The
lion (
Panthera leo) is one of the four
big cats in the
genus Panthera and a member of the family
Felidae. With some males exceeding 250 kg (550 lb) in weight,
[4] it is the second-largest living cat after the
tiger. Wild lions currently exist in
sub-Saharan Africa and in
Asia (where an
endangered remnant population resides in
Gir Forest National Park in
India) while other types of lions have disappeared from
North Africa and
Southwest Asia in historic times. Until the late
Pleistocene,
about 10,000 years ago, the lion was the most widespread large land
mammal after humans. They were found in most of Africa, across Eurasia
from western Europe to India, and in the Americas from the
Yukon to
Peru.
[5] The lion is a
vulnerable species,
having seen a major population decline in its African range of 30–50%
per two decades during the second half of the 20th century.
[2]
Lion populations are untenable outside designated reserves and national
parks. Although the cause of the decline is not fully understood,
habitat loss and conflicts with humans are currently the greatest causes
of concern. Within Africa, the
West African lion population is particularly endangered.
Lions live for 10–14 years in the wild, while in captivity they can
live longer than 20 years. In the wild, males seldom live longer than 10
years, as injuries sustained from continual fighting with rival males
greatly reduce their longevity.
[6] They typically inhabit
savanna and
grassland, although they may take to
bush and
forest. Lions are unusually
social
compared to other cats. A pride of lions consists of related females
and offspring and a small number of adult males. Groups of female lions
typically hunt together, preying mostly on large
ungulates. Lions are
apex and
keystone predators,
although they scavenge as opportunity allows. While lions do not
typically hunt humans, some have been known to do so. Sleeping mainly
during the day, lions are primarily
nocturnal, although bordering on
crepuscular[7][8] in nature.
Highly distinctive, the male lion is easily recognised by its mane,
and its face is one of the most widely recognised animal symbols in
human
culture. Depictions have existed from the
Upper Paleolithic period, with carvings and paintings from the
Lascaux and
Chauvet Caves,
through virtually all ancient and medieval cultures where they once
occurred. It has been extensively depicted in sculptures, in 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
zoos over the world since the late eighteenth century. Zoos are cooperating worldwide in breeding programs for the endangered
Asiatic subspecies.
Etymology
The lion's name, similar in many
Romance languages, is derived from the
Latin leo;
[9] and the
Ancient Greek λέων (
leon).
[10] The
Hebrew word
לָבִיא (
lavi) may also be related.
[11] It was one of the many species originally described by
Linnaeus, who gave it the name
Felis leo, in his eighteenth-century work,
Systema Naturae.
[3]
Taxonomy and evolution
The lion's closest relatives are the other species of the genus
Panthera: the
tiger, the
jaguar, and the
leopard.
P. leo evolved in
Africa between 1 million and 800,000 years ago, before spreading throughout the
Holarctic region.
[12] It appeared in the fossil record in Europe for the first time 700,000 years ago with the subspecies
Panthera leo fossilis at
Isernia in
Italy. From this lion derived the later cave lion (
Panthera leo spelaea), which appeared about 300,000 years ago.
[13] Lions died out in northern
Eurasia at the end of the last
glaciation, about 10,000 years ago;
[14] this may have been secondary to the
extinction of
Pleistocene megafauna.
[15]
Subspecies
Traditionally, 12 recent
subspecies
of lion were recognised, distinguished by mane appearance, size, and
distribution. Because these characteristics are very insignificant and
show a high individual variability, most of these forms were probably
not true subspecies, especially as they were often based upon zoo
material of unknown origin that may have had "striking, but abnormal"
morphological characteristics.
[16] Today, only eight subspecies are usually accepted,
[14][17] although one of these, the
Cape lion, formerly described as
Panthera leo melanochaita, is probably invalid.
[17] Even the remaining seven subspecies might be too many. While the status of the Asiatic lion (
P. l. persica) as a subspecies is generally accepted, the systematic relationships among African lions are still not completely resolved.
Mitochondrial
variation in living African lions seemed to be modest according to some
newer studies, therefore all sub-Saharan lions sometimes have been
considered a single subspecies, however, a recent study revealed lions
from western and central Africa differ genetically from lions of
southern or eastern Africa. According to this study,
Western African lions
are more closely related to Asian lions than to South or East African
lions. These findings might be explained by a late Pleistocene
extinction event of lions in western and central Africa and a subsequent
recolonization of these parts from Asia.
[18]
Previous studies, which were focused mainly on lions from eastern and
southern parts of Africa, already showed these can be possibly divided
in two main
clades: one to the west of the
Great Rift Valley and the other to the east. Lions from
Tsavo in eastern Kenya are much closer genetically to lions in
Transvaal (South Africa), than to those in the
Aberdare Range in western Kenya.
[19]
Another study revealed there are three major types of lions, one North
African–Asian, one southern African and one middle African.
[20] Conversely, Per Christiansen found that using skull morphology allowed him to identify the subspecies
krugeri,
nubica,
persica, and
senegalensis, while there was overlap between
bleyenberghi with
senegalensis and
krugeri. The Asiatic lion
persica was the most distinctive, and the Cape lion had characteristics allying it more with
P. l. persica than the other sub-Saharan lions. He had analysed 58 lion skulls in three European museums.
[21]
By far most lions kept in zoos are hybrids of different subspecies. Approximately 77% of the captive lions registered by the
International Species Information System
are of unknown origin. Nonetheless, they might carry genes which are
extinct in the wild, and might be therefore important to maintain
overall genetic variability of the lion.
[17]
It is believed that those lions, imported to Europe before the middle
of the nineteenth century, were mainly either Barbary lions from North
Africa or lions from the Cape.
[22]
Recent
Range map of the commonly accepted subspecies of the lion in the late twentieth century
Eight recent (
Holocene) subspecies are recognised today:
- P. l. persica, known as the Asiatic lion or South Asian, Persian, or Indian lion, once was widespread from Turkey, across Southwest Asia, to India and Pakistan, however, large prides and daylight activity made them easier to poach than tigers or leopards;[23] now around 400 exist in and near the Gir Forest of India.[24]
Genetic evidence suggests its ancestors split from the ancestors of
sub-Saharan African lions between 203 and 74 thousand years ago.[14]
- P. l. leo, known as the Barbary lion, originally ranged from Morocco to Egypt. It is extinct in the wild due to excessive hunting, as the last wild Barbary lion was killed in Morocco in 1922.[25] This was one of the largest of the lion subspecies,[26]
with reported lengths of 3.0–3.3 m (9.8–10.8 ft) and weights of more
than 200 kg (440 lb) for males. It appears to be more closely related to
the Asiatic rather than sub-Saharan lions. A number of animals in
captivity are likely to be Barbary lions,[27] particularly the 90 animals descended from the Moroccan Royal collection at Rabat Zoo.[28]
- P. l. senegalensis, known as the West African lion, is found in western Africa, from Senegal to the Central African Republic.[29][30]
- P. l. azandica, known as the northeast Congo Lion, is found in the northeastern parts of the Congo.[29]
- P. l. nubica, known as the East African or Masai lion is found in East Africa, from Ethiopia and Kenya to Tanzania and Mozambique;[30] a local population is known as the Tsavo lion.
