Lion

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For other uses, see Lion (disambiguation).

Lion

Temporal range: Pleistocene–

Present 

PreꞒ

Pg

N
↓

Male lion in Okonjima, Namibia

 Female (lioness) in Okonjima

Conservation status

Vulnerable (IUCN 3.1) [2]

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. fossilis
P. l. sinhaleyus

Historical and

present

distribution of the

lion

in Africa, Asia an

d Europe

The lion (Panthera leo) is a large felid of the genus Panthera native mainly to Africa. It

has a muscular, deep-chested body, short, rounded head, round ears, and a hairy tuft at
the end of its tail. It is sexually dimorphic; adult male lions have a prominent mane. With
a typical head-to-body length of 184 to 208 cm (72 to 82 in) they are larger than females
at 160 to 184 cm (63 to 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, 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 wild cats, but when persecuted, it adapts to being
active at night and at twilight. During the Neolithic period, the lion ranged throughout
Africa, Southeast Europe, and Western and South Asia, but it has been reduced to
fragmented populations in sub-Saharan Africa and one 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 Ancient Egypt, and depictions have occurred in virtually all ancient and medieval
cultures in the lion's historic and current range.

Contents

 1Etymology
 2Taxonomy
o 2.1Subspecies
o 2.2Fossil records
o 2.3Evolution
o 2.4Hybrids
 3Description
o 3.1Size
o 3.2Mane
o 3.3Colour 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 10.1Books
 11External links

Etymology
The word 'lion' is derived from Latin: leo[4] and Ancient Greek: λέων (leon).[5] The
word lavi (Hebrew: ‫ ) ָלבִיא‬may also be related.[6] The generic name Panthera is traceable
to the classical Latin word 'panthēra' and the ancient Greek word πάνθηρ 'panther'.
[7]
 Panthera is phonetically similar to the Sanskrit word पाण्डर pând-ara meaning 'pale
yellow, whitish, white'.[8]

Taxonomy

The upper cladogram is based on the 2006 study, [9][10] the lower one on the 2010[11] and 2011[12] 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 Lorenz
Oken in 1816.[13] 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 mostly by the size and colour of their manes and skins. [14]
Subspecies
Range map showing distribution of subspecies and clades

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][15][16] 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:[17]

 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.[17] Synonyms include P. l. persica (Meyer, 1826), P. l.
senegalensis (Meyer, 1826), P. l. kamptzi (Matschie, 1900), and P. l. azandica (Allen, 1924).
[1]
 Multiple authors referred to it as 'northern lion' and 'northern subspecies'. [18][19]
 P. l. melanochaita (Smith, 1842) − includes the extinct Cape lion and lion
populations in East and Southern African regions.[17] 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][14] It has been referred to as 'southern
subspecies' and 'southern lion'.[19]
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 Ethiopia is a contact zone between the
two subspecies.[20] Genome-wide data of a wild-born historical lion sample from Sudan
showed that it clustered with P. l. leo in mtDNA-based phylogenies, but with a high
affinity to P. l. melanochaita. This result suggested that the taxonomic position of lions in
Central Africa may require revision. [21]
Fossil records
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: [22]

 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. [23]
 P. leo 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.[24][25]
 P. spelaea, or the cave lion, lived in Eurasia and Beringia during the Late
Pleistocene. It became extinct due to climate warming or human expansion latest by
11,900 years ago.[26] Bone fragments excavated in European, North Asian, Canadian
and Alaskan caves indicate that it ranged from Europe across Siberia into western
Alaska.[27] It likely derived from P. fossilis,[28] and was genetically isolated and highly
distinct from the modern lion in Africa and Eurasia. [29][28] It is depicted
in Paleolithic cave paintings, ivory carvings, and clay busts. [30]
 P. atrox, or the American lion, ranged in the Americas from Canada to
possibly Patagonia.[31] It arose when a cave lion population in Beringia became
isolated south of the Cordilleran Ice Sheet about 370,000 years ago.[32][33] A fossil
from Edmonton dates to 11,355 ± 55 years ago.[34]
Evolution

