordinatit groups an stalk thair chosen prey. Houiver, thay are nae parteecularly kent for thair stamina – for instance, a lioness' hert maks up anly 0.57% o her body wecht (a male's is aboot 0.45% o his body wecht), whauras a hyena's hert is close tae 1% o its body wecht.[45] Thus, thay anly run fast in short bursts,[46] an need tae be close tae thair prey afore stairtin the attack. Thay tak advantage o factors that reduce veesibility; mony kills tak place naur some form o cover or at nicht.[47] Thay sneak up tae the victim till thay reach a distance o approximately 30 metres (98 feet) or less.

Lioness in a burst o speed while huntin.

The prey consists mainly o medium-sized mammals, wi a preference for wildebeest, zebras, buffalo, an warthogs in Africae an nilgai, wild boar, an several deer species in Indie. Mony ither species are huntit, based on availability, mainly ungulates wichin atween 50 an 300 kg (110 an 660 lb) lik kudu, hartebeest, gemsbok, an eland.[4] Occasionally, thay tak relatively smaw species lik Thomson's gazelle or springbok. Lions huntin in groups are capable o takin doun maist ainimals, even adults in guid heal, but in maist pairts o thair range thay rarely attack verra lairge prey lik fully grown male giraffes due tae the danger o injury. Giraffes an buffaloes are awmaist invulnerable tae a solitary lion as well.[48]

Extensive studies shaw that lionesses normally prey on mammals wi an average wecht o 126 kg (278 lb), while kills made bi male lions average 399 kg (880 lb).[49] In Africae, wildebeest rank at the tap o preferred prey (makin nearly hauf o the lion prey in the Serengeti) follaed bi zebra.[50] Lions dae nae prey on fully grown adult elephants; maist adult hippopotamuses, rhinoceroses, an smawer gazelles, impala, an ither agile antelopes are generally excludiy. Houiver, giraffes an buffaloes are eften taken in certain regions. For instance, in Kruger Naitional Pairk, giraffes are regularly huntit.[51] In Manyara Pairk, Cape buffaloes constitute as much as 62% o the lion's diet,[52] due tae the heich nummer density o buffaloes. Occasionally hippopotamus is an aa taken, but adult rhinoceroses are generally avoidit. Warthogs are eften taken dependin on availability.[53] The lions o Savuti, Botswana, hae adaptit tae huntin young elephants during the dry saison, an a pride o 30 lions haes been recordit killin individuals atween the ages o fower an eleven years.[54] In the Kalahari desert in Sooth Africae, black-maned lions mey chase baboons up a tree, wait patiently, then attack them whan thay try tae escape:

Baboons trapped up a tree bi lions.

Lions an aa attack domestic livestock an in Indie cattle contribute signeeficantly tae thair diet.[23] Lions are capable o killin ither predators lik leopards, cheetahs, hyenas, an wild dugs, awbesit (unlik maist felids) thay seldom devour the competitors efter killin them. A lion mey gorge itsel an eat up tae 30 kg (66 lb) in ane sittin;[55] if it is unable tae consume aw the kill it will rest for a few oors afore consumin mair. On a het day, the pride mey retreat tae shade leavin a male or twa tae staund guard.[56] An adult lioness requires an average o aboot 5 kg (11 lb) o meat per day, a male aboot 7 kg (15 lb).[57]

Four lionesses catch a cape buffalo.

Acause lionesses hunt in open spaces whaur thay are easy seen bi thair prey, cooperative huntin increases the likelihood o a successfu hunt; this is especially true wi lairger species. Teamwork an aa enables them tae defend thair kills easier against ither lairge predators lik hyenas, that mey be attractit bi vultures frae kilometres away in open savannas. Lionesses dae maist o the huntin; males attached tae prides dae nae uisually pairteecipate in huntin, except in the case o lairger quarry lik giraffe an buffalo. In teepical hunts, ilka lioness haes a favored poseetion in the group, either stalkin prey on the "weeng" then attackin, or muivin a smawer distance in the centre o the group an capturin prey in flicht frae ither lionesses.[58] Thare is evidence that male lions are juist as successfu at huntin as females; thay are solo hunters wha ambush prey in smaw bush.[59] Young lions first display stalkin behaviour aboot three months o age, awbesit thay dae nae pairteecipate in huntin till thay are awmaist a year auld. Thay begin tae hunt effectively whan thay are naur twa yeir auld.[60]

Predator competeetion[eedit | eedit soorce]

