The lion (Panthera leo) is one of the big
cats in the genus Panthera and a member of the family Felidae. The commonly used term African lion collectively denotes the several
subspecies in Africa. With some males exceeding 250 kg (550 lb) in
weight, it is the second-largest living cat after the tiger, barring hybrids like the liger. Wild lions currently exist in sub-Saharan Africa and
in India (where an endangeredremnant population resides in and
around Gir Forest National Park). In ancient historic times,
their range was in most of Africa, including North
Africa, and
across Eurasia from Greece and southeastern Europe to India. In the late Pleistocene, about 10,000 years ago, the lion
was the most widespread large land mammal after humans: Panthera leo spelaea lived in northern and western
Europe and Panthera leo atrox lived in the Americas from the Yukon to Peru. The lion is classified as a vulnerable species by
the International Union for Conservation of Nature (IUCN), having seen a major
population decline in its African range of 30–50% over two decades during the
second half of the twentieth century. 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 the
greatest causes of concern. Within Africa, the West African lionpopulation
is particularly endangered.
In the wilderness, males seldom live longer than 10 to 14
years, as injuries sustained from continual fighting with rival males greatly
reduce their longevity. In captivity they can live more than 20 years.
They typically inhabit savanna and grassland, although they may take to bushand 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 are also expert scavengers obtaining over 50 percent of their
food by scavenging as opportunity allows. While lions do not typically hunt
humans, some have. Sleeping mainly during the day, lions are active primarily
at night (nocturnal), although sometimes at twilight (crepuscular).
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 in France
dated to 17,000 years ago, 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, and the Ancient Greek λέων (leon). The Hebrewword לָבִיא (lavi) may also be related. It was one of the
species originally described by Linnaeus, who gave it the name Felis leo, in his eighteenth-century
work, Systema Naturae.
Taxonomy and evolution
The lion's closest relatives are the
other species of the genus Panthera: the tiger, the snow
leopard, the jaguar, and the leopard. Studies from 2006 and 2009 concluded
that the jaguar is a sister
species to the
lion and the leopard is a sister taxon to the jaguar/lion clade while 2010
and 2011 studies have swapped the positions leopard and jaguar. P. leo evolved in Africa between 1 million and 800,000 years
ago, before spreading throughout the Holarctic region. 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. Lions died out in northern Eurasia at the end of the last glaciation, about 10,000 years ago; this may have
been secondary to the extinction of Pleistocene megafauna.[21][22]
Subspecies
Range
map of the commonly accepted subspecies of the lion in the late twentieth
century
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. Today,
only eight subspecies are usually accepted, although one of these,
the Cape lion, formerly described as Panthera leo melanochaita, is probably invalid. 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 have sometimes 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, West-Central African lions are more closely related to North
African and Asian lions than to Southern or Eastern African lions. These
findings might be explained by a late Pleistocene extinction event of lions in
western and central Africa, and a subsequent recolonisation of these parts from
Asia.
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. Another study revealed there are three major types of
lions, one North African–Asian, one southern African and one middle
African. 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.
Based on recent genetic studies, the Cat
Classification Task Force of the IUCN SSC Cat Specialist Group has
provisionally proposed to assign the lions occurring in Asia (P. l. persica) and West, Central and North Africa (P. l. senegalensis, and P. l. leo) to the subspecies Panthera leo leo, and the lions inhabiting Southern (P. l. bleyenberghi and P. l. krugeri) and Eastern Africa (P. l. azandica and P. l. nubica) to the subspecies Panthera leo
melanochaita. The U.S. Fish and Wildlife Service has followed this revised
taxonomic classification, as being based on "the best available scientific
and commercial information", in listing these two subspecies as,
respectively, endangered and threatened.
The majority of 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 that are extinct in the wild, and might be
therefore important to maintain overall genetic variability of the
lion. 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
Recent
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Subspecies
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Description
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Is found in Gir Forest National Park of northwestern India. Once was widespread from Turkey, across Southwest Asia, to India
and Pakistan, now 523 exist in and near the Gir Forest in
the Saurashtran region of Gujarat. Genetic evidence suggests its
ancestors split from the ancestors of sub-Saharan African lions between 203
and 74 thousand years ago. Its closest relatives are North and West-Central African lions.