- P. l. bleyenberghi, known as the southwest African or Katanga lion, is found in southwestern Africa, Namibia, Botswana, Angola, Katanga (Democratic Republic of the Congo), Zambia, and Zimbabwe.[30]
- P. l. krugeri, known as the southeast African or Transvaal lion, is found in the Transvaal region of southeastern Africa, including Kruger National Park.[30]
- P. l. melanochaita, known as the Cape lion, became extinct in the wild around 1860. Results of mitochondrial DNA
research do not support its status as a distinct subspecies. The Cape
lion probably was only the southernmost population of the extant P. l. krugeri.[17]
- A newly discerned lion subspecies could exist in captivity in Ethiopia's capital city of Addis Ababa.[31] Researchers compared the microsatellite variations over ten loci
of fifteen lions in captivity with those of six different wild lion
populations. They determined that these lions are genetically unique and
presumably that "their wild source population is similarly unique."
These lions—with males that have a distinctly dark and luxuriant mane
seam to define a new subspecies perhaps native only to Ethiopia. These lions were part of a collection of the late Haile Selassie I of Ethiopia.[32]
Pleistocene
Several additional subspecies of lion existed in prehistoric times:
- P. l. fossilis, known as the Middle Pleistocene European cave lion, flourished about 500,000 years ago; fossils have been recovered from Germany and Italy.
It was larger than today's African lions, reaching sizes comparable to
the American cave lion and slightly larger than the Upper Pleistocene
European cave lion.[14][33]
- P. l. spelaea,
known as the European cave lion, Eurasian cave lion, or Upper
Pleistocene European cave lion, occurred in Eurasia 300,000 to 10,000
years ago.[14] This species is known from Paleolithic cave paintings, ivory carvings, and clay busts,[34]
indicating it had protruding ears, tufted tails, perhaps faint
tiger-like stripes, and at least some had a ruff or primitive mane
around their necks, possibly indicating males.[35]
- P. l. atrox, known as the American lion or American cave lion, was abundant in the Americas from Canada to Peru in the Pleistocene Epoch until about 10,000 years ago. This form is the sister clade of P. l. spelaea, and likely arose when an early P. l. spelaea population became isolated south of the North American continental ice sheet about 0.34 Mya.[36] One of the largest purported lion subspecies to have existed, its body length is estimated to have been 1.6–2.5 m (5–8 ft).[37]
Dubious
- P. l. youngi or Panthera youngi, flourished 350,000 years ago.[5] Its relationship to the extant lion subspecies is obscure, and it probably represents a distinct species.
- P. l. sinhaleyus, known as the Sri Lanka lion, appears to have become extinct around 39,000 years ago. It is only known from two teeth found in deposits at Kuruwita. Based on these teeth, P. Deraniyagala erected this subspecies in 1939.[38]
- P. l. vereshchagini, the Beringian cave lion of Yakutia (Russia), Alaska (USA), and the Yukon Territory (Canada), has been considered a subspecies separate from P. l. spelaea on morphological grounds. However, mitochondrial DNA sequences obtained from cave lion fossils from Europe and Alaska were indistinguishable.[36]
- P. l. mesopotamica or Mesopotamian lion, flourished during the Neo-Assyrian Period (approximately 1000–600 BC).[39]
It inhabited the Mesopotamian Plain where it probably represents a
distinct sub-species. Nearly all ancient Mesopotamian representations of
male lions demonstrate full underbelly hair in which until recently was
only identified in the Barbary lion (Panthera leo leo) from
Northern Africa. Ancient evidence from adjacent landmasses reveal no
substantiation for lions with underbelly hair in this manner so that the
distinct phenotype of depicted lions in ancient Mesopotamia (including
Babylon, Elam and ancient Persia) represent an extinct sub-species. Many
of the images of these lions are derived from lion hunting sculptures
so that the extinction of this sub-species likely resulted from
overhunting in the ancient world.
- P. l. europaea, known as the European lion, was probably identical with Panthera leo persica or Panthera leo spelea. It became extinct around 100 AD due to persecution and over-exploitation. It inhabited the Balkans, the Italian Peninsula, southern France, and the Iberian Peninsula. It was a very popular object of hunting among ancient Romans and Greeks.
- P. l. maculatus, known as the marozi
or spotted lion, sometimes is believed to be a distinct subspecies, but
may be an adult lion that has retained its juvenile spotted pattern. If
it was a subspecies in its own right, rather than a small number of
aberrantly coloured individuals, it has been extinct since 1931. A less
likely identity is a natural leopard-lion hybrid commonly known as a leopon.[40]
Hybrids
Video of lioness and her cubs in the wild, South Africa
Lions have been known to
breed with
tigers (most often the
Siberian and
Bengal subspecies) to create hybrids called
ligers and
tiglons (or tigons).
[41] They also have been crossed with
leopards to produce
leopons,
[42] and
jaguars to produce
jaglions. The
marozi is reputedly a spotted lion or a naturally occurring leopon, while the
Congolese spotted lion
is a complex lion-jaguar-leopard hybrid called a lijagulep. Such
hybrids were once commonly bred in zoos, but this is now discouraged due
to the emphasis on conserving species and subspecies. Hybrids are still
bred in private menageries and in zoos in
China.
The
liger is a cross between a male lion and a tigress.
[43]
Because the growth-inhibiting gene from the female tiger mother is
absent, the growth-promoting gene passed on by the male lion father is
unimpeded by a regulating gene and the resulting ligers grow far larger
than either parent. They share physical and behavioural qualities of
both parent species (spots and stripes on a sandy background). Male
ligers are sterile, but female ligers often are fertile. Males have
about a 50% chance of having a mane, but if they grow them, their manes
will be modest: around 50% the size of a pure lion mane.
Ligers are much bigger than normal lions, typically 3.0 to 3.7 m (10
to 12 feet) in length, and can weigh 360 to 450 kg (800 to 1,000
pounds).
[43]
The less common
tiglon or
tigon is a cross between a lioness and a male tiger.
[44] In contrast to ligers, tigons are often relatively small in comparison to their parents, because of reciprocal gene effects.
[45]
Characteristics
The lion is the tallest (at the shoulder) of all living cats, averaging approximately 5 cm (2.0 in) taller than the
tiger.
Behind only the tiger, the lion is the second largest living felid in
length and weight. Its skull is very similar to that of the tiger,
although the frontal region is usually more depressed and flattened,
with a slightly shorter postorbital region. The lion's skull has broader
nasal openings than the tiger, however, 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.
[46]
Lion coloration varies from light buff to yellowish, reddish, or dark
ochraceous brown. The underparts are generally lighter and the tail tuft
is black. Lion cubs are born with brown
rosettes
(spots) on their body, rather like those of a leopard. Although these
fade as lions reach adulthood, faint spots often may still be seen on
the legs and underparts, particularly on lionesses.
Lions are the only members of the cat family to display obvious
sexual dimorphism
– that is, males and females look distinctly different. They also have
specialised roles that each gender plays in the pride. For instance, the
lioness, the hunter, lacks the male's thick mane. The colour of the
male's mane varies from blond to black, generally becoming darker as the
lion grows older.
During confrontations with others, the mane makes the lion look larger
Weights for adult lions range between 150–250 kg (330–550 lb) for males and 120–182 kg (264–400 lb) for females.
[4] Nowell and Jackson report average weights of 181 kg (399 lb) for males and 126 kg (278 lb) for females.
[25]
Lions tend to vary in size depending on their environment and area,
resulting in a wide spread in recorded weights. For instance, lions in
southern Africa tend to be about 5% heavier than those in
East Africa, in general.