red Panthera spelaea
blue P. atrox
green P. leo

Maximal range of the modern lion

and its prehistoric relatives
in the late Pleistocene
The Panthera lineage is estimated to have genetically diverged from the common
ancestor of the Felidae around 9.32 to 4.47 million years ago to 11.75 to 0.97 million
years ago,[9][35][36] and the geographic origin of the genus is most likely northern Central
Asia.[37] Results of analyses differ in the phylogenetic relationship of the lion; it was
thought to form a sister group with the jaguar (P. onca) that diverged 3.46 to 1.22 million
years ago,[9] but also with the leopard (P. pardus) that diverged 3.1 to 1.95 million years
ago[11][12] to 4.32 to 0.02 million years ago. Hybridisation between lion and snow
leopard (P. uncia) ancestors possibly continued until about 2.1 million years ago. [36] The
lion-leopard clade was distributed in the Asian and African Palearctic since at least the
early Pliocene.[37] The earliest fossils recognisable as lions were found at Olduvai
Gorge in Tanzania and are estimated to be up to 2 million years old. [35]
Estimates for the divergence time of the modern and cave lion lineages range from
529,000 to 392,000 years ago based on mutation rate per generation time of the
modern lion. There is no evidence for gene flow between the two lineages, indicating
that they did not share the same geographic area. [21] The Eurasian and American cave
lions became extinct at the end of the last glacial
period without mitochondrial descendants on other continents.[29][38][39] 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 to 81,800 years ago. [40] They shared a common
ancestor probably between 98,000 and 52,000 years ago. [21] Due to the expansion of the
Sahara between 83,100 and 26,600 years ago, lion populations in West and North
Africa became separated. As the rainforest decreased and thus gave rise to more open
habitats, lions moved from West to Central Africa. Lions from North Africa dispersed to
southern Europe and Asia between 38,800 and 8,300 years ago. [40]
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. [41] A whole genome-wide sequence of lion
samples showed that samples from West Africa shared alleles with samples from
Southern Africa, and samples from Central Africa shared alleles with samples from
Asia. This phenomenon indicates that Central Africa was a melting pot of lion
populations after they had become isolated, possibly migrating through corridors in
the Nile Basin during the early Holocene.[21]
Hybrids
Further information: Panthera hybrid
In zoos, lions have been bred with tigers to create hybrids for the curiosity of visitors or
for scientific purpose.[42][43] The liger is bigger than a lion and a tiger, whereas
most tigons are relatively small compared to their parents because of reciprocal gene
effects.[44][45] The leopon is a hybrid between a lion and leopard.[46]

Description
A tuft at the end of the tail is a distinct characteristic of the lion.
Skeleton

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. [47][48]
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.[49]
Of the living felid species, the lion is rivaled only by the tiger in length, weight, and
height at the shoulder.[50] 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. [51][52]
Size
The size and weight of adult lions varies across global range and habitats. [53][54][55]
[56]
 Accounts of a few individuals that were larger than average exist from Africa and
India.[47][57][58][59]

Average Female lions Male lions

Head-and-body
160–184 cm (63–72 in)[60] 184–208 cm (72–82 in)[60]
length

Tail length 72–89.5 cm (28.3–35.2 in)[60] 82.5–93.5 cm (32.5–36.8 in)[60]

 118.37–143.52 kg (261.0–316.4 lb) in 186.55–225 kg (411.3–496.0 lb) in
Southern Africa,[53] Southern Africa,[53]
Weight
119.5 kg (263 lb) in East Africa,[53] 174.9 kg (386 lb) in East Africa,[53]
110–120 kg (240–260 lb) in India[54] 160–190 kg (350–420 lb) in India[54]

Mane

A six-year-old male with a large mane at Phinda Private Game Reserve, South Africa

