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Lion
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Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Carnivora
Suborder: Feliformia
Family: Felidae
Subfamily: Pantherinae
Genus: Panthera
Species: P. leo[1]
Binomial name
Panthera leo[1]
(Linnaeus, 1758)[3]
Subspecies
P. l. leo
P. l. melanochaita
P. l. sinhaleyus
The lion (Panthera leo) is a species in the family Felidae; it is a muscular, deep-chested cat with
a short, rounded head, a reduced neck and round ears, and a hairy tuft at the end of its tail. It
is sexually dimorphic; male lions have a prominent mane, which is the most recognisable feature
of the species. With a typical head-to-body length of 184–208 cm (72–82 in) they are larger than
females at 160–184 cm (63–72 in). It is a social species, forming groups called prides. A lion
pride consists of a few adult males, related females and cubs. Groups of female lions usually
hunt together, preying mostly on large ungulates. The lion is an apex and keystone predator,
although some lions scavenge when opportunities occur, and have been known to hunt humans,
although the species typically does not.
Typically, the lion inhabits grasslands and savannas but is absent in dense forests. It is usually
more diurnal than other big cats, but when persecuted it adapts to being active at night and at
twilight. In the Pleistocene, the lion ranged throughout Eurasia, Africa and North America but
today it has been reduced to fragmented populations in Sub-Saharan Africa and one critically
endangered population in western India. It has been listed as Vulnerable on the IUCN Red
List since 1996 because populations in African countries have declined by about 43% since the
early 1990s. Lion populations are untenable outside designated protected areas. Although the
cause of the decline is not fully understood, habitat loss and conflicts with humans are the
greatest causes for concern.
One of the most widely recognised animal symbols in human culture, the lion has been
extensively depicted in sculptures and paintings, on national flags, and in contemporary films and
literature. Lions have been kept in menageries since the time of the Roman Empire and have
been a key species sought for exhibition in zoological gardens across the world since the late
18th century. Cultural depictions of lions were prominent in the Upper Paleolithic period; carvings
and paintings from the Lascaux and Chauvet Caves in France have been dated to 17,000 years
ago, and depictions have occurred in virtually all ancient and medieval cultures that coincided
with the lion's former and current ranges.
Contents
1Etymology
2Taxonomy
o 2.1Subspecies
o 2.2Fossil records
o 2.3Evolution
o 2.4Dubious subspecies
o 2.5Hybrids
3Description
o 3.1Mane
o 3.2Colour variation
4Distribution and habitat
o 4.1Historical range
5Behaviour and ecology
o 5.1Group organisation
o 5.2Hunting and diet
5.2.1Predator competition
o 5.3Reproduction and life cycle
o 5.4Health
o 5.5Communication
6Conservation
o 6.1In Africa
o 6.2In Asia
o 6.3Captive breeding
7Interactions with humans
o 7.1In zoos and circuses
o 7.2Hunting and games
o 7.3Man-eating
8Cultural significance
o 8.1Sub-Saharan Africa
o 8.2Near East
o 8.3Far East
o 8.4Western world
9See also
10References
o 10.1Cited texts
11Further reading
12External links
Etymology
The word 'lion' is derived from Latin: leo[4] and Ancient Greek: λέων (leon).[5] The
word lavi (Hebrew: )לָ ִביאmay also be related.[6]
Taxonomy
Two cladograms proposed for Panthera. The upper cladogram is based on the 2006 study,[7][8] and the
lower one on the 2010[9] and 2011[10] studies.
Felis leo was the scientific name used by Carl Linnaeus in 1758, who described the lion in his
work Systema Naturae.[3] The genus name Panthera was coined by German naturalist Lorenz
Oken in 1816.[11] Between the mid-18th and mid-20th centuries, 26 lion specimens were
described and proposed as subspecies, of which 11 were recognised as valid in 2005.[1] They
were distinguished on the basis of appearance, size and colour of mane. Because these
characteristics show much variation between individuals, most of these forms were probably not
true subspecies, especially because they were often based upon museum material with "striking,
but abnormal" morphological characteristics.[12]
Subspecies
In the 19th and 20th centuries, several lion type specimens were described and proposed
as subspecies, with about a dozen recognised as valid taxa until 2017.[1] Between 2008 and
2016, IUCN Red List assessors used only two subspecific names: P. l. leo for African lion
populations and P. l. persica for the Asiatic lion population.[2][13][14] In 2017, the Cat Classification
Task Force of the Cat Specialist Group revised lion taxonomy, and recognises two subspecies
based on results of several phylogeographic studies on lion evolution, namely:[15]
P. l. leo (Linnaeus, 1758) − the nominate lion subspecies includes the Asiatic lion,
the regionally extinct Barbary lion, and lion populations in West and northern parts of Central
Africa.[15] Synonyms include P. l. persica (Meyer, 1826), P. l. senegalensis (Meyer, 1826), P. l.
kamptzi (Matschie, 1900), and P. l. azandica (Allen, 1924).[1] Some authors referred to it as
'Northern lion' and 'northern subspecies'.[16][17]
P. l. melanochaita (Smith, 1842) − includes the extinct Cape lion and lion populations
in East and Southern African regions.[15] Synonyms include P. l. somaliensis (Noack 1891), P. l.
massaica (Neumann, 1900), P. l. sabakiensis (Lönnberg, 1910), P. l. bleyenberghi (Lönnberg,
1914), P. l. roosevelti (Heller, 1914), P. l. nyanzae (Heller, 1914), P. l. hollisteri (Allen, 1924), P. l.