Subadult male lion an spotted hyena

Lions an spottit hyenas occupy the same ecological niche, meanin thay compete for prey an carrion in the auries whaur thay coexist. A review o data across several studies indicates a dietary owerlap o 58.6%.[61] Lions teepically ignore spottit hyenas unless the lions are on a kill or are bein harassed bi the hyenas, while the latter tend tae visibly react tae the presence o lions whether thare is fuid or nae. Lions seize the kills o spottit hyenas: in the Ngorongoro crater, it is common for lions tae subsist largely on kills stolen frae hyenas, causin the hyenas tae increase thair kill rate.[62] On the ither haund, in Northren Botswana's Chobe Naitional Pairk, the situation is reversed: hyenas frequently challenge lions an steal thair kills: thay obtain fuid frae 63% o aw lion kills.[63] whan confrontit on a kill bi lions, spottit hyenas mey either leave or wait patiently at a distance o 30–100 m (98–328 ft) till the lions hae finished,[64] but thay are an aa bauld enough tae feed alangside lions, an even force the lions off a kill. The twa species mey attack ane anither even whan thare is no fuid involved for no apparent reason.[65][66] Lion predation can accoont for up tae 71% o hyena daiths in Etosha. Spottit hyenas hae adaptit bi frequently mobbin lions that enter thair territories.[67] Experiments on captive spottit hyenas revealed that specimens wi no prior experience wi lions act indifferently tae the sicht o them, but will react fearfully tae the scent.[62] The size o male lions allows them occasionally tae confront hyenas in itherwise evenly matched brawls an sae tae tip the balance in favour o the lions.

Lions tend tae dominate smawer felines lik cheetahs an leopards whaur thay co-occur, stealin thair kills an killin thair cubs an even adults whan gien the chance. The cheetah haes a 50% chance o losin its kill tae lions or ither predators.[68] Lions are major killers o cheetah cubs, up tae 90% o thaim are lost in thair first weeks o life due tae attacks bi ither predators. Cheetahs evit competeetion bi huntin at different times o the day an hide thair cubs in thick brush. Leopards an aa uise siclike tactics, but hae the advantage that thay can tae subsist a fair bit better on smaw prey nor on aither lions or cheetahs. An aa, unlik cheetahs, leopards can clim trees an uise them tae haud thair cubs an kills away frae lions; houiver, lionesses will occasionally be successfu in climin tae retrieve leopard kills.[69] Seemilarly, lions dominate African wild dugs; thay nae juist tak thair kills but preying on young an (rarely) adult dugs an aa. Population densities o wild dugs are laich in auries whaur lions are mair abundant.[70] Houiver, thare are a few reportit cases o auld an woondit lions fawin prey tae wild dugs.[51][71]

The Nile crocodile is the anly sympatric predator (besides humans) that can singly threaten the lion. Dependin on the size o the crocodile an the lion, aither can lose kills or carrion tae the ither. Lions hae been kent tae kill crocodiles venturin ontae laund,[72] while the reverse is true for lions enterin watterways, as evidenced bi the occasional lion claw foond in crocodile stomachs.[73]

Man-eatin[eedit | eedit soorce]

Main article: Man-eater

While lions dae nae uisually hunt fowk, some (uisually males) seem tae seek oot human prey; ane well-publicised case includes the Tsavo maneaters, whaur 28 offeecially recordit railway wirkers biggin the Kenyae-Uganda Railway war taken bi lions ower nine months during the construction o a brig ower the Tsavo River in Kenyae in 1898.[74] The hunter wha kill the lions wrote a beuk detailin the ainimals' predatory behaviour. The lions war lairger than normal, lacked manes, an ane seemed tae suffer frae tuith decay. The infirmity theory, includin tuith decay, is nae favored bi aw researchers; an analysis o teeth an jaws o man-eatin lions in museum collections suggests that while tuith decay mey explain some incidents, prey depletion in human-dominatit auries is a mair likely cause o lion predation on humans.[75]

In thair analysis o Tsavo an general man-eatin, Kerbis Peterhans an Gnoske acknowledge that sick or injurt ainimals mey be mair prone tae man-eatin, but that the behaviour is "nae unuisual, nor necessarily 'aberrant'" whaur the opportunity exists; if inducements lik access tae livestock or human corpses are present, lions will regularly prey upon human beins. The authors note that the relationship is well-attestit amang ither pantherines an primates in the paleontological record.[76]

The lion's proclivity for man-eatin haes been seestematically examined. American an Tanzanian scientists report that man-eatin behaviour in rural auries o Tanzanie increased a fair bit frae 1990 tae 2005. At least 563 veelagers war attacked an mony eaten ower this period – a nummer far exceedin the mair famed "Tsavo" incidents o a century earlier. The incidents occurred naur aboots Selous Naitional Pairk in Rufiji Destrict an in Lindi Province near the Mozambican border. While the expansion o villagers intae bush kintra is ane concern, the authors argue that conservation policy must mitigate the danger acause, in this case, conservation contreibutes directly tae human daiths. Cases in Lindi hae been documentit whaur lions seize humans frae the center o substantial veelages.[77] Anither study o 1,000 fowk attacked bi lions in soothren Tanzanie atween 1988 an 2009 foond that the weeks follaein the full muin (whan thare wis less muinlight) war a strang indicator o increased nicht attacks on fowk.[78]

The Tsavo Man-Eaters on display in the Field Museum o Naitural History in Chicago, Illinois