Male with short mane at Pendjari National Park, Benin, West Africa

The male lion's mane is the most recognisable feature of the species. [14] It may have
evolved around 320,000–190,000 years ago. [61] 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. Male lions usually aim for the backs or hindquarters of rivals, rather
than their necks.[62][63] Cool ambient temperature in European and North American zoos
may result in a heavier mane.[64] Asiatic lions usually have sparser manes than average
African lions.[65]
Almost all male lions in Pendjari National Park are either maneless or have very short
manes.[66] Maneless lions have also been reported in Senegal, in Sudan's Dinder
National Park and in Tsavo East National Park, Kenya.[67] 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.[68] Increased testosterone may be the
cause of maned lionesses reported in northern Botswana. [69]
Colour variation
Further information: White lion
The white lion is a rare morph with a genetic condition called leucism which is caused
by a double recessive allele. It is not albino; it has normal pigmentation in the eyes and
skin. White lions have occasionally been encountered in and around Kruger National
Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. They
were removed from the wild in the 1970s, thus decreasing the white lion gene pool.
Nevertheless, 17 births have been recorded in five prides between 2007 and 2015.
[70]
 White lions are selected for breeding in captivity.[71] They have reportedly been bred in
camps in South Africa for use as trophies to be killed during canned hunts.[72]

Distribution and habitat

Lion in Gir National Park

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 rainforests and rarely enters closed forests. 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.[47] Lions occur in savannah grasslands with scattered acacia trees,
which serve as shade.[73] The Asiatic lion now survives only in and around Gir National
Park in Gujarat, western India. Its habitat is a mixture of dry savannah forest and very
dry, deciduous scrub forest.[15]
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 southern Europe and Asia, the lion once ranged in regions where climatic conditions
supported an abundance of prey.[78] In Greece, it was common as reported
by Herodotus in 480 BC; it was considered rare by 300 BC and extirpated by AD 100.
[47]
 It was present in the Caucasus until the 10th century.[52] It lived in Palestine until
the Middle Ages, and in Southwest Asia until the late 19th century. By the late 19th
century, it had been extirpated in most of Turkey. [79] The last live lion in Iran was sighted
in 1942 about 65 km (40 mi) northwest of Dezful,[80] although the corpse of a lioness was
found on the banks of the Karun river in Khūzestān Province in 1944.[81] It once ranged
from Sind and Punjab in Pakistan to Bengal and the Narmada River in central India.[82]

Behaviour and ecology

Lions spend much of their time resting; they are inactive for about twenty 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 fifty minutes eating.[84]
Group organisation

Five tree-climbing lions in Uganda

A lioness (left) and two males in Masai Mara

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 changes only 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. Female 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.[57]
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. [91]
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,[48][103][104] while domestic livestock may contribute significantly to their
diet.[104] They usually avoid fully grown
adult elephants, rhinoceroses and hippopotamus and small prey like dik-
dik, hyrax, hare and monkey.[103][105] Unusual prey include porcupines and small reptiles.
Lions kill other predators such as leopard, cheetah and spotted hyena but seldom
consume them.[106]

A skeletal mount of a lion attacking a common eland, on display at The Museum of Osteology

Four lionesses catching a buffalo in the Serengeti

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
zebra and wildebeest, while larger prey like buffalo and giraffe are riskier. [91] In Chobe
National Park, large prides have been observed hunting African bush elephants up to
around 15 years old in exceptional cases, with most of the victims being calves and
subadults.[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
run quickly only 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. They usually pull it down by the rump and kill
by a strangling bite to the throat. They also kill prey by enclosing its mouth and nostrils
in their jaws.[115]
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;[81] 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.[56] Carrion is thought to provide a large part of lion diet.
[120]

Predator competition

Lion attacked by spotted hyenas in Sabi Sand Game Reserve

Lioness stealing a kill from a leopard in Kruger National Park

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 animal can lose their kills to
the other. Lions have been observed killing crocodiles that ventured onto land.
[137]
 Crocodiles may also kill and eat lions, evidenced by the occasional lion claw found in
crocodile stomachs.[138]
Reproduction and life cycle

Lions mating at Masai Mara

A lion cub in Masai Mara

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 the 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—one male could
patiently let the cubs play with his tail or his mane, while another may snarl and bat the
cubs away.[148]

Video of a lioness and her cubs in Phinda Reserve

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, adolescents 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

Lions in a tree near Lake Nakuru

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 tapeworm species of the 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

Head rubbing among pride members is a common social behaviour

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—nuzzling 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

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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]