krugeri (Roberts, 1929), P. l. vernayi (Roberts, 1948), and P. l. webbiensis (Zukowsky, 1964).[1][18] It
has been referred to as 'southern subspecies'.[17]
Lion samples from some parts of the Ethiopian Highlands cluster genetically with those from
Cameroon and Chad, while lions from other areas of Ethiopia cluster with samples from East
Africa. Researchers therefore assume that Ethiopia is a contact zone between the two
subspecies.[19]
Fossil records
The skull of an American lion on display at the National Museum of Natural History
Other lion subspecies or sister species to the modern lion existed in prehistoric times:[21]
P. l. sinhaleyus was a fossil carnassial excavated in Sri Lanka, which was attributed to a lion.
It is thought to have become extinct around 39,000 years ago.[22]
P. leo fossilis, P. fossilis or P. spelaea fossilis was larger than the modern lion and lived in
the Middle Pleistocene. Bone fragments were excavated in caves in the United Kingdom,
Germany, Italy and Czech Republic.[23][24]
Panthera spelaea lived in Eurasia and Beringia during the Late Pleistocene. It became
extinct due to climate warming latest by 11,900 years ago.[25] Bone fragments excavated in
European, North Asian, Canadian and Alaskan caves indicate that it ranged from Europe
across Siberia into western Alaska.[26] It likely derived from P. fossilis,[27] and was genetically
isolated and highly distinct from the lion in Africa and Asia.[28][27] It is depicted
in Paleolithic cave paintings, ivory carvings, and clay busts.[29]
P. l. atrox or P. atrox ranged in the Americas from Canada to possibly Patagonia.[30][30] The
American lion arose when a cave lion population in Beringia became isolated south of
the Cordilleran Ice Sheet about 370,000 years ago.[31][32] A fossil from Edmonton dates to
11,355 ± 55 years ago.[33]
Evolution
The maximal range of the modern lion and its prehistoric relatives in the late Pleistocene: red
indicates Panthera spelaea, blue P. atrox, and green P. leo
The lion's closest relatives are the other species of the genus Panthera, namely tiger, snow
leopard, jaguar, and leopard. Results of phylogenetic studies published in 2006 and 2009
indicate that the jaguar and the lion belong to one sister group that diverged about 2.06 million
years ago.[7][8] Results of later studies indicate that the leopard and the lion belong to the same
sister group, which diverged 3.1–1.95 million years ago.[9][10] The geographic origin of
the Panthera is most likely northern Central Asia. The leopard-lion clade was distributed in the
Asian and African Palearctic since at least the Early Pliocene. The clade consisting of lion and
Eurasian cave lion is thought to have diverged in Africa or Holarctic Asia,[34] 2.93–1.23 million
years ago.[27] Hybridisation between lion and snow leopard ancestors, however, may have
continued until about 2.1 million years ago.[35] The Eurasian and American cave lions became
extinct at the end of the last glacial period without mitochondrial descendants on other
continents.[28][36][37]
The modern lion was probably widely distributed in Africa during the Middle Pleistocene and
started to diverge in sub-Saharan Africa during the Late Pleistocene. Lion populations in East
and Southern Africa became separated from populations in West and North Africa when the
equatorial rainforest expanded 183,500–81,800 years ago. The expansion of the Sahara 83,100–
26,600 years ago caused the separation of populations in West and North Africa. As the
rainforest decreased, thus giving rise to more open habitats, lions moved from West to Central
Africa. Lions from North Africa dispersed to southern Europe and Asia 38,800–8,300
ago.[38] Extinction of lions in southern Europe, North Africa and the Middle East interrupted gene
flow between lion populations in Asia and Africa. Genetic evidence revealed
numerous mutations in lion samples from East and Southern Africa, which indicates that this
group has a longer evolutionary history than genetically less diverse lion samples from Asia and
West and Central Africa.[39] Results of phylogeographic research indicate that the two living lion
subspecies diverged about 245,000 years ago.[19]
Dubious subspecies
Description
A tuft at the end of the tail is a distinct characteristic of the lion
The lion is a muscular, deep-chested cat with a short, rounded head, a reduced neck and round
ears. Its fur varies in colour from light buff to silvery grey, yellowish red and dark brown. The
colours of the underparts are generally lighter. A new-born lion has dark spots, which fade as the
cub reaches adulthood, although faint spots often may still be seen on the legs and underparts.