Author Robert R. Frump wrote in The Man-eaters of Eden that Mozambican refugees regularly crossin Kruger Naitional Pairk at nicht in Sooth Africae are attacked an eaten bi the lions; pairk offeecials hae concedit that man-eatin is a problem thare. Frump believes thoosands mey hae been killt in the decades efter apartheid sealed the pairk an forced the refugees tae cross the pairk at nicht. For nearly a century afore the border wis sealed, Mozambicans haed regularly walked athort the pairk in daytime wi little harm.[79]

Packer estimates mair nor 200 Tanzanians are killt ilka year bi lions, crocodiles, elephants, hippos, an snakes, an that the nummers could be dooble that amount, wi lions thocht tae kill at least 70 o those. Packer haes documented that atween 1990 an 2004, lions attacked 815 fowk in Tanzanie, killin 563. Packer an Ikanda are amang the few conservationists wha believe wastren conservation efforts must tak accoont o these matters nae juist acause o ethical concerns aboot human life, but an aa for the lang term success o conservation efforts an lion preservation.[77]

A man-eatin lion wis killt bi gemme scouts in Soothren Tanzanie in Aprile 2004. It is believed tae hae killt an eaten at least 35 fowk in a series o incidents coverin several veelages in the Rufiji Delta coastal region.[80] Dr Rolf D. Baldus, the GTZ wildlife programme coordinator, commentit that it wis likely that the lion preyed on humans acause it haed a lairge abscess unnerneath a molar that wis cracked in several places. He further commentit that "This lion probably experienced a lot o pain, pairteecularly whan it wis chewin."[81] GTZ is the German development cooperation agency an haes been wirkin wi the Tanzanian govrenment on wildlife conservation for nearly twa decades. As in ither cases this lion wis lairge, lacked a mane, an haed a tuith problem.

The "Aw-Africae" record o man-eatin generally is considered tae be nae Tsavo, but incidents in the early 1930s throu the late 1940s in whit wis then Tanganyika (nou Tanzanie). George Rushby, gemme warden an professional hunter, eventually dispatched the pride, that ower three generations is thocht tae hae killt an eaten 1,500 tae 2,000 fowk in what is nou Njombe destrict.[82]

Reproduction an life cycle[eedit | eedit soorce]

Maist lionesses will hae reproduced bi the time thay are fower years o age.[83] Lions dae nae mate at ony specific time o year, an the females are polyestrous.[84] As wi ither cats' penises, the male lion's penis haes spines that pyntin backward. During widrawal o the penis, the spines rake the waws o the female's vagina, that mey cause ovulation.[85] A lioness mey mate wi mair nor ane male whan she is in heat.[86]

Matin lions.

The average gestation period is aroond 110 days,[84] the female giein birth tae a litter o ane tae fower cubs in a secluded den (that mey be a thicket, a reed-bed, a cave, or some ither sheltered aurie) uisually away frae the rest o the pride. She will eften hunt bi hersel while the cubs are still helpless, stayin relatively close tae the thicket or den whaur the cubs are kept.[87] The cubs themsels are born blind – thair ees dae nae open till roughly a week efter birth. Thay weigh 1.2–2.1 kg (2.6–4.6 lb) at birth an are awmaist helpless, beginnin tae crawl a day or twa efter birth an walkin aroond three weeks o age.[88] The lioness muives her cubs tae a new den steid several times a month, carrying them ane bi ane bi the nape o the neck, tae prevent scent frae biggin up at a single den steid an thus avoidin the attention o predators that mey harm the cubs.[87]

Uisually, the mither daes nae integrate hersel an her cubs back intae the pride until the cubs are sax tae aicht weeks auld.[87] Whiles this introduction tae pride life occurs earlier, houiver, parteecularly if ither lionesses hae gien birth at aboot the same time. For instance, lionesses in a pride eften synchronise thair reproductive cycles sae that thay cooperate in the raising an souklin o the young (ance the cubs are past the initial stage o isolation wi thair mither), wha suckle indiscriminately frae ony or aw o the nouricin females in the pride. In addition tae greater pertection, the synchronization o births an aa haes an advantage in that the cubs end up bein roughly the same size, an thus hae an equal chance o survival. If ane lioness gies birth tae a litter o cubs a couple o months efter anither lioness, for instance, then the younger cubs, bein much smawer than thair aulder brethren, uisually are dominatit bi lairger cubs at mealtimes – consequently, daith bi starvation is mair common amang the younger cubs.

A pregnant lioness (richt).