West Asia: (Armenia, Azerbaijan, Baluchistan, Georgia, Iran, Iraq, Israel, Jordan, Kuwait, Lebanon, Mesopotamia, North Caucasus, Oman, Saudi Arabia, Syria, Turkey, United Arab Emirates and Yemen)
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Formerly found in Algeria, Morocco, Tunisia, and Egypt, this is the nominate lion subspecies
from North Africa. It is extinct in the wild due to excessive hunting; the
last, known Barbary lion in the wilderness was killed in Morocco in
1920. This is reportedly one of the largest subspecies, 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. Besides the Senegal lion of
Western and Central Africa, it is more closely related to the Asiatic lion
than to other African lions. A number of animals in captivity are likely
to be Barbary lions, particularly the 90 animals descended from the
Moroccan Royal collection at Rabat Zoo.
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Found in Western and Central Africa,
from Senegal to the Central African Republic. It is currently listed as critically endangeredin 2015. It is among the smallest of
the Sub-Saharan African lions.
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A newly discerned lion subspecies
could exist in captivity in Ethiopia's capital city of Addis Ababa. 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 seem 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.
Northeastern Africa: (Ethiopia)
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Found in East Africa, from Ethiopia and Kenya to Tanzania and Mozambique;[41] a local population is known as
the Tsavo lion.
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Found in the northeastern parts of
the Congo, adjacent to Uganda. It is currently extinct
in Rwanda.
Central or Eastern
Africa: (Central African Republic, Democratic Republic of the Congo, Rwanda, South Sudan, Sudan, Uganda)
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Found in southwestern Africa
and the Democratic Republic of the Congo. It is among the largest subspecies
of African lions.
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Characteristics
A skeletal mount of an African lion attacking a common eland on display at The Museum of
Osteology, Oklahoma City,
Oklahoma.
Lions are muscular, deep-chested felids
with short, rounded heads, reduced necks and round ears. Of the living,
non-hybrid felids, the lion is second only to the tiger 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 and broader nasal openings than that of a 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. Lion colouration varies from light buff to silverly gray, to
yellowish red and dark brown. The underparts are generally lighter and
cubs are born with dark rosettes (spots)
on their body. The spots fade as lions reach adulthood, although faint spots
often may still be seen on the legs and underparts.
Lions are the only members of the cat
family to display obvious sexual dimorphism –
males are more robust with broader heads and have a prominent mane which grows
downwards and backward and may cover most of the head, neck, shoulders and
chest. The mane is typically brownish and tinged with yellow, rust and black
hairs. The most distinctive characteristic shared by both females and males is
that the tail ends in a dark, 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 5 1⁄2 months of age and is readily identifiable at
7 months.
Nowak indicates the typical weight range
of lions as 150 to 250 kg (331 to 551 lb) for males and 120 to
182 kg (265 to 401 lb) for females. The size of adult lions
varies across their range, with those from the southern African populations in
Zimbabwe, the Kalahari and Kruger Park (P. l. bleyenberghi, P. l. vernayi, or P. l. krugeri) averaging around 189.6 kg
(418 lb) and 126.9 kg (280 lb) in males and females
respectively, compared to 174.9 kg (386 lb) and 119.5 kg
(263 lb) of male and female lions from East Africa. Reported body measurements
in males are head-body lengths ranging from 170 to 250 cm (5 ft
7 in to 8 ft 2 in), tail lengths of 90–105 cm (2 ft
11 in–3 ft 5 in). In females reported head-body lengths range
from 140 to 175 cm (4 ft 7 in to 5 ft 9 in), tail
lengths of 70–100 cm (2 ft 4 in–3 ft
3 in), 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 centimetres
less in length.
Record measurements from hunting records
are supposedly a total length of nearly 3.6 m (12 ft) for a
male Southwest African lion shot near Mucusso, southern Angola in October 1973, and a weight of
313 kg (690 lb) for a male Southeast African lion shot outside Hectorspruit in eastern Transvaal, South
Africa in 1936. Another notably outsized male lion, which was a East African lion was
shot near Mount Kenya, weighed in at 272 kg (600 lb). There are also
reports of large Asiatic lions.
Mane
During
agonistic confrontations with other lions, the mane makes the lion appear
larger.
The mane of the adult male lion, unique among cats, is
one of the most distinctive characteristics of the species. In rare cases a
female lion can have a mane.The male lion's mane serves two main purposes:
Defense of the neck from opponents, and attracting females for mating.
The presence, absence, colour, and size
of the mane is associated with genetic precondition, sexual maturity, climate,
and testosteroneproduction; the rule of thumb is the darker and fuller the
mane, the healthier the lion. Sexual selection of mates by lionesses favours
males with the densest, darkest mane. 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.
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, which had the
thickest, most