The lion is the only member of the cat family that displays obvious sexual dimorphism. Males
have broader heads and a prominent mane that grows downwards and backwards covering most
of the head, neck, shoulders, and chest. The mane is typically brownish and tinged with yellow,
rust and black hairs.[45][46]
The tail of all lions ends in a dark, hairy tuft that in some lions conceals an approximately 5 mm
(0.20 in)-long, hard "spine" or "spur" that is formed from the final, fused sections of tail bone. The
functions of the spur are unknown. The tuft is absent at birth and develops at around 5 1⁄2 months
of age. It is readily identifiable by the age of seven months.[47]
Of the living felid species, the lion is rivaled only by the tiger in length, weight and height at the
shoulder.[48] Its skull is very similar to that of the tiger, although the frontal region is usually more
depressed and flattened, and has a slightly shorter postorbital region and broader nasal
openings than those of the tiger. Due to the amount of skull variation in the two species, usually
only the structure of the lower jaw can be used as a reliable indicator of species.[49][50]
The size and weight of adult lions varies across global range and habitats.[51][52][53][54] Accounts of a
few individuals that were larger than average exist from Africa and India.[45][55][56][57]
Head-and-body
160–184 cm (63–72 in)[58] 184–208 cm (72–82 in)[58]
length
Mane
The lion's mane is the most recognisable feature of the species.[18] It starts growing when lions
are about a year old. Mane colour varies and darkens with age; research shows its colour and
size are influenced by environmental factors such as average ambient temperature. Mane length
apparently signals fighting success in male–male relationships; darker-maned individuals may
have longer reproductive lives and higher offspring survival, although they suffer in the hottest
months of the year. The presence, absence, colour and size of the mane are associated with
genetic precondition, sexual maturity, climate and testosterone production; the rule of thumb is
that a darker, fuller mane indicates a healthier animal. In Serengeti National Park, female lions
favour males with dense, dark manes as mates.[59][60] The main purpose of the mane is thought be
the protection of the neck and throat in territorial fights with rivals.[61] Cool ambient temperature in
European and North American zoos may result in a heavier mane.[62] Asiatic lions usually have
sparser manes than average African lions.[63]
Almost all male West African lions in Pendjari National Park are either maneless or have very
short manes.[64] Maneless lions have also been reported in Senegal, in Sudan's Dinder National
Park and in Tsavo East National Park, Kenya.[65] The original male white lion from Timbavati in
South Africa was also maneless. The hormone testosterone has been linked to mane
growth; castrated lions often have little to no mane because the removal of the gonads inhibits
testosterone production.[66] Increased testosterone may be the cause of maned lionesses
reported in northern Botswana.[67]
The lion's mane may have evolved around 320,000–190,000 years ago.[68] Cave paintings of
extinct Eurasian cave lions almost exclusively show hunting animals without a mane; some
suggest that this is evidence that they were indeed maneless.[69]
Colour variation
Further information: White lion
African lions live in scattered populations across Sub-Saharan Africa.The lion prefers grassy
plains and savannahs, scrub bordering rivers and open woodlands with bushes. It is absent
from rainforest and rarely enters closed forest. On Mount Elgon, the lion has been recorded up to
an elevation of 3,600 m (11,800 ft) and close to the snow line on Mount Kenya.[45] Lions occur in
savannah grasslands with scattered acacia trees, which serve as shade.[73] The Asiatic lion now
only survives in and around Gir Forest National Park in Gujarat, western India. Its habitat is a
mixture of dry savannah forest and very dry, deciduous scrub forest.[13]
Historical range
In Africa, the range of the lion originally spanned most of the central rainforest zone and
the Sahara desert.[74] In the 1960s, it became extinct in North Africa, except in the southern part
of Sudan.[75][76][77] In Eurasia, the lion once ranged from India to Europe; a Bronze Age statue of a
lion from either southern Italy or southern Spain from c. 1000–1200 years BC, the "Mari-Cha
Lion", was put on display at the Louvre Abu Dhabi;[78] Herodotus reported that lions had been
common in Greece in 480 BC; they attacked the baggage camels of the Persian king Xerxes on
his march through the country. Aristotle considered them rare by 300 BC, and by 100 AD, they
had been extirpated.[74] Until the 10th century, lions survived in the Caucasus, their last European
outpost.[50] The species was eradicated in Palestine by the Middle Ages, and from most of the
rest of Asia after the arrival of readily available firearms in the 18th century. Between the late
19th and late 20th centuries, it became extinct in Southwest Asia. By the late 19th century, the
lion had been extirpated in most of northern India and Turkey.[79][80] The last live lion in Iran was
sighted in 1942, about 65 km (40 mi) northwest of Dezful.[81] The corpse of a lioness was found
on the banks of the Karun river in Khūzestān Province in 1944. No subsequent reliable records
exist from Iran.[82]
Lions spend much of their time resting; they are inactive for about 20 hours per day.[83] Although
lions can be active at any time, their activity generally peaks after dusk with a period of
socialising, grooming and defecating. Intermittent bursts of activity continue until dawn, when
hunting most often takes place. They spend an average of two hours a day walking and
50 minutes eating.[84]
Group organisation
The lion is the most social of all wild felid species, living in groups of related individuals with their
offspring. Such a group is called a "pride". Groups of male lions are called
"coalitions".[85] Females form the stable social unit in a pride and do not tolerate outside
females.[86] Membership only changes with the births and deaths of lionesses,[87] although some
females leave and become nomadic.[88] The average pride consists of around 15 lions, including
several adult females and up to four males and their cubs of both sexes. Large prides, consisting
of up to 30 individuals, have been observed.[89] The sole exception to this pattern is the Tsavo
lion pride that always has just one adult male.[90] Male cubs are excluded from their maternal
pride when they reach maturity at around two or three years of age.[88]
Some lions are "nomads" that range widely and move around sporadically, either in pairs or
alone.[85] Pairs are more frequent among related males who have been excluded from their birth
pride. A lion may switch lifestyles; nomads can become residents and vice versa.[91] Interactions
between prides and nomads tend to be hostile, although pride females in estrus allow nomadic
males to approach them.[92] Males spend years in a nomadic phase before gaining residence in a
pride.[93] A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain
residency at between 3.5 and 7.3 years of age.[94] In Kruger National Park, dispersing male lions
move more than 25 km (16 mi) away from their natal pride in search of their own territory.