In addition tae stairvation, cubs an aa face mony ither dangers, lik predation bi jackals, hyenas, leopards, mairtial eagles, an snakes. Even buffaloes, should thay catch the scent o lion cubs, eften stampede toward the thicket or den whaur thay are bein kept, daein thair best tae trample the cubs tae daith while wardin off the lioness. Furthermair, whan ane or mair new males oust the previous male(s) associatit wi a pride, the conqueror(s) eften kill ony existin young cubs,[89] perhaps acause females dae nae acome fertile an receptive till thair cubs mature or dee. Aw in aw, as mony as 80% o the cubs will dee afore the age o twa.[90]

whan first introduced tae the rest o the pride, the cubs initially lack confidence whan confrontit wi adult lions ither than thair mither. Thay suin begin tae immerse thairsels in the pride life, houiver, playin amang thairsels or attemptin tae initiate play wi the adults. Lionesses wi cubs o thair ain are mair likely tae be tolerant o anither lioness's cubs than lionesses wioot cubs. The tolerance o the male lions toward the cubs varies – whiles, a male will patiently let the cubs play wi his tail or his mane, whauras anither mey snarl an bat the cubs away.[91]

Male lions are generally mair likely tae share fuid caught bi the pride wi cubs than wi lionesses, but rarely share thair ain catches wi ithers.

Weanin occurs efter sax tae seiven months. Male lions reach maturity at aboot 3 years o age an, at 4–5 years o age, are capable o challengin an displacing the adult male(s) associatit wi anither pride. Thay begin tae age an weaken atween 10 an 15 years o age at the latest,[92] if thay hae nae already been creetically injured while defendin the pride (ance ousted frae a pride bi rival males, male lions rarely manage a seicont tak-ower). This leaves a short window for thair ain affspring tae be born an matur. If thay are able tae procreate as suin as thay tak ower a pride, potentially, thay mey hae mair affspring reachin maturity afore thay an aa are displaced. A lioness eften will attempt tae defend her cubs fiercely frae a usurpin male, but such actions are rarely successfu. He uisually kills aw o the existin cubs wha are less nor twa years auld. A lioness is weaker an much lichter than a male; success is mair likely whan a group o three or fower mithers within a pride jyn forces against ane male.[89]

Contrary tae popular belief, it is nae anly males that are oostit frae thair pride tae acome nomads, awbesit maist females certainly dae remain wi thair birth pride. Houiver, whan the pride acomes too lairge, the next generation o female cubs mey be forced tae leave tae eke oot thair ain territory. Furthermair, whan a new male lion taks ower the pride, subadult lions, baith male an female, mey be evictit.[93] Life is harsh for a female nomad. Nomadic lionesses rarely manage tae raise thair cubs tae maturity, wioot the pertection o ither pride members.

Heal[eedit | eedit soorce]

awbesit adult lions hae no naitural predators, evidence suggests that the majority dee violently frae humans or ither lions.[94] Lions eften inflict serious injuries on ilka ither, either members o different prides encoonterin ilka ither in territorial disputes, or members o the same pride fechtin at a kill.[95] Crippled lions an lion cubs mey faw victim tae hyenas, leopards, or be trampled bi buffalo or elephants, an careless lions mey be maimed whan huntin prey.[96]

Lions seekin refuge frae flies bi climbin a tree.

Various species o tick commonly infest the lugs, neck an groin regions o maist lions.[97][98] Adult forms o several species o the tapeworm genus Taenia hae been isolatit frae thairms, the lions haein ingestit larval forms frae antelope meat.[99] Lions in the Ngorongoro Crater war afflictit bi an ootbreak o stable flee (Stomoxys calcitrans) in 1962; this resultit in lions acomin covered in bluidy bare patches an emaciatit. Lions soucht unsuccessfully tae evade the bitin flees bi climbin trees or crawlin intae hyena burrows; mony perished or emigratit as the population dropped frae 70 tae 15 individuals.[100] A mair recent ootbreak in 2001 killt sax lions.[101] Lions, especially in captivity, are vulnerable tae the canine distemper virus (CDV), feline immunodeficiency virus (FIV), an feline infectious peritonitis (FIP).[27] CDV is spread throu domestic dugs an ither carnivores; a 1994 outbreak in Serengeti Naitional Pairk resultit in mony lions developin neurological seemptoms lik seizurs. During the ootbreak, several lions dee'd frae pneumonia an encephalitis.[102] FIV, that is seemilar tae HIV while nae kent tae adversely affect lions, is worrisome eneuch in its effect in domestic cats that the Species Survival Plan recommends systematic testin in captive lions. It occurs wi heich tae endemic frequency in several wild lion populations, but is maistly absent frae Asiatic an Namibian lions.[27]

Communication[eedit | eedit soorce]

Heid rubbin an lickin are common social behaviours athin a pride

whan restin, lion socialization occurs throu a nummer o behaviours, an the ainimal's expressive movements are heichly developed. The maist common peacefu tactile gestures are heid rubbin an social lickin,[103] that hae been compared wi gruimin in primates.[104] Heid rubbin – nuzzlin ane's foreheid, face an neck against anither lion – appears tae be a form o greetin,[105] as it is seen eften efter an ainimal haes been apart frae others, or efter a fecht or confrontation. Males tend tae rub ither males, while cubs an females rub females.[106] Social lickin eften occurs in tandem wi heid rubbin; it is generally mutual an the recipient appears tae express pleasure. The heid an neck are the maist common pairts o the body licked, that mey hae arisen oot o utility, as a lion canna lick these auries individually.[107]