[47]
Head and body length is 170–250 cm (5 ft 7 in – 8 ft 2 in) in males
and 140–175 cm (4 ft 7 in – 5 ft 9 in) in females; shoulder height is up
to 123 cm (4 ft) in males and as low as 91 cm (3 ft) in females.
[48] The tail length is 90–105 cm (2 ft 11 in – 3 ft 5 in) in males and 70–100 cm in females (2 ft 4 in – 3 ft 3 in).
[4] The longest known lion, at nearly 3.6 m (12 ft) in total length, was a black-maned male shot near Mucsso, southern
Angola in October 1973; the heaviest lion known in the wild was a man-eater shot in 1936 just outside
Hectorspruit in eastern
Transvaal,
South Africa and weighed 313 kg (690 lb).
[49] Another notably outsized male lion, which was shot near
Mount Kenya, weighed in at 272 kg (600 lb).
[25]
Lions in captivity tend to be larger than lions in the wild – the
heaviest lion on record is a male at Colchester Zoo in England named
Simba in 1970, which weighed 375 kg (826 lb).
[50]
However, the frequently cited maximum head and body length of 250 cm
(8 ft 2 in) fits rather to extinct Pleistocene forms, like the
American lion, with even large modern lions measuring several centimeters less in length.
[51]
The most distinctive characteristic shared by both females and males
is that the tail ends in a hairy tuft. In some lions, the tuft conceals a
hard "spine" or "spur", approximately 5 mm long, formed of the final
sections of tail bone fused together. The lion is the only felid to have
a tufted tail – the function of the tuft and spine are unknown. Absent
at birth, the tuft develops around 5½ months of age and is readily
identifiable at 7 months.
[52]
Mane
The mane of the adult male lion, unique among cats, is one of the
most distinctive characteristics of the species. It makes the lion
appear larger, providing an excellent intimidation display; this aids
the lion during confrontations with other lions and with the species'
chief competitor in Africa, the
spotted hyena.
[53] The presence, absence, colour, and size of the mane is associated with genetic precondition, sexual maturity, climate, and
testosterone
production; the rule of thumb is the darker and fuller the mane, the
healthier the lion. Sexual selection of mates by lionesses favors males
with the densest, darkest mane.
[54] Research in
Tanzania
also suggests mane length 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.
[55]
In prides including a coalition of two or three males, it is possible
that lionesses solicit mating more actively with the males who are more
heavily maned.
[54]
A
maneless male lion, who is devoid of much body hair as well, Tsavo East National Park, Kenya
Scientists once believed that the distinct status of some subspecies could be justified by
morphology, including the size of the mane. Morphology was used to identify subspecies such as the
Barbary lion and
Cape lion. Research has suggested, however, that environmental factors influence the colour and size of a lion's mane, such as the
ambient temperature.
[55] The cooler ambient temperature in European and North American
zoos, for example, may result in a heavier mane. Thus the mane is not an appropriate marker for identifying subspecies.
[17][56] The males of the Asiatic subspecies, however, are characterised by sparser manes than average African lions.
[57]
In the
Pendjari National Park area almost all males are maneless or have very weak manes.
[58] Maneless male lions have also been reported from
Senegal and from
Tsavo East National Park
in Kenya, and the original male white lion from Timbavati also was
maneless. The testosterone hormone has been linked to mane growth,
therefore
castrated lions often have minimal to no mane, as the removal of the gonads inhibits testosterone production.
[59]
Cave paintings of extinct European
cave lions almost exclusively show animals with no manes, suggesting that either they were maneless,
[35] or that the paintings depict lionesses as seen hunting in a group.
White lions
White lions owe their colouring to a recessive gene; they are rare forms of the subspecies
Panthera leo krugeri
The
white lion is not a distinct subspecies, but a special
morph with a genetic condition,
leucism,
[16] that causes paler colouration akin to that of the
white tiger; the condition is similar to
melanism, which causes
black panthers. They are not albinos, having normal pigmentation in the eyes and skin. White Transvaal lion (
Panthera leo krugeri) individuals occasionally have been encountered in and around
Kruger National Park and the adjacent
Timbavati Private Game Reserve in eastern South Africa, but are more commonly found in
captivity, where breeders deliberately select them. The unusual cream colour of their coats is due to a
recessive gene.
[60] Reportedly, they have been bred in camps in South Africa for use as trophies to be killed during
canned hunts.
[61]
Behaviour
Lions spend much of their time resting and are inactive for about 20 hours per day.
[62]
Although lions can be active at any time, their activity generally
peaks after dusk with a period of socializing, grooming, and defecating.
Intermittent bursts of activity follow through the night hours until
dawn, when hunting most often takes place. They spend an average of two
hours a day walking and 50 minutes eating.
[63]
Group organization
Two lionesses and a mature male of a pride, northern Serengeti
Two males and a female in Masai Mara, southwest Kenya
Lions are the most socially inclined of all wild
felids, most of which remain quite solitary in nature. The lion is a predatory
carnivore with two types of social organization. Some lions are
residents, living in groups centering around related lionesses, called
prides.
[64] The pride usually consists of five or six females, their cubs of both sexes, and one or two males (known as a
coalition
if more than one) who mate with the adult females (although extremely
large prides, consisting of up to 30 individuals, have been observed).
The number of adult males in a coalition is usually two, but may
increase to four and decrease again over time. Male cubs are excluded
from their maternal pride when they reach maturity.
The second organizational behaviour is labeled
nomads, who range widely and move about sporadically, either singularly or in pairs.
[64]
Pairs are more frequent among related males who have been excluded from
their birth pride. Note that a lion may switch lifestyles; nomads may
become residents and vice versa. Males have to go through this lifestyle
and some never are able to join another pride. A female who becomes a
nomad has much greater difficulty joining a new pride, as the females in
a pride are related, and they reject most attempts by an unrelated
female to join their family group.
The area a pride occupies is called a
pride area, whereas that by a nomad is a
range.
[64] The males associated with a pride tend to stay on the fringes, patrolling their
territory. Why
sociality
– the most pronounced in any cat species – has developed in lionesses
is the subject of much debate. Increased hunting success appears an
obvious reason, but this is less than sure upon examination: coordinated
hunting does allow for more successful predation, but also ensures that
non-hunting members reduce per capita caloric intake, however, some
take a role raising cubs, who may be left alone for extended periods of
time. Members of the pride regularly tend to 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 and they are the first to consume the prey
at the site it is taken. Other benefits include possible
kin selection
(better to share food with a related lion than with a stranger),
protection of the young, maintenance of territory, and individual
insurance against injury and hunger.
[25]
Lionesses do most of the hunting for their pride. They are more
effective hunters as they are smaller, swifter and more agile than the
males, and unencumbered by the heavy and conspicuous mane, which causes
overheating during exertion. They act as a coordinated group with
members who perform the same role consistently in order to stalk and
bring down the prey successfully. Smaller prey is eaten at the location
of the hunt, thereby being shared among the hunters; when the kill is
larger it often is dragged to the pride area. There is more sharing of
larger kills,
[65]
although pride members often behave aggressively toward each other as
each tries to consume as much food as possible. If near the conclusion
of the hunt, males have a tendency to dominate the kill once the
lionesses have succeeded. They are more likely to share this with the
cubs than with the lionesses, but males rarely share food they have
killed by themselves.
Both males and females can defend the pride against intruders, but
the male lion is better-suited for this purpose due to its stockier,
more powerful build.