Females lions stay closer to their natal pride. Therefore, female lions in an area are more closely
related to each other than male lions in the same area.[95]
The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a
"range".[85] Males associated with a pride tend to stay on the fringes, patrolling their territory. The
reasons for the development of sociality in lionesses – the most pronounced in any cat species –
are the subject of much debate. Increased hunting success appears to be an obvious reason, but
this is uncertain upon examination; coordinated hunting allows for more successful predation but
also ensures non-hunting members reduce per capita calorific intake. Some females, however,
take a role raising cubs that may be left alone for extended periods. Members of the pride tend to
regularly play the same role in hunts and hone their skills. The health of the hunters is the
primary need for the survival of the pride; hunters are the first to consume the prey at the site it is
taken. Other benefits include possible kin selection; sharing food within the family; protecting the
young, maintaining territory and individual insurance against injury and hunger.[55]
Both males and females defend the pride against intruders, but the male lion is better-suited for
this purpose due to its stockier, more powerful build. Some individuals consistently lead the
defence against intruders, while others lag behind.[96] Lions tend to assume specific roles in the
pride; slower-moving individuals may provide other valuable services to the
group.[97] Alternatively, there may be rewards associated with being a leader that fends off
intruders; the rank of lionesses in the pride is reflected in these responses.[98] The male or males
associated with the pride must defend their relationship with the pride from outside males who
may attempt to usurp them.
Asiatic lion prides differ in group composition. Male Asiatic lions are solitary or associate with up
to three males, forming a loose pride while females associate with up to 12 other females,
forming a stronger pride together with their cubs. Female and male lions associate only when
mating.[99] Coalitions of males hold territory for a longer time than single lions. Males in coalitions
of three or four individuals exhibit a pronounced hierarchy, in which one male dominates the
others and mates more frequently.[100]
Hunting and diet
The lion is a generalist hypercarnivore[101] and is considered to be both an apex and keystone
predator due to its wide prey spectrum.[102] Its prey consists mainly of mammals –
particularly ungulates – weighing 190–550 kg (420–1,210 lb) with a preference for blue
wildebeest, plains zebra, African buffalo, gemsbok and giraffe. Lions also hunt common
warthog depending on availability, although the species is below the preferred weight range.[103] In
India, sambar deer and chital are the most commonly recorded wild prey,[46][103][104] while domestic
livestock may contribute significantly to their diet.[104] They usually avoid fully grown
adult elephants, rhinoceroses and hippopotamus, as well as small prey like dik-
dik, hyrax, hare and vervet monkey.[105] Unusual prey items include porcupines and small reptiles.
Lions kill other predators such as leopard, cheetah and spotted hyena but seldom consume
them.[106]
Young lions first display stalking behaviour at around three months of age, although they do not
participate in hunting until they are almost a year old and begin to hunt effectively when nearing
the age of two.[107] Single lions are capable of bringing down prey much larger than themselves,
such as zebra and wildebeest, while larger prey like buffalo and giraffes are riskier.[91] In Chobe
National Park, large prides have been observed hunting subadult and even adult African bush
elephants.[108][109] In typical hunts, each lioness has a favoured position in the group, either stalking
prey on the "wing", then attacking, or moving a smaller distance in the centre of the group and
capturing prey fleeing from other lionesses. Males attached to prides do not usually participate in
group hunting.[110] Some evidence suggests, however, that males are just as successful as
females; they are typically solo hunters who ambush prey in small bushland.[111]
Lions are not particularly known for their stamina; for instance, a lioness' heart comprises only
0.57% of her body weight and a male's is about 0.45% of his body weight, whereas a hyena's
heart comprises almost 1% of its body weight.[112] Thus, lions only run quickly in short
bursts[113] and need to be close to their prey before starting the attack. They take advantage of
factors that reduce visibility; many kills take place near some form of cover or at night.[114] The
lion's attack is short and powerful; they attempt to catch prey with a fast rush and final leap, and
usually kill prey by strangulation,[115] which can cause cerebral ischemia or asphyxia and results
in hypoxaemia or hypoxia. They also kill prey by enclosing its mouth and nostrils in their jaws,
which also results in asphyxia.[54]
Lions typically consume prey at the location of the hunt but sometimes drag large prey into
cover.[116] They tend to squabble over kills, particularly the males. Cubs suffer most when food is
scarce but otherwise all pride members eat their fill, including old and crippled lions, which can
live on leftovers.[91] Large kills are shared more widely among pride members.[117] An adult lioness
requires an average of about 5 kg (11 lb) of meat per day while males require about 7 kg
(15 lb).[118] Lions gorge themselves and eat up to 30 kg (66 lb) in one session;[82] if it is unable to
consume all of the kill, it rests for a few hours before continuing to eat. On hot days, the pride
retreats to shade with one or two males standing guard.[116] Lions defend their kills from
scavengers such as vultures and hyenas.[91]
Lions scavenge on carrion when the opportunity arises; they scavenge animals dead from
natural causes such as disease or those that were killed by other predators. Scavenging lions
keep a constant lookout for circling vultures, which indicate the death or distress of an
animal.[119] Most carrion on which both hyenas and lions feed upon are killed by hyenas rather
than lions.[54] Carrion is thought to provide a large part of lion diet.[120]
Predator competition
Lions and spotted hyenas occupy a similar ecological niche and where they coexist they
compete for prey and carrion; a review of data across several studies indicates a dietary overlap
of 58.6%.[121] Lions typically ignore spotted hyenas unless the lions are on a kill or are being
harassed by the hyenas, while the latter tend to visibly react to the presence of lions, with or
without the presence of food. Lions seize the kills of spotted hyenas; in the Ngorongoro crater it
is common for lions to subsist largely on kills stolen from hyenas, causing the hyenas to increase
their kill rate.[122] In Botswana's Chobe National Park, the situation is reversed; hyenas frequently
challenge lions and steal their kills, obtaining food from 63% of all lion kills.[123] When confronted
on a kill by lions, spotted hyenas may either leave or wait patiently at a distance of 30–100 m
(100–330 ft) until the lions have finished.[124] Hyenas are bold enough to feed alongside lions and
to force the lions off a kill. The two species attack one another even when there is no food
involved for no apparent reason.[125][126] Lion predation can account for up to 71% of hyena deaths
in Etosha National Park. Spotted hyenas have adapted by frequently mobbing lions that enter
their territories.[127] When the lion population in Kenya's Masai Mara National Reserve declined,
the spotted hyena population increased rapidly.[128] Experiments on captive spotted hyenas show
that specimens without prior experience with lions act indifferently to the sight of them, but will
react fearfully to lion scent.[122]
Lions tend to dominate cheetahs and leopards, steal their kills and kill their cubs and even adults
when given the chance.[129] Cheetahs in particular often lose their kills to lions or other
predators.[130] A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125
cheetah cubs born between 1987 and 1990.[131] Cheetahs avoid their competitors by using
different temporal and habitat niches.[132] Leopards are able to take refuge in trees; lionesses,
however, occasionally attempt to retrieve leopard kills from trees.[133] Lions similarly
dominate African wild dogs, taking their kills and preying on young and rarely adult dogs.