A lion in captivity roaring

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Lions hae an array o facial expressions an body posturs that serve as veesual gesturs.[108] Thair repertoire o vocalisations is an aa lairge; variations in intensity an pitch, rather than discrete signals, appear central tae communication. Lion soonds include snarlin, hissin, coughin, miaowin, woofin, an roarin. Lions tend tae roar in a verra characteristic manner, stairtin wi a few deep, lang roars that trail off intae a series o shorter anes.[109][110] Thay maist eften roar at nicht; the soond, that can be heard frae a distance o 8 kilometres (5.0 mi), is uised tae advertise the ainimal's presence.[111] Lions hae the loudest roar o ony big cat.

Distribution an habitat[eedit | eedit soorce]

Twa male Asiatic lions. The wild population o the endangered Asiatic lions is restrictit tae the Gir Wid Naitional Pairk in wastren Indie.[112]

In Africae, lions can be foond in savanna grasslands wi scattered Acacia trees, that serve as shade;[113] thair habitat in Indie is a mixtur o dry savanna wid an verra dry deciduous scrub wid.[114] The habitat o lions oreeginally spanned the soothren pairts o Eurasie, rangin frae Greece tae Indie, an maist o Africae except the central rainwid-zone an the Sahara desert. Herodotus reportit that lions haed been common in Greece in 480 BC; thay attacked the baggage camels o the Persie keeng Xerxes on his march throu the kintra. Aristotle considered them rare bi 300 BC. Bi 100 AD thay war extirpated.[115] A population o Asiatic lions survived till the tent century in the Caucasus, thair last European ootpost.[116]

The species wis eradicatit frae Palestine bi the Middle Ages an frae maist o the rest o Asie efter the arrival o readily available firearms in the aichteent century. Atween the late nineteent an early twentiet century, thay became extinct in North Africae an Soothwast Asie. Bi the late nineteenth century, the lion haed disappeared frae Turkey an maist o northren Indie,[27][117] while the last sichtin o a live Asiatic lion in Iran wis in 1941 (atween Shiraz an Jahrom, Fars Province), awbesit the corpse o a lioness wis foond on the banks o the Karun river, Khūzestān Province in 1944. Thare are no subsequent reliable reports frae Iran.[55] The subspecies nou survives anly in an aroond the Gir Wid o northwastren Indie.[118] Approximately 500 lions live in the aurie o the 1,412 km2 (545 sq mi) sanctuary in the state o Gujarat, that covers maist o the wid. Thair nummers hae increased frae 180 tae 523 ainimals mainly acause the naitural prey species hae recovered.[119][120]

Population an conservation status[eedit | eedit soorce]

Main article: Lion huntin
The Asiatic lion, whose habitat ance ranged frae the Mediterranean tae north-wast Indian subcontinent, is the day foond anly in the Gir Wid o Gujarat, Indie. It wis estimatit that anly 523 Asiatic lions survive in the wild.[121]

Maist lions nou live in eastren an soothren Africae, an thair nummers thare are rapidly decreasin, wi an estimatit 30–50% decline per 20 years in the late hauf o the 20t century.[2] Estimates o the African lion population range atween 16,500 an 47,000 livin in the wild in 2002–2004,[122][123] doun frae early 1990s estimates that ranged as heich as 100,000 an perhaps 400,000 in 1950. Primary causes o the decline include disease an human interference.[2] Habitat loss an conflicts wi humans are considered the maist signeeficant threats tae the species.[124][125] The remainin populations are aften geografically isolatit frae ane anither, that can lead tae inbreedin, an consequently, reduced genetic diversity. Tharefore the lion is considered a vulnerable species bi the Internaitional Union for Conservation o Naitur, while the Asiatic subspecies is endangered.[126] The lion population in the region o Wast Africae is isolatit frae lion populations o Central Africae, wi nae or naur nae exchynge o breedin individuals. The nummer o matur individuals in Wast Africae is estimatit bi twa separate recent surveys at 850–1,160 (2002/2004). Thare is disagreement ower the size o the lairgest individual population in Wast Africae: the estimates range frae 100 tae 400 lions in Burkina Faso's Arly-Singou ecoseestem.[2] Anither population in northwastren Africae is foond in Waza Naitional Pairk, whaur approximately 14–21 ainimals persist.[127]

Black maned male lion, shot in the Sotik Plains, Kenyae (Mey 1909)