[66] Some individuals consistently lead the defence against intruders, while others lag behind.
[67] Lions tend to assume specific roles in the pride. Those lagging behind may provide other valuable services to the group.
[68]
An alternative hypothesis is that there is some reward associated with
being a leader who fends off intruders and the rank of lionesses in the
pride is reflected in these responses.
[69]
The male or males associated with the pride must defend their
relationship to the pride from outside males who attempt to take over
their relationship with the pride. Females form the stable
social unit in a pride and do not tolerate outside females;
[70] membership only changes with the births and deaths of lionesses,
[71] although some females do leave and become nomadic.
[72] Subadult males on the other hand, must leave the pride when they reach maturity at around 2–3 years of age.
[72]
Hunting and diet
While a lioness such as this one has very sharp teeth, prey is usually killed by strangulation
The lioness is the one who does the hunting for the pride. The male
lion associated with the pride usually stays and watches its young while
waiting for the lionesses to return from the hunt. Typically, several
lionesses work together and encircle the herd from different points.
Once they have closed with a herd, they usually target the closest prey.
The attack is short and powerful; they attempt to catch the victim with
a fast rush and final leap. The prey usually is killed by
strangulation,
[73] which can cause
cerebral ischemia or
asphyxia (which results in
hypoxemic, or "general",
hypoxia).
The prey also may be killed by the lion enclosing the animal's mouth
and nostrils in its jaws (which would also result in asphyxia). Smaller
prey, though, may simply be killed by a swipe of a lion's paw.
[4]
Lions are powerful animals that usually hunt in coordinated groups
and stalk their chosen prey. However, they are not particularly known
for their stamina – for instance, a lioness' heart makes up only 0.57%
of her body weight (a male's is about 0.45% of his body weight), whereas
a hyena's heart is close to 1% of its body weight.
[74] Thus, they only run fast in short bursts,
[75]
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.
[76] They sneak up to the victim until they reach a distance of approximately 30 metres (98 ft) or less.
Lioness in a burst of speed while hunting in the Serengeti
The
prey consists mainly of large mammals, with a preference for
wildebeest,
zebras,
buffalo, and
warthogs in Africa and
nilgai,
wild boar, and several
deer species in India. Many other species are hunted, based on availability. Mainly this will include
ungulates weighing between 50 and 300 kg (110–660 lb) such as
kudu,
hartebeest,
gemsbok, and
eland.
[4] Occasionally, they take relatively small species such as
Thomson's gazelle or
springbok.
Lions hunting in groups are capable of taking down most animals, even
healthy adults, but in most parts of their range they rarely attack very
large prey such as fully grown male
giraffes due to the danger of injury.
Extensive statistics collected over various studies show that lions
normally feed on mammals in the range 190–550 kg (420–1210 lb). In
Africa, wildebeest rank at the top of preferred prey (making nearly half
of the lion prey in the
Serengeti) followed by zebra.
[77] Most adult
hippopotamuses,
rhinoceroses,
elephants, and smaller
gazelles,
impala,
and other agile antelopes are generally excluded. However giraffes and
buffalos are often taken in certain regions. For instance, in Kruger
National Park, giraffes are regularly hunted.
[78] In Manyara Park, Cape buffaloes constitute as much as 62% of the lion's diet,
[79]
due to the high number density of buffaloes. Occasionally hippopotamus
is also taken, but adult rhinoceroses are generally avoided. Even though
smaller than 190 ;kg (420 lb), warthogs are often taken depending on
availability.
[80] In some areas, lions specialise in hunting atypical prey species; this is the case at the
Savuti river, where they prey on elephants.
[81]
Park guides in the area reported that the lions, driven by extreme
hunger, started taking down baby elephants, and then moved on to
adolescents and, occasionally, fully grown adults during the night when
elephants' vision is poor.
[82] Lions also attack domestic livestock; in India cattle contribute significantly to their diet.
[57] Lions are capable of killing other predators such as
leopards,
cheetahs,
hyenas, and
wild dogs,
though (unlike most felids) they seldom devour the competitors after
killing them. They also scavenge animals either dead from
natural causes
(disease) or killed by other predators, and keep a constant lookout for
circling vultures, being keenly aware that they indicate an animal dead
or in distress.
[83] A lion may gorge itself and eat up to 30 kg (66 lb) in one sitting;
[84]
if it is unable to consume all the kill it will rest for a few hours
before consuming more. On a hot day, the pride may retreat to shade
leaving a male or two to stand guard.
[85] An adult lioness requires an average of about 5 kg (11 lb) of meat per day, a male about 7 kg (15.5 lb).
[86]
Four lionesses take down a cape buffalo in the central Serengeti, Tanzania
Because lionesses hunt in open spaces where they are easily seen by
their prey, cooperative hunting increases the likelihood of a successful
hunt; this is especially true with larger species. Teamwork also
enables them to defend their kills more easily against other large
predators such as hyenas, which may be attracted by
vultures
from kilometres away in open savannas. Lionesses do most of the
hunting; males attached to prides do not usually participate in hunting,
except in the case of larger quarry such as giraffe and buffalo. In
typical hunts, each lioness has a favored 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 in flight from other
lionesses.
[87]
Young lions first display stalking behaviour around three months of
age, although they do not participate in hunting until they are almost a
year old. They begin to hunt effectively when nearing the age of two.
[88]
Predator competition
Lions and
spotted hyenas
occupy the same ecological niche (and hence compete) where they
coexist. A review of data across several studies indicates a dietary
overlap of 58.6%.
[89]
Lions typically ignore spotted hyenas, unless they are on a kill or are
being harassed by them, while the latter tend to visibly react to the
presence of lions, whether there is food or not. 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. Lions are quick to
follow the calls of hyenas feeding, a fact that was proven by Dr. Hans
Kruuk, who found that lions repeatedly approached him whenever he played
the tape-recorded calls of hyenas feeding.
[90]
When confronted on a kill by lions, spotted hyenas will either leave or
wait patiently at a distance of 30–100 m (100–350 ft) until the lions
have finished.
[91]
In some cases, spotted hyenas are bold enough to feed alongside lions,
and may occasionally force the lions off a kill. The two species may act
aggressively toward one another even when there is no food involved.
Lions may charge at hyenas and maul them for no apparent reason: one
male lion was filmed killing two matriarch hyenas on separate occasions
without eating them,
[92] and lion predation can account for up to 71% of hyena deaths in
Etosha. Spotted hyenas have adapted by frequently mobbing lions that enter their territories.
[93]
Experiments on captive spotted hyenas revealed that specimens with no
prior experience with lions act indifferently to the sight of them, but
will react fearfully to the scent.
[90]
The size of male lions allows them occasionally to confront hyenas in
otherwise evenly matched brawls and so to tip the balance in favour of
the lions.
Lions tend to dominate smaller felines such as
cheetahs and
leopards
where they co-occur, stealing their kills and killing their cubs and
even adults when given the chance. The cheetah has a 50% chance of
losing its kill to lions or other predators.
[94]
Lions are major killers of cheetah cubs, up to 90% of which are lost in
their first weeks of life due to attacks by other predators. Cheetahs
avoid competition by hunting at different times of the day and hide
their cubs in thick brush. Leopards also use such tactics, but have the
advantage of being able to subsist much better on small prey than either
lions or cheetahs. Also, unlike cheetahs, leopards can climb trees and
use them to keep their cubs and kills away from lions; however,
lionesses will occasionally be successful in climbing to retrieve
leopard kills.