Population densities of wild dogs are low in areas where lions are more abundant.[134] However,
there are a few reported cases of old and wounded lions falling prey to wild dogs.[135][136] Lions also
charge at Nile crocodiles; depending on the size of the crocodile and the lion, either can lose kills
or carrion to the other. Lions have been observed killing crocodiles that ventured onto
land.[137] Lions also enter waterways, evidenced by the occasional lion claw found in crocodile
stomachs.[138]
Reproduction and life cycle
Most lionesses reproduce by the time they are four years of age.[139] Lions do not mate at a
specific time of year and the females are polyestrous.[140] Like those of other cats, the male lion's
penis has spines that point backward. During withdrawal of the penis, the spines rake the walls of
the female's vagina, which may cause ovulation.[141][142] A lioness may mate with more than one
male when she is in heat.[143] Generation length of the lion is about seven years.[144] The average
gestation period is around 110 days;[140] the female gives birth to a litter of between one and four
cubs in a secluded den, which may be a thicket, a reed-bed, a cave, or some other sheltered
area, usually away from the pride. She will often hunt alone while the cubs are still helpless,
staying relatively close to the den.[145] Lion cubs are born blind – their eyes open around seven
days after birth. They weigh 1.2–2.1 kg (2.6–4.6 lb) at birth and are almost helpless, beginning to
crawl a day or two after birth and walking around three weeks of age.[146] To avoid a buildup of
scent attracting the attention of predators, the lioness moves her cubs to a new den site several
times a month, carrying them one-by-one by the nape of the neck.[145]
Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are
six to eight weeks old.[145] Sometimes this introduction to pride life occurs earlier, particularly if
other lionesses have given birth at about the same time.[91][147] When first introduced to the rest of
the pride, lion cubs lack confidence when confronted with adults other than their mother. They
soon begin to immerse themselves in the pride life, however, playing among themselves or
attempting to initiate play with the adults.[147] Lionesses with cubs of their own are more likely to
be tolerant of another lioness's cubs than lionesses without cubs. Male tolerance of the cubs
varies – sometimes a male will patiently let the cubs play with his tail or his mane, whereas
another may snarl and bat the cubs away.[148]
Video of a lioness and her cubs in Phinda Private Game Reserve, South Africa
Pride lionesses often synchronise their reproductive cycles and communal rearing and suckling
of the young, which suckle indiscriminately from any or all of the nursing females in the pride.
The synchronisation of births is advantageous because the cubs grow to being roughly the same
size and have an equal chance of survival, and sucklings are not dominated by older
cubs.[91][147] Weaning occurs after six or seven months. Male lions reach maturity at about three
years of age and at four to five years are capable of challenging and displacing adult males
associated with another pride. They begin to age and weaken at between 10 and 15 years of age
at the latest.[149]
When one or more new males oust the previous males associated with a pride, the victors
often kill any existing young cubs, perhaps because females do not become fertile and receptive
until their cubs mature or die. Females often fiercely defend their cubs from a usurping male but
are rarely successful unless a group of three or four mothers within a pride join forces against the
male.[150] Cubs also die from starvation and abandonment, and predation by leopards, hyenas
and wild dogs.[136][91] Up to 80% of lion cubs will die before the age of two.[151] Both male and
female lions may be ousted from prides to become nomads, although most females usually
remain with their birth pride. When a pride becomes too large, however, the youngest generation
of female cubs may be forced to leave to find their own territory. When a new male lion takes
over a pride, adolescent lions – both male and female – may be evicted.[152] Lions of both sexes
may be involved in group homosexual and courtship activities; males will also head-rub and roll
around with each other before simulating sex together.[153][154]
Health
Although adult lions have no natural predators, evidence suggests most die violently from attacks
by humans or other lions.[155] Lions often inflict serious injuries on members of other prides they
encounter in territorial disputes or members of the home pride when fighting at a kill.[156] Crippled
lions and cubs may fall victim to hyenas and leopards or be trampled by buffalo or elephants.