Conservation o baith African an Asian lions haes required the setup an maintenance o naitional parks an gemme reserves; amang the best kent are Etosha Naitional Pairk in Namibie, Serengeti Naitional Pairk in Tanzanie, an Kruger Naitional Pairk in eastren Sooth Africae. The Ewaso Lions Project protects lions in the Samburu Naitional Reserve, Buffalo Springs Naitional Reserve an Shaba Naitional Reserve o the Ewaso Ng'iro ecoseestem in Northren Kenyae.[128] Ootside these auries, the issues arising frae lions' interaction wi livestock an fowk uisually results in the elimination o the umwhile.[129] In Indie, the last refuge o the Asiatic lion is the 1,412 km2 (545 sq mi) Gir Wid Naitional Pairk in wastren Indie, that haed approximately 180 lions in 1974 an aboot 400 in 2010.[119] As in Africae, numerous human habitations are close bi wi the resultant problems atween lions, livestock, locals an wildlife offeecials.[130] The Asiatic Lion Reintroduction Project plans tae establish a seicont independent population o Asiatic lions at the Kuno Wildlife Sanctuary in the Indian state o Madhya Pradesh.[131] It is important tae stairt a seicont population tae serve as a gene puil for the last survivin Asiatic lions an tae help develop an maintain genetic diversity enabling the species tae survive.

The umwhile popularity o the Barbary lion as a zoo ainimal haes meant that scattered lions in captivity are as lik tae be descendit frae Barbary lion stock. This includes lions at Port Lympne Wild Ainimal Pairk in Kent, Ingland that are descendit frae ainimals ained bi the Keeng o Morocco.[132] Anither eleven ainimals believed tae be Barbary lions war foond in Addis Ababa zoo, descendants o ainimals ained bi Emperor Haile Selassie. WildLink Internaitional, in collaboration wi Oxford Varsity, launched thair ambeetious Internaitional Barbary Lion Project wi the aim o identifyin an breedin Barbary lions in captivity for eventual reintroduction intae a naitional pairk in the Atlas Moontains o Morocco.[22]

Follaein the discovery o the decline o lion population in Africae, several coordinatit efforts involvin lion conservation hae been organised in an attempt tae stem this decline. Lions are ane species included in the Species Survival Plan, a coordinatit attempt bi the Association o Zoos an Aquariums tae increase its chances o survival. The plan wis oreeginally stairtit in 1982 for the Asiatic lion, but wis suspendit whan it wis foond that maist Asiatic lions in North American zoos war nae genetically pure, haein been hybridised wi African lions. The African lion plan stairtit in 1993, focusin especially on the Sooth African subspecies, awbesit thare are difficulties in assessin the genetic diversity o captive lions, syne maist individuals are o unkent origin, makin maintenance o genetic diversity a problem.[27]

In captivity[eedit | eedit soorce]

Lions are pairt o a group o exotic ainimals that are the core o zoo exhibits syne the late aichteent century; members o this group are invariably lairge vertebrates an include elephants, rhinoceroses, hippopotamuses, lairge primates, an ither big cats; zoos sought tae gather as mony o these species as possible.[133] awbesit mony modren zoos are mair selective aboot thair exhibits,[134] thare are mair nor 1,000 African an 100 Asiatic lions in zoos an wildlife pairks aroond the warld. Thay are considered an ambassador species an are kept for tourism, eddication an conservation purposes.[135] Lions can reach an age o ower 20 years in captivity; Apollo, a resident lion o Honolulu Zoo in Honolulu, Hawaii, dee'd at age 22 in August 2007. His twa sisters, born in 1986, war still alive in August 2007.[136] Breedin programs need tae note origins tae avoid breedin different subspecies an thus reducing conservation value.[137] Houiver, several Asiatic-African lion crosses hae been bred.[138]

Male African lion o the Transvaal subspecies (P. l. krugeri)

At the ancient Egyptian ceeties o Taremu an Per-Bast war temples tae the lioness goddesses o Egyp, Sekhmet an Bast an at Taremu thare wis a temple tae the son o the deity, Maahes the lion prince, whaur live lions war kept an allaed tae roam within his temple. The Greeks cried the ceety Leontopolis, the "Ceety o Lions" an documentit that practice. Lions war kept an bred bi Assirie keengs as early as 850 BC,[115] an Alexander the Great wis said tae hae been presentit wi tame lions bi the Malhi o northren Indie.[139] Later in Roman times, lions war kept bi emperors tae tak pairt in the gladiator arenas or for executions (see bestiarii, damnatio ad bestias, an venatio). Roman notables, includin Sulla, Pompey, an Julius Caesar, eften ordered the mass slaughter o hunders o lions at a time.[140] In the East, lions war tamed bi Indian princes, an Marco Polo reportit that Kublai Khan kept lions inside.[141] The first European "zoos" spread amang noble an ryal faimilies in the thirteent century, an till the seiventeent century war cried seraglios; at that time, thay came tae be cried menageries, an extension o the cabinet o curiosities. Thay spread frae Fraunce an Italy during the Renaissance tae the rest o Europe.[142] In England, awbesit the seraglio tradeetion wis less developed, Lions war kept at the Touer o Lunnon in a seraglio established bi Keeng John in the thirteent century,[143][144] probably stocked wi ainimals frae an earlier menagerie stairtit in 1125 bi Henry I at his huntin lodge in Woodstock, near Oxford; whaur lions haed reportedly been stocked bi William o Malmesbury.[145]