[95] Similarly, lions dominate
African wild dogs,
not only taking their kills but also preying on young and (rarely)
adult dogs. Population densities of wild dogs are low in areas where
lions are more abundant.
[96]
However, in Kruger National Park, there have been records of wild dogs
killing lions and there is one report of eight dogs killing and eating
an adult male.
[97]
The
Nile crocodile is the only
sympatric
predator (besides humans) that can singly threaten the lion. Depending
on the size of the crocodile and the lion, either can lose kills or
carrion to the other. Lions have been known to kill crocodiles venturing
onto land,
[98] while the reverse is true for lions entering waterways, as evidenced by the occasional lion claw found in crocodile stomachs.
[99]
Reproduction and life cycle
Most lionesses will have reproduced by the time they are four years of age.
[100] Lions do not
mate at any specific time of year, and the females are
polyestrous.
[101] As with other
cats' penises, 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.
[102] A lioness may mate with more than one male when she is
in heat;
[103]
during a mating bout, which could last several days, the couple
copulates twenty to forty times a day, often forgoing eating. Lions
reproduce very well in captivity.
The average gestation period is around 110 days,
[101]
the female giving birth to a litter of one to 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 rest of the pride. She will often hunt by
herself while the cubs are still helpless, staying relatively close to
the thicket or den where the cubs are kept.
[104]
The cubs themselves are born blind – their eyes do not open until
roughly a week 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.
[105]
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, to prevent scent from
building up at a single den site and thus avoiding the attention of
predators that may harm the cubs.
[104]
Usually, the mother does not integrate herself and her cubs back into the pride until the cubs are six to eight weeks old.
[104]
Sometimes this introduction to pride life occurs earlier, however,
particularly if other lionesses have given birth at about the same time.
For instance, lionesses in a pride often synchronise their reproductive
cycles so that they cooperate in the raising and suckling of the young
(once the cubs are past the initial stage of isolation with their
mother), who suckle indiscriminately from any or all of the nursing
females in the pride. In addition to greater protection, the
synchronization of births also has an advantage in that the cubs end up
being roughly the same size, and thus have an equal chance of survival.
If one lioness gives birth to a litter of cubs a couple of months after
another lioness, for instance, then the younger cubs, being much smaller
than their older brethren, usually are dominated by larger cubs at
mealtimes – consequently, death by starvation is more common amongst the
younger cubs.
A pregnant lioness (right)
In addition to starvation, cubs also face many other dangers, such as
predation by jackals, hyenas, leopards, martial eagles, and snakes.
Even buffaloes, should they catch the scent of lion cubs, often stampede
toward the thicket or den where they are being kept, doing their best
to trample the cubs to death while warding off the lioness. Furthermore,
when one or more new males oust the previous male(s) associated with a
pride, the conqueror(s) often kill any existing young cubs,
[106]
perhaps because females do not become fertile and receptive until their
cubs mature or die. All in all, as many as 80% of the cubs will die
before the age of two.
[107]
When first introduced to the rest of the pride, the cubs initially
lack confidence when confronted with adult lions other than their
mother. They soon begin to immerse themselves in the pride life,
however, playing amongst themselves or attempting to initiate play with
the adults. Lionesses with cubs of their own are more likely to be
tolerant of another lioness's cubs than lionesses without cubs. The
tolerance of the male lions toward 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.
[108]
Male lions are generally more likely to share food with cubs than with
lionesses, unless they have caught it, they rarely share their own
catches with others
[clarification needed]
Weaning occurs after six to seven months. Male lions reach maturity
at about 3 years of age and, at 4–5 years of age, are capable of
challenging and displacing the adult male(s) associated with another
pride. They begin to age and weaken between 10 and 15 years of age at
the latest,
[109]
if they have not already been critically injured while defending the
pride (once ousted from a pride by rival males, male lions rarely manage
a second take-over). This leaves a short window for their own offspring
to be born and mature. If they are able to procreate as soon as they
take over a pride, potentially, they may have more offspring reaching
maturity before they also are displaced. A lioness often will attempt to
defend her cubs fiercely from a usurping male, but such actions are
rarely successful. He usually kills all of the existing cubs who are
less than two years old. A lioness is weaker and much lighter than a
male; success is more likely when a group of three or four mothers
within a pride join forces against one male.
[106]
Contrary to popular belief, it is not only males that are ousted from
their pride to become nomads, although most females certainly do remain
with their birth pride. However, when the pride becomes too large, the
next generation of female cubs may be forced to leave to eke out their
own territory. Furthermore, when a new male lion takes over the pride,
subadult lions, both male and female, may be evicted.
[110]
Life is harsh for a female nomad. Nomadic lionesses rarely manage to
raise their cubs to maturity, without the protection of other pride
members. One scientific study reports that both males and females may
interact
homosexually[clarification needed].
[111][112]
Health
Although adult lions have no natural predators, evidence suggests that the majority die violently from humans or other lions.
[113]
Lions often inflict serious injuries on each other, either members of
different prides encountering each other in territorial disputes, or
members of the same pride fighting at a kill.
[114]
Crippled lions and lion cubs may fall victim to hyenas, leopards, or be
trampled by buffalo or elephants, and careless lions may be maimed when
hunting prey.
[115]
Lions seeking refuge from flies by climbing a tree, Lake Nakuru, Kenya
Various species of
tick commonly infest the ears, neck and groin regions of most lions.
[116][117] Adult forms of several species of the tapeworm genus
Taenia have been isolated from intestines, the lions having ingested larval forms from antelope meat.
[118] Lions in the
Ngorongoro Crater were afflicted by an outbreak of stable fly (
Stomoxys calcitrans)
in 1962; this resulted in lions becoming covered in bloody bare patches
and emaciated. Lions sought unsuccessfully to evade the biting flies by
climbing trees or crawling into hyena burrows; many perished or
emigrated as the population dropped from 70 to 15 individuals.
[119] A more recent outbreak in 2001 killed six lions.
[120] Lions, especially in captivity, are vulnerable to the
canine distemper virus (CDV),
feline immunodeficiency virus (FIV), and
feline infectious peritonitis (FIP).
[16] CDV is spread through 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.
[121] FIV, which is similar to
HIV
while not known to adversely affect lions, is worrisome enough in its
effect in domestic cats that the Species Survival Plan recommends
systematic testing in captive lions. It occurs with high to endemic
frequency in several wild lion populations, but is mostly absent from
Asiatic and Namibian lions.
[16]
Communication
Head rubbing and licking are common social behaviours within a pride
When resting, lion socialization occurs through a number of
behaviours, and the animal's expressive movements are highly developed.
The most common peaceful tactile gestures are head rubbing and
social licking,
[122] which have been compared with grooming in primates.
[123] Head rubbing – nuzzling one's forehead, face and neck against another lion – appears to be a form of greeting,
[124]
as it 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.
[125]
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, which may have
arisen out of utility, as a lion cannot lick these areas individually.
[126]
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A lion in captivity roaring
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Lions have an array of facial expressions and body postures that serve as visual gestures.
[127]
Their repertoire of vocalizations is also large; variations in
intensity and pitch, rather than discrete signals, appear central to
communication. Lion sounds include snarling, hissing, coughing,
miaowing, woofing, and roaring. Lions tend to
roar in a very characteristic manner, starting with a few deep, long roars that trail off into a series of shorter ones.