Careless lions may be maimed when hunting prey.[157]
Ticks commonly infest the ears, neck and groin regions of lions.[158][159] Adult forms of several
species of the tapeworm genus Taenia have been isolated from lion intestines, having been
ingested as larvae in antelope meat.[160] Lions in the Ngorongoro Crater were afflicted by an
outbreak of stable fly (Stomoxys calcitrans) in 1962; this resulted in lions becoming emaciated
and covered in bloody, bare patches. Lions sought unsuccessfully to evade the biting flies by
climbing trees or crawling into hyena burrows; many died or migrated and the local population
dropped from 70 to 15 individuals.[161] A more recent outbreak in 2001 killed six lions.[162]
Captive lions have been infected with canine distemper virus (CDV) since at least the mid
1970s.[163] CDV is spread by domestic dogs and other carnivores; a 1994 outbreak in Serengeti
National Park resulted in many lions developing neurological symptoms such as seizures. During
the outbreak, several lions died from pneumonia and encephalitis.[164] Feline immunodeficiency
virus and lentivirus also affect captive lions.[165][166]
Communication
When resting, lion socialisation occurs through a number of behaviours; the animal's expressive
movements are highly developed. The most common peaceful, tactile gestures are head rubbing
and social licking,[167] which have been compared with the role of allogrooming among
primates.[168] Head rubbing – the nuzzling of the forehead, face and neck against another lion –
appears to be a form of greeting[169] and is seen often after an animal has been apart from others
or after a fight or confrontation. Males tend to rub other males, while cubs and females rub
females.[170] Social licking often occurs in tandem with head rubbing; it is generally mutual and the
recipient appears to express pleasure. The head and neck are the most common parts of the
body licked; this behaviour may have arisen out of utility because lions cannot lick these areas
themselves.[171]
Lion roar
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A captive lion roaring
Lions have an array of facial expressions and body postures that serve as visual gestures.[172] A
common facial expression is the "grimace face" or flehmen response, which a lion makes when
sniffing chemical signals and involves an open mouth with bared teeth, raised muzzle, wrinkled
nose closed eyes and relaxed ears.[173] Lions also use chemical and visual marking; males
will spray and scrape plots of ground and objects within the territory.[172]
The lion's repertoire of vocalisations is large; variations in intensity and pitch appear to be central
to communication. Most lion vocalisations are variations of growling, snarling, meowing and
roaring. Other sounds produced include purring, puffing, bleating and humming. Roaring is used
to advertise its presence. Lions most often roar at night, a sound that can be heard from a
distance of 8 kilometres (5.0 mi).[174] They tend to roar in a very characteristic manner starting
with a few deep, long roars that subside into a series of shorter ones.[175][176]
Conservation
The lion is listed as Vulnerable on the IUCN Red List.[2]
In Africa
Several large and well-managed protected areas in Africa host large lion populations. Where an
infrastructure for wildlife tourism has been developed, cash revenue for park management and
local communities is a strong incentive for lion conservation.[2] Most lions now live in East and
Southern Africa; their numbers are rapidly decreasing, and fell by an estimated 30–50% in the
late half of the 20th century. Primary causes of the decline include disease and human
interference.[2] In 1975, it was estimated that since the 1950s, lion numbers had decreased by
half to 200,000 or fewer.[177] Estimates of the African lion population range between 16,500 and
47,000 living in the wild in 2002–2004.[178][75]
In the Republic of the Congo, Odzala-Kokoua National Park was considered a lion stronghold in
the 1990s. By 2014, no lions were recorded in the protected area so the population is considered
locally extinct.[179] The West African lion population is isolated from the one in Central Africa, with
little or no exchange of breeding individuals. In 2015, it was estimated that this population
consists of about 400 animals, including fewer than 250 mature individuals. They persist in three
protected areas in the region, mostly in one population in the WAP protected area complex,
shared by Benin, Burkina Faso and Niger. This population is listed as Critically
Endangered.[14] Field surveys in the WAP ecosystem revealed that lion occupancy is lowest in the
W National Park, and higher in areas with permanent staff and thus better protection.[180] A
population occurs in Cameroon's Waza National Park, where between approximately 14 and 21
animals persisted as of 2009.[181] In addition, 50 to 150 lions are estimated to be present in
Burkina Faso's Arly-Singou ecosystem.[182] In 2015, an adult male lion and a female lion were
sighted in Ghana's Mole National Park. These were the first sightings of lions in the country in 39
years.[183]
In 2005, Lion Conservation Strategies were developed for West and Central Africa, and or East
and Southern Africa. The strategies seek to maintain suitable habitat, ensure a sufficient wild
prey base for lions, reduce factors that lead to further fragmentation of populations, and make
lion–human coexistence sustainable.[184][185] Lion depredation on livestock is significantly reduced
in areas where herders keep livestock in improved enclosures. Such measures contribute to
mitigating human–lion conflict.[186] In 2015, a population of up to 200 lions that was previously
thought to have been extirpated was filmed in the Alatash National Park, Ethiopia, close to the
Sudanese border.[187][188]
In Asia