Seraglios served as expressions o the nobility's pouer an wealth. Ainimals lik big cats an elephants, in parteecular, seembolised pouer, an would be pittit in fechts against ilka ither or domesticatit ainimals. Bi extension, menageries an seraglios served as demonstrations o the dominance o humanity ower naitur. Consequently, the defeat o such naitural "lairds" bi a cou in 1682 astonished the spectators, an the fecht o an elephant afore a rhinoceros drew jeers. Such fechts would slowly fade oot in the seiventeent century wi the spread o the menagerie an thair appropriation bi the commoners. The tradeetion o keepin big cats as pets would last intae the nineteent century; at that time it wis seen as heich eccentric.[146]

Albrecht Dürer, Lions sketch. (circa 1520)

The presence o lions at the Touer o Lunnon wis intermittent, bein restocked whan a monarch or his consort, lik Margaret o Anjou the wife o Henry VI, either soucht or war gien ainimals. Records indicate thay war kept in puir condeetions thare in the seiventeent century, in contrast tae mair open condeetions in Florence at the time.[147] The menagerie wis open tae the public bi the aichteent century; admission wis a sum o three hauf-pence or the supply o a cat or dug for feedin tae the lions.[148] A rival menagerie at the Exeter Exchynge an aa exhibitit lions till the early nineteent century.[149] The Touer menagerie wis closed doun bi William IV,[148] an ainimals transferred tae the Lunnon Zoo, that opened its gates tae the public on 27 Aprile 1828.[150]

The wild ainimals trade flourished alongside improved colonial trade o the nineteent century. Lions war considered fairly common an wanpricey. Awbesit thay wad barter heicher nor teegers, thay war less costly than lairger, or mair difficult tae transport ainimals lik the giraffe an hippopotamus, an much less nor giant pandas.[151] Lik ither ainimals, lions war seen as little mair nor a naitural, boundless commodity that wis mercilessly exploited wi terrible losses in capture an transportation.[152] The widely reproduced imagery o the heroic hunter chasin lions would dominate a lairge pairt o the century.[153] Splorers an hunters exploitit a popular Manichean division o ainimals intae "guid" an "evil" tae add thrillin value tae thair adventurs, castin thairsels as heroic figurs. This resultit in big cats, always suspectit o bein man-eaters, representin "baith the fear o naitur an the satisfaction o havin owercome it."[154]

Lion at Melbourne Zoo enjoyin an elevatit gressy aurie wi some tree shelter

Lions war kept in cramped an squalid condeetions at Lunnon Zoo till a lairger lion hoose wi roomier cages wis biggit in the 1870s.[155] Further changes teuk place in the early twentiet century, whan Carl Hagenbeck designed enclosurs mair closely resemblin a naitural habitat, wi concrete 'rocks', mair open space an a moat insteid o bars. He designed lion enclosurs for baith Melbourne Zoo an Sydney's Taronga Zoo, amang ithers, in the early twentiet century. awbesit his designs war popular, the auld bars an cage enclosurs prevailed till the 1960s in mony zoos.[156] In the later decades o the twentiet century, lairger, mair naitural enclosurs an the uise o wire mesh or laminatit glass insteid o lawered dens allaed veesitors tae come closer norr ever tae the ainimals, wi some attractions even placin the den on grund heicher than visitors, lik the Cat Wid/Lion Owerleuk o Oklahoma City Zoological Pairk.[27] Lions are nou placed in gey muckler naturalistic auries; modren recommendit guidelines mair closely approximate condeetions in the wild wi closer attention tae the lions' needs, heichlichtin the need for dens in separate auries, elevatit poseetions in baith sun an shade whaur lions can sit an adequate grund cover an drainage as well as sufficient space tae roam. Thare hae an aa been instances whaur a lion wis kept bi a private individual, lik the lioness Elsa, wha wis raised bi George Adamson an his wife Joy Adamson an came tae develop a strang bond wi them, parteecularly the latter. The lioness later achieved fame, her life bein documentit in a series o beuks an films.

Baitin an tamin[eedit | eedit soorce]

Main articles: Lion-baitin and Lion tamin
Nineteent century etchin o a lion tamer in a cage o lions

Lion-baitin is a bluid sport involvin the baitin o lions in combat wi ither ainimals, uisually dugs. Records o it exist in auncient times throu till the seiventeent century. It wis finally banned in Vienna bi 1800 an Ingland in 1835.[157][158]