[128] [129]
They most often roar at night; the sound, which can be heard from a
distance of 8 kilometres (5.0 mi), is used to advertise the animal's
presence.
[130] Lions have the loudest roar of any big cat.
Distribution and habitat
In Africa, lions can be found in savanna grasslands with scattered
Acacia trees which serve as shade;
[132] their habitat in India is a mixture of dry savanna forest and very dry deciduous scrub forest.
[133] The habitat of lions originally spanned the southern parts of
Eurasia, ranging from
Greece to
India, and most of
Africa except the central
rainforest-zone and the
Sahara desert.
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. By 100 AD they were extirpated.
[134] A population of
Asiatic lions survived until the tenth century in the
Caucasus, their last
European outpost.
[135]
The species was eradicated from
Palestine by the
Middle Ages
and from most of the rest of Asia after the arrival of readily
available firearms in the eighteenth century. Between the late
nineteenth and early twentieth century, they became extinct in
North Africa and
Southwest Asia. By the late nineteenth century, the lion had disappeared from
Turkey and most of northern India,
[16][136] while the last sighting of a live Asiatic lion in
Iran was in 1941 (between
Shiraz and Jahrom,
Fars Province), although the corpse of a lioness was found on the banks of the
Karun river,
Khūzestān Province in 1944. There are no subsequent reliable reports from
Iran.
[84] The subspecies now survives only in and around the
Gir Forest of northwestern India.
[23] Approximately 400 lions live in the area of the 1,412 km
2 (545 sq mi) sanctuary in the state of
Gujarat,
which covers most of the forest. Their numbers have increased from 180
in 1974 mainly because the natural prey species have recovered.
[24][137]
Population and conservation status
Main article:
Lion hunting
The Asiatic lion, whose habitat once ranged from the Mediterranean to
north-west Indian subcontinent, is today found only in the
Gir Forest of
Gujarat, India. It is estimated that only 320 Asiatic lions survive in the wild.
[138]
Most lions now live in eastern and southern Africa, and their numbers
there are rapidly decreasing, with an estimated 30–50% decline per 20
years in the late half of the 20th century.
[2] Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004,
[139][140]
down from early 1990s estimates that ranged as high as 100,000 and
perhaps 400,000 in 1950. Primary causes of the decline include disease
and human interference.
[2] Habitat loss and conflicts with humans are considered the most significant threats to the species.
[141][142] The remaining populations are often geographically isolated from one another, which can lead to
inbreeding, and consequently, reduced
genetic diversity. Therefore the lion is considered a
vulnerable species by the
International Union for Conservation of Nature, while the Asiatic subspecies is
endangered.
[143]
The lion population in the region of West Africa is isolated from lion
populations of Central Africa, with little or no exchange of breeding
individuals. The number of mature individuals in West Africa is
estimated by two separate recent surveys at 850–1,160 (2002/2004). There
is disagreement over the size of the largest individual population in
West Africa: the estimates range from 100 to 400 lions in
Burkina Faso's
Arly-Singou ecosystem.
[2] Another population in northwestern Africa is found in
Waza National Park, where approximately 14–21 animals persist.
[144]
Black maned male lion, shot in the Sotik Plains, Kenya (May 1909)
Conservation of both African and Asian lions has required the setup
and maintenance of national parks and game reserves; among the best
known are
Etosha National Park in
Namibia,
Serengeti National Park in Tanzania, and
Kruger National Park in eastern
South Africa. The
Ewaso Lions Project protects lions in the
Samburu National Reserve,
Buffalo Springs National Reserve and
Shaba National Reserve of the Ewaso Nyiro ecosystem in Northern
Kenya.
[145]
Outside these areas, the issues arising from lions' interaction with
livestock and people usually results in the elimination of the former.
[146] In India, the last refuge of the Asiatic lion is the 1,412 km
2 (545 sq mi)
Gir Forest National Park in western
India which had about 180 lions in 1974 and about 400 in 2010.
[24]
As in Africa, numerous human habitations are close by with the
resultant problems between lions, livestock, locals and wildlife
officials.
[147] The
Asiatic Lion Reintroduction Project plans to establish a second independent population of
Asiatic lions at the
Kuno Wildlife Sanctuary in the Indian state of
Madhya Pradesh.
[148] It is important to start a second population to serve as a
gene pool for the last surviving Asiatic lions and to help develop and maintain
genetic diversity enabling the species to survive.
The former popularity of the Barbary lion as a zoo animal has meant
that scattered lions in captivity are likely to be descended from
Barbary lion stock. This includes twelve lions at
Port Lympne Wild Animal Park in
Kent,
England that are descended from animals owned by the
King of Morocco.
[149] Another eleven animals believed to be Barbary lions were found in
Addis Ababa zoo, descendants of animals owned by
Emperor Haile Selassie. WildLink International, in collaboration with
Oxford University, launched their 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.
[56]
Following the discovery of the decline of lion population in Africa, several coordinated efforts involving lion
conservation have been organised in an attempt to stem this decline. Lions are one species included in the
Species Survival Plan, a coordinated attempt by the
Association of Zoos and Aquariums
to increase its chances of survival. The plan was originally started in
1982 for the Asiatic lion, but was suspended when it was found that
most Asiatic lions in North American zoos were not
genetically pure,
having been hybridised with African lions. The African lion plan
started in 1993, focusing especially on the South African subspecies,
although there are difficulties in assessing the genetic diversity of
captive lions, since most individuals are of unknown origin, making
maintenance of genetic diversity a problem.
[16]
Man-eaters
While lions do not usually hunt people, some (usually males) seem to seek out human prey; well-publicised cases include the
Tsavo maneaters, where 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 in 1898, and the 1991
Mfuwe man-eater, which killed six people in the Luangwa River Valley in
Zambia.
[150]
In both, the hunters who killed the lions wrote books detailing the
animals' predatory behaviour. The Mfuwe and Tsavo incidents bear
similarities: the lions in both incidents were larger than normal,
lacked manes, and seemed to suffer from tooth decay (although only one
of the Tsavo man-eaters did). The infirmity theory, including tooth
decay, is not favored 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.
[151]
In their analysis of Tsavo and man-eating generally, 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 that the relationship is
well-attested amongst other pantherines and primates in the
paleontological record.
[152]
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 more famed "Tsavo" incidents of a
century earlier. The incidents occurred near
Selous National Park in
Rufiji District and in
Lindi Province near the
Mozambican
border. While the expansion of villagers into bush country is one
concern, the authors argue that conservation policy must mitigate the
danger because, in this case, conservation contributes directly to human
deaths. Cases in Lindi have been documented where lions seize humans
from the center of substantial villages.
[153] 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 attacks on people.
[154]
Author Robert R. Frump wrote in
The Man-eaters of Eden that Mozambican refugees regularly crossing Kruger National Park at night in
South Africa
are attacked and eaten by the lions; park officials have conceded that
man-eating is a problem there. Frump believes thousands may have been
killed in the decades after
apartheid
sealed the park and forced the refugees to cross the park at night. For
nearly a century before the border was sealed, Mozambicans had
regularly walked across the park in daytime with little harm.
[155]
Packer estimates more than 200 Tanzanians are killed each year by lions,
crocodiles, elephants, hippos, and
snakes,
and that the numbers could be double that amount, with lions thought to
kill at least 70 of those. Packer has documented that between 1990 and
2004, lions attacked 815 people in Tanzania, killing 563. Packer and
Ikanda are among the few
conservationists
who believe western conservation efforts must take account of these
matters not just because of ethical concerns about human life, but also
for the long term success of conservation efforts and lion preservation.