A lioness in Gir Forest National Park
The last refuge of the Asiatic lion population is the 1,412 km2 (545 sq mi) Gir Forest National
Park and surrounding areas in the region of Saurashtra or Kathiawar
Peninsula in Gujarat State, India. The population has risen from approximately 180 lions in 1974
to about 400 in 2010.[189] It is geographically isolated, which can lead to inbreeding and
reduced genetic diversity. Since 2008, the Asiatic lion has been listed as Endangered on
the IUCN Red List.[13] By 2015, the population had grown to 523 individuals inhabiting an area of
7,000 km2 (2,700 sq mi) in Saurashtra.[190][191][192] The Asiatic Lion Census conducted in 2017
recorded about 650 individuals.[193]
The presence of numerous human habitations close to the National Park results in conflict
between lions, local people and their livestock.[194][190] Some consider the presence of lions a
benefit, as they keep populations of crop damaging herbivores in check.[195] The establishment of
a second, independent Asiatic lion population in Kuno Wildlife Sanctuary, located in Madhya
Pradesh was planned but in 2017, the Asiatic Lion Reintroduction Project seemed unlikely to be
implemented.[196][197]
Captive breeding
Two captive male Asiatic lions in Sanjay Gandhi National Park, India
Lions imported to Europe before the middle of the 19th century were possibly foremost Barbary
lions from North Africa, or Cape lions from Southern Africa.[198] Another 11 animals thought to be
Barbary lions kept in Addis Ababa Zoo are descendants of animals owned by Emperor Haile
Selassie. WildLink International in collaboration with Oxford University launched an ambitious
International Barbary Lion Project with the aim of identifying and breeding Barbary lions in
captivity for eventual reintroduction into a national park in the Atlas Mountains of
Morocco.[199] However, a genetic analysis showed that the captive lions at Addis Ababa Zoo were
not Barbary lions, but rather closely related to wild lions in Chad and Cameroon.[200]
In 1982, the Association of Zoos and Aquariums started a Species Survival Plan for the Asiatic
lion to increase its chances of survival. In 1987, it was found that most lions in North American
zoos were hybrids between African and Asiatic lions.[201] Breeding programs need to note origins
of the participating animals to avoid cross-breeding different subspecies and thus reducing their
conservation value.[202] Captive breeding of lions was halted to eliminate individuals of unknown
origin and pedigree. Wild-born lions were imported to American zoos from Africa between 1989
and 1995. Breeding was continued in 1998 in the frame of an African lion Species Survival
Plan.[203]
About 77% of the captive lions registered in the International Species Information System in 2006
were of unknown origin; these animals might have carried genes that are extinct in the wild and
may therefore be important to the maintenance of the overall genetic variability of the lion.[62]
Lion in Melbourne Zoo enjoying an elevated grassy area with some tree shelter
Lions are part of a group of exotic animals that have been central to zoo exhibits since the late
18th century. Although many modern zoos are more selective about their exhibits,[204] there are
more than 1,000 African and 100 Asiatic lions in zoos and wildlife parks around the world. They
are considered an ambassador species and are kept for tourism, education and conservation
purposes.[205] Lions can reach an age of over 20 years in captivity; Apollo, a resident lion
of Honolulu Zoo in Honolulu, Hawaii, died at age 22 in August 2007. His two sisters, born in
1986, were still alive in August 2007.[206]
The first European "zoos" spread among noble and royal families in the 13th century, and until
the 17th century were called seraglios; at that time they came to be called menageries, an
extension of the cabinet of curiosities. They spread from France and Italy during
the Renaissance to the rest of Europe.[207] In England, although the seraglio tradition was less
developed, lions were kept at the Tower of London in a seraglio established by King John in the
13th century;[208][209] this was probably stocked with animals from an earlier menagerie started in
1125 by Henry I at his hunting lodge in Woodstock, Oxfordshire, where according to William of
Malmesbury lions had been stocked.[210]
Lions were kept in cramped and squalid conditions at London Zoo until a larger lion house with
roomier cages was built in the 1870s.[211] Further changes took place in the early 20th century
when Carl Hagenbeck designed enclosures with concrete "rocks", more open space and a moat
instead of bars, more closely resembling a natural habitat. Hagenbeck designed lion enclosures
for both Melbourne Zoo and Sydney's Taronga Zoo; although his designs were popular, the use
of bars and caged enclosures prevailed in many zoos until the 1960s.[212] In the late 20th century,
larger, more natural enclosures and the use of wire mesh or laminated glass instead of lowered
dens allowed visitors to come closer than ever to the animals; some attractions such as the Cat
Forest/Lion Overlook of Oklahoma City Zoological Park placed the den on ground level, higher
than visitors.[12]
Lion taming has been part of both established circuses and individual acts such as Siegfried &
Roy. The practice began in the early 19th century by Frenchman Henri Martin and
American Isaac Van Amburgh, who both toured widely and whose techniques were copied by a
number of followers.[213] Van Amburgh performed before Queen Victoria in 1838 when he toured
Great Britain. Martin composed a pantomime titled Les Lions de Mysore ("the lions of Mysore"),
an idea that Amburgh quickly borrowed. These acts eclipsed equestrianism acts as the central
display of circus shows and entered public consciousness in the early 20th century with cinema.