Lion tamin refers tae the practice o tamin lions for enterteenment, either as pairt o an established circus or as an indiveedual act, lik Siegfried & Roy. The term is an aa eften uised for the tamin an display o ither big cats lik teegers, leopards, an cougars. The practice wis pioneered in the first hauf o the nineteent century bi Frenchman Henri Martin an American Isaac Van Amburgh wha baith toured widely, an whase techniques war copied bi a nummer o follaers.[159] Van Amburgh performed afore Queen Victoria in 1838 whan he toured Great Breetain. Martin componed a pantomime titled Les Lions de Mysore ("the lions o Mysore"), an idea that Amburgh quickly borraed. These acts eclipsed equestrianism acts as the central display o circus shaws, but truly entered public consciousness in the early twentiet century wi cinema. In demonstratin the superiority o human ower ainimal, lion tamin served a purpose seemilar tae ainimal fichts o previous centuries.[159] The ultimate pruif o a tamer's dominance an control ower a lion is demonstratit bi placin his heid in the lion's gab. The nou iconic lion tamer's chair wis possibly first uised bi American Clyde Beatty (1903–1965).[160]

Notes[eedit | eedit soorce]

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  3. 3.0 3.1 3.2 Linnaeus, Carolus (1758). Systema naturae per regna tria naturae :secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. (in Latin) 1 (10th ed.). Holmiae (Laurentii Salvii). p. 41. Retrieved 8 September 2008. 
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  10. Liddell, Henry George; Scott, Robert (1980). A Greek-English Lexicon (Abridged Edition). United Kingdom: Oxford University Press. p. 411. ISBN 0-19-910207-4. 
  11. Simpson, John; Weiner, Edmund, ed. (1989). "Lion". Oxford English Dictionary (2nd ed.). Oxford: Clarendon Press. ISBN 0-19-861186-2. 
  12. Heptner, V. G.; Sludskii, A. A. (1992). Mammals of the Soviet Union, Volume II, Part 2. Leiden u.a.: Brill. ISBN 90-04-08876-8. 
  13. Schaller, p. 28.
  14. Smuts, G. L.; Robinson, G.A.; Whyte, I.J. (1980). "Comparative growth of wild male and female lions (Panthera leo)". Journal of Zoology 190 (3): 365–373. doi:10.1111/j.1469-7998.1980.tb01433.x. 
  15. Mazák, Vratislav (April 2004) Der Tiger, Westarp Wissenschaften, 5th ed., pp. 178 ff., ISBN 3-89432-759-6.
  16. Wood, The Guinness Book of Animal Facts and Feats. Sterling Pub Co Inc (1983), ISBN 978-0-85112-235-9
  17. 17.0 17.1 Nowell, Kristin; Jackson, Peter (1996). "Panthera Leo". Wild Cats: Status Survey and Conservation Action Plan (PDF). Gland, Switzerland: IUCN/SSC Cat Specialist Group. pp. 17–21, 37–41. ISBN 2-8317-0045-0. 
  18. Trivedi, Bijal P. (2005). "Are Maneless Tsavo Lions Prone to Male Pattern Baldness?". National Geographic. Retrieved 7 July 2007. 
  19. Trivedi, Bijal P. (22 August 2002). "Female Lions Prefer Dark-Maned Males, Study Finds". National Geographic News. National Geographic. Retrieved 1 September 2007. 
  20. 20.0 20.1 West, Peyton M.; Packer, Craig (August 2002). "Sexual Selection, Temperature, and the Lion's Mane". Science 297 (5585): 1339–43. doi:10.1126/science.1073257. PMID 12193785. 
  21. Barnett, Ross; Yamaguchi, Nobuyuki; Barnes, Ian; Cooper, Alan (August 2006). "Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation". Conservation Genetics 7 (4): 507–14. doi:10.1007/s10592-005-9062-0. 
  22. 22.0 22.1 Yamaguchi, Nobuyuki; Haddane, B. (2002). "The North African Barbary Lion and the Atlas Lion Project". International Zoo News 49: 465–81. 
  23. 23.0 23.1 Menon, Vivek (2003). A Field Guide to Indian Mammals. Delhi: Dorling Kindersley India. ISBN 0-14-302998-3. 
  24. Schoe, Marjolein; Sogbohossou, Etotépé A.; Kaandorp, Jacques and de Iongh, Hans (2010) Progress Report – collaring operation Pendjari Lion Project, Benin. Funded by the Dutch Zoo Conservation Fund.
  25. Munson, Linda (3 March 2006). "Contraception in felids". Theriogenology 66 (1): 126–34. doi:10.1016/j.theriogenology.2006.03.06. PMID 16626799. 
  26. Koenigswald, Wighart von (2002). Lebendige Eiszeit: Klima und Tierwelt im Wandel (in German). Stuttgart: Theiss. ISBN 3-8062-1734-3. 
  27. 27.0 27.1 27.2 27.3 27.4 27.5 Grisham, Jack (2001). "Lion". In Bell, Catherine E. Encyclopedia of the World's Zoos. 2: G–P. Chofago: Fitzroy Dearborn. pp. 733–39. ISBN 1-57958-174-9. 
  28. McBride, Chris (1977). The White Lions of Timbavati. Johannesburg: E. Stanton. ISBN 0-949997-32-3. 
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