[153]
A man-eating lion was killed by game scouts in Southern Tanzania in
April 2004. It is believed to have killed and eaten at least 35 people
in a series of incidents covering several villages in the Rufiji Delta
coastal region.
[156]
Dr Rolf D. Baldus, the GTZ wildlife programme coordinator, commented
that it was likely that the lion preyed on humans because it had a large
abscess underneath a
molar
which was cracked in several places. He further commented that "This
lion probably experienced a lot of pain, particularly when it was
chewing."
[157]
GTZ is the German development cooperation agency and has been working
with the Tanzanian government on wildlife conservation for nearly two
decades. As in other cases this lion was large, lacked a mane, and had a
tooth problem.
The "All-Africa" record of man-eating generally is considered to be
not Tsavo, but incidents in the early 1930s through the late 1940s in
what was then
Tanganyika
(now Tanzania). George Rushby, game warden and professional hunter,
eventually dispatched the pride, which over three generations is thought
to have killed and eaten 1,500 to 2,000 people in what is now
Njombe district.
[158]
In captivity
Lions are part of a group of exotic animals that are the core of
zoo
exhibits since the late eighteenth century; members of this group are
invariably large vertebrates and include elephants, rhinoceroses,
hippopotamuses, large primates, and other big cats; zoos sought to
gather as many of these species as possible.
[159] Although many modern zoos are more selective about their exhibits,
[160]
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.
[161] 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.
[162] Breeding programs need to note origins to avoid breeding different subspecies and thus reducing conservation value.
[163]
Male African lion of the Transvaal subspecies (
P. l. krugeri)
At the ancient Egyptian cities of
Taremu and
Per-Bast were temples to the lioness goddesses of Egypt,
Sekhmet and
Bast and at Taremu there was a temple to the son of the deity,
Maahes
the lion prince, where live lions were kept and allowed to roam within
his temple. The Greeks called the city Leontopolis, the "City of Lions"
and documented that practice. Lions were kept and bred by
Assyrian kings as early as 850 BC,
[134] and
Alexander the Great was said to have been presented with tame lions by the
Malhi of northern
India.
[164] Later in
Roman times, lions were kept by emperors to take part in the gladiator arenas or for executions (see
bestiarii,
damnatio ad bestias, and
venatio). Roman notables, including
Sulla,
Pompey, and
Julius Caesar, often ordered the mass slaughter of hundreds of lions at a time.
[165] In the East, lions were tamed by Indian princes, and
Marco Polo reported that
Kublai Khan kept lions inside.
[166]
The first European "zoos" spread amongst noble and royal families in
the thirteenth century, and until the seventeenth 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.
[167] 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 thirteenth century,
[168][169] probably stocked with animals from an earlier menagerie started in 1125 by
Henry I at his palace in
Woodstock, near
Oxford; where lions had been reported stocked by
William of Malmesbury.
[170]
Seraglios served as expressions of the nobility's power and wealth. Animals such as big cats and
elephants,
in particular, symbolised power, and would be pitted in fights against
each other or domesticated animals. By extension, menageries and
seraglios served as demonstrations of the dominance of humanity over
nature. Consequently, the defeat of such natural "lords" by a cow in
1682 astonished the spectators, and the flight of an elephant before a
rhinoceros drew jeers. Such fights would slowly fade out in the
seventeenth century with the spread of the menagerie and their
appropriation by the commoners. The tradition of keeping big cats as
pets would last into the nineteenth century, at which time it was seen
as highly eccentric.
[171]
The presence of lions at the Tower of London was intermittent, being restocked when a monarch or his consort, such as
Margaret of Anjou the wife of
Henry VI,
either sought or were given animals. Records indicate they were kept in
poor conditions there in the seventeenth century, in contrast to more
open conditions in
Florence at the time.
[172]
The menagerie was open to the public by the eighteenth century;
admission was a sum of three half-pence or the supply of a cat or dog
for feeding to the lions.
[173] A rival menagerie at the
Exeter Exchange also exhibited lions until the early nineteenth century.
[174] The Tower menagerie was closed down by
William IV,
[173] and animals transferred to the
London Zoo which opened its gates to the public on 27 April 1828.
[175]
Animal species disappear when they cannot peacefully orbit the center of gravity that is man.
Pierre-Amédée Pichot, 1891
[176]
The wild animals trade flourished alongside improved colonial trade
of the nineteenth century. Lions were considered fairly common and
inexpensive. Although they would barter higher than tigers, they were
less costly than larger, or more difficult to transport animals such as
the giraffe and hippopotamus, and much less than
giant pandas.
[177]
Like other animals, lions were seen as little more than a natural,
boundless commodity that was mercilessly exploited with terrible losses
in capture and transportation.
[178] The widely reproduced imagery of the heroic hunter chasing lions would dominate a large part of the century.
[179] Explorers and hunters exploited a popular
Manichean
division of animals into "good" and "evil" to add thrilling value to
their adventures, casting themselves as heroic figures. This resulted in
big cats, always suspected of being man-eaters, representing "both the
fear of nature and the satisfaction of having overcome it."
[180]
Lion at
Melbourne Zoo enjoying an elevated grassy area with some tree shelter
Lions were kept in cramped and squalid conditions at London Zoo until
a larger lion house with roomier cages was built in the 1870s.
[181] Further changes took place in the early twentieth century, when
Carl Hagenbeck
designed enclosures more closely resembling a natural habitat, with
concrete 'rocks', more open space and a moat instead of bars. He
designed lion enclosures for both
Melbourne Zoo and Sydney's
Taronga Zoo,
among others, in the early twentieth century. Though his designs were
popular, the old bars and cage enclosures prevailed until the 1960s in
many zoos.
[182] In the later decades of the twentieth 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, with some attractions even placing the den on ground higher
than visitors, such as the Cat Forest/Lion Overlook of
Oklahoma City Zoological Park.
[16]
Lions are now housed in much larger naturalistic areas; modern
recommended guidelines more closely approximate conditions in the wild
with closer attention to the lions' needs, highlighting the need for
dens in separate areas, elevated positions in both sun and shade where
lions can sit and adequate ground cover and drainage as well as
sufficient space to roam.
[161]
There have also been instances where a lion was kept by a private individual, such as the lioness
Elsa, who was raised by
George Adamson and his wife
Joy Adamson
and came to develop a strong bond with them, particularly the latter.
The lioness later achieved fame, her life being documented in a series
of books and films.
Baiting and taming
Nineteenth century
etching of a lion tamer in a cage of lions
Lion-baiting is a
blood sport involving the
baiting
of lions in combat with other animals, usually dogs. Records of it
exist in ancient times through until the seventeenth century. It was
finally banned in
Vienna by 1800 and England in 1825.
[183][184]
Lion taming refers to the practice of taming lions for entertainment, either as part of an established
circus or as an individual act, such as
Siegfried & Roy. The term is also often used for the taming and display of other big cats such as
tigers,
leopards, and
cougars. The practice was pioneered in the first half of the nineteenth century by Frenchman Henri Martin and American
Isaac Van Amburgh who both toured widely, and whose techniques were copied by a number of followers.
[185] 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, but truly entered public
consciousness in the early twentieth century with cinema. In
demonstrating the superiority of human over animal, lion taming served a
purpose similar to animal fights of previous centuries.