In demonstrating the superiority of human over animal, lion taming served a purpose similar to
animal fights of previous centuries.[213] The ultimate proof of a tamer's dominance and control
over a lion is demonstrated by the placing of the tamer's head in the lion's mouth. The now-iconic
lion tamer's chair was possibly first used by American Clyde Beatty (1903–1965).[214]
Hunting and games
Main article: Lion hunting
See also: Lion baiting
Bas-relief of a wounded lioness from Nineveh, ca. 645–635 BC during the Neo-Assyrian period
Lion hunting has occurred since ancient times and was often a royal pastime; intended to
demonstrate the power of the king over nature. The earliest surviving record of lion hunting is
an ancient Egyptian inscription dated circa 1380 BC that mentions Pharaoh Amenhotep III killing
102 lions "with his own arrows" during the first ten years of his rule. The Assyrians would release
captive lions in a reserved space for the king to hunt; this event would be watched by spectators
as the king and his men, on horseback or chariots, killed the lions with arrows and spears. Lions
were also hunted during the Mughal Empire, where Emperor Jahangir is said to have excelled at
it.[215] In Ancient Rome, lions were kept by emperors for hunts as well as gladiator
fights and executions.[216]
The Maasai people have traditionally viewed the killing of lions as a rite of passage. Historically,
lions were hunted by individuals, however, due to reduced lion populations, elders discourage
solo lion hunts.[217] During the European colonisation of Africa in the 19th century, the hunting of
lions was encouraged because they were considered as vermin and lion hides
fetched £1 each.[218] The widely reproduced imagery of the heroic hunter chasing lions would
dominate a large part of the century.[219] Trophy hunting of lions in recent years has been met with
controversy; notably with the killing of Cecil the lion in mid-2015.[220]
Man-eating
The Tsavo maneaters of East Africa on display in the Field Museum of Natural History
Lions do not usually hunt humans but some – usually males – seem to seek them out. One well-
publicised case is the Tsavo maneaters; in 1898, 28 officially recorded railway workers building
the Kenya-Uganda Railway were taken by lions over nine months during the construction of a
bridge over the Tsavo River in Kenya.[221] The hunter who killed the lions wrote a book detailing
the animals' predatory behaviour; they were larger than normal and lacked manes, and one
seemed to suffer from tooth decay. The infirmity theory, including tooth decay, is not favoured by
all researchers; an analysis of teeth and jaws of man-eating lions in museum collections
suggests that while tooth decay may explain some incidents, prey depletion in human-dominated
areas is a more likely cause of lion predation on humans.[222]
In their analysis of man-eating – including the Tsavo incident – Kerbis Peterhans and Gnoske
acknowledge that sick or injured animals may be more prone to man-eating but that the
behaviour is "not unusual, nor necessarily 'aberrant'" where the opportunity exists; if inducements
such as access to livestock or human corpses are present, lions will regularly prey upon human
beings. The authors note the relationship is well-attested among other pantherines and primates
in the fossil record.[223]
The lion's proclivity for man-eating has been systematically examined. American and Tanzanian
scientists report that man-eating behaviour in rural areas of Tanzania increased greatly from
1990 to 2005. At least 563 villagers were attacked and many eaten over this period – a number
far exceeding the Tsavo attacks. The incidents occurred near Selous National Park in Rufiji
District and in Lindi Province near the Mozambican border. While the expansion of villages into
bush country is one concern, the authors argue conservation policy must mitigate the danger
because in this case, conservation contributes directly to human deaths. Cases in Lindi in which
lions seize humans from the centres of substantial villages have been documented.[224] Another
study of 1,000 people attacked by lions in southern Tanzania between 1988 and 2009 found that
the weeks following the full moon, when there was less moonlight, were a strong indicator of
increased night-time attacks on people.[225]
According to Robert R. Frump, Mozambican refugees regularly crossing Kruger National Park,
South Africa, at night are attacked and eaten by lions; park officials have said man-eating is a
problem there. Frump said thousands may have been killed in the decades
after apartheid sealed the park and forced refugees to cross the park at night. For nearly a
century before the border was sealed, Mozambicans had regularly crossed the park in daytime
with little harm.[226]
Cultural significance
Upper Paleolithic cave painting depicting lions, found in the Chauvet Cave, France.[227]
Granite statue of the Egyptian goddess Sekhmet from the Luxor Temple, dated 1403–1365 BC, exhibited in
the National Museum of Denmark
Roaring and striding lion from the Throne Room of Nebuchadnezzar II, 6th century BC, from Babylon, Iraq
The Lion Capital of Ashoka, which has been a emblem of India
Dorothy meets the Cowardly Lion, from the first edition of The Wonderful Wizard of Oz. Art by W. W.
Denslow
See also
Cats portal
Mammals portal
Animals portal
Biology portal
Africa portal
India portal
Tiger versus lion
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Cited texts
Baratay, E.; Hardouin-Fugier, E. (2002). Zoo: a history of zoological gardens in the West. London:
Reaktion Books. ISBN 978-1-86189-111-2.
Blunt, W. (1975). The Ark in the Park: The Zoo in the Nineteenth Century. London: Hamish
Hamilton. ISBN 978-0-241-89331-9.
de Courcy, C. (1995). The Zoo Story. Ringwood, Victoria: Penguin Books. ISBN 978-0-14-023919-5.
Jackson, D. (2010). Lion. London: Reaktion Books. ISBN 978-1861896551.
Schaller, G. B. (1972). The Serengeti lion: A study of predator–prey relations. Chicago: University of
Chicago Press. ISBN 978-0-226-73639-6.
Scott, J.; Scott, A. (2002). Big Cat Diary: Lion. New York: Harper Collins. ISBN 9780007146666.
Further reading
Schnitzler, A.; Hermann, L. (19 August 2019). "Chronological distribution of the
tiger Panthera tigris and the Asiatic lion Panthera leo persica in their common range in
Asia". Mammal Review. Wiley Online Library. doi:10.1111/mam.12166. Retrieved 23
September 2019.
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