[185]
The ultimate proof of a tamer's dominance and control over a lion is
demonstrated by placing his head in the lion's mouth. The now iconic
lion tamer's chair was possibly first used by American
Clyde Beatty (1903–1965).
[186]
Cultural depictions
Georgian lion from
Colchis (3000 BC).
The lion has been an icon for humanity for thousands of years,
appearing in cultures across Europe, Asia, and Africa. Despite incidents
of attacks on humans, lions have enjoyed a positive depiction in
culture as strong but noble. A common depiction is their representation
as "
king of the jungle" or "king of beasts"; hence, the lion has been a popular symbol of royalty and stateliness,
[187] as well as a symbol of bravery; it is featured in several
fables of the sixth century BC
Greek storyteller
Aesop.
[188]
Representations of lions date back to the early
Upper Paleolithic. The
lioness-headed ivory carving from Vogelherd cave in the
Swabian Alb in southwestern
Germany, dubbed
Löwenmensch (lion-human) in German. The sculpture has been determined to be at least 32,000 years old from the
Aurignacian culture,
[14] but it may date to as early as 40,000 years ago.
[189] The sculpture has been interpreted as
anthropomorphic, giving human characteristics to an animal, however, it also may represent a
deity.
Two lions were depicted mating in the Chamber of Felines in 15,000-year-old
Paleolithic cave paintings in the
Lascaux caves. Cave lions also are depicted in the
Chauvet Cave, discovered in 1994; this has been dated at 32,000 years of age,
[34] though it may be of similar or younger age to Lascaux.
[190]
Ancient Egypt venerated the lioness (the fierce hunter) as their
war deities and among those in the
Egyptian pantheon are,
Bast,
Mafdet,
Menhit,
Pakhet,
Sekhmet,
Tefnut, and the
Sphinx;
[187] The
Nemean lion was symbolic in Ancient Greece and Rome, represented as the constellation and zodiac sign
Leo, and described in mythology, where its skin was borne by the hero
Heracles.
[191]
The lion was a prominent symbol in ancient
Mesopotamia (from
Sumer up to
Assyrian and
Babylonian times), where it was strongly associated with kingship.
[192] The classic Babylonian lion motif, found as a statue, carved or painted on walls, is often referred to as the
striding lion of Babylon. It is in Babylon that the biblical
Daniel is said to have been delivered from the lion's den.
[193]
In the
Puranic texts of
Hinduism,
Narasimha ("man-lion") a half-lion, half-man incarnation or (
avatar) of
Vishnu, is worshipped by his devotees and saved the child devotee
Prahlada from his father, the evil demon king
Hiranyakashipu;
[194] Vishnu takes the form of half-man/half-
lion, in Narasimha, having a human torso and lower body, but with a lion-like face and claws.
[195] Singh is an
ancient Indian vedic name meaning "lion" (
Asiatic lion), dating back over 2000 years to
ancient India. It was originally only used by
Rajputs a
Hindu Kshatriya or military
caste in India. After the birth of the
Khalsa brotherhood in 1699, the
Sikhs also adopted the name "Singh" due to the wishes of
Guru Gobind Singh. Along with millions of Hindu Rajputs today, it is also used by over 20 million
Sikhs worldwide.
[196] Found famously on numerous
flags and
coats of arms all across Asia and Europe, the Asiatic lions also stand firm on the
National Emblem of India.
[197] Farther south on the
Indian subcontinent, the Asiatic lion is symbolic for the
Sinhalese,
[198] Sri Lanka's ethnic majority; the term derived from the Indo-Aryan
Sinhala, meaning the "lion people" or "people with lion blood", while a sword wielding lion is the central figure on the national
flag of Sri Lanka.
[199]
The Asiatic lion is a common motif in
Chinese art. They were first used in art during the late
Spring and Autumn Period (fifth or sixth century BC), and became much more popular during the
Han Dynasty (206 BC – AD 220), when
imperial guardian lions
started to be placed in front of imperial palaces for protection.
Because lions have never been native to China, early depictions were
somewhat unrealistic; after the introduction of
Buddhist art to China in the
Tang Dynasty
(after the sixth century AD), lions usually were depicted without
wings, their bodies became thicker and shorter, and their manes became
curly.
[200] The
lion dance is a form of traditional dance in
Chinese culture
in which performers mimic a lion's movements in a lion costume, often
with musical accompaniment from cymbals, drums, and gongs. They are
performed at
Chinese New Year, the
August Moon Festival and other celebratory occasions for good luck.
[201]
The
island nation of
Singapore derives its name from the
Malay words
singa (lion) and
pora (city/fortress), which in turn is from the
Tamil-
Sanskrit சிங்க
singa सिंह siṃha and
पुर புர
pura, which is cognate to the
Greek πόλις,
pólis.
[202] According to the
Malay Annals, this name was given by a fourteenth-century
Sumatran Malay prince named
Sang Nila Utama, who, on alighting the island after a thunderstorm, spotted an auspicious beast on shore which appeared to be a lion.
[203]
The name of the nomadic
Hadendoa people, inhabiting parts of
Sudan,
Egypt, and
Eritrea, is made up of
haɖa 'lion' and
(n)ɖiwa 'clan'. Other variants are
Haɖai ɖiwa,
Hanɖiwa, and
Haɖaatʼar (children of lioness).
"Lion" was the nickname of several medieval warrior rulers with a reputation for bravery, such as the English King
Richard the Lionheart,
[187] Henry the Lion, (
German:
Heinrich der Löwe),
Duke of Saxony,
William the Lion, King of Scotland, and
Robert III of Flanders nicknamed "The Lion of Flanders"—a major
Flemish national icon up to the present. Lions are frequently depicted on
coats of arms, either as a device on shields themselves, or as
supporters, but the lioness is much more infrequent.
[204] The formal language of
heraldry, called
blazon,
employs French terms to describe the images precisely. Such
descriptions specified whether lions or other creatures were "rampant"
or "passant", that is whether they were rearing or crouching.
[205] The lion is used as a symbol of sporting teams, from national association football teams such as
England,
Scotland and
Singapore to famous clubs such as the
Detroit Lions[206] of the NFL,
Chelsea[207] and
Aston Villa of the English
Premier League,
[208] (and the Premiership itself),
Eintracht Braunschweig of the
Bundesliga, and to a host of smaller clubs around the world.
Lions continue to be featured in modern literature, from the messianic
Aslan in
The Lion, the Witch and the Wardrobe and following books from
The Chronicles of Narnia series written by
C. S. Lewis,
[209] to the comedic
Cowardly Lion in
The Wonderful Wizard of Oz.
[210]
The advent of moving pictures saw the continued presence of lion
symbolism; one of the most iconic and widely recognised lions is
Leo the Lion, which has been the mascot for
Metro-Goldwyn-Mayer (MGM) studios since the 1920s.
[211] The 1960s saw the appearance of what is possibly the most famous lioness, the Kenyan animal
Elsa in the movie
Born Free,
[212] based on the true-life international bestselling book of the same title.
[213]
The lion's role as King of the Beasts has been used in cartoons, from
the 1950s manga which gave rise to the first Japanese colour TV
animation series,
Kimba the White Lion, Leonardo Lion of
King Leonardo and his Short Subjects, both from the 1960s, up to the 1994
Disney animated feature film
The Lion King,
[214][215] which also featured the popular song "
The Lion Sleeps Tonight" in its soundtrack. A lion appears on the 50-rand
South African banknote.
Heraldic depictions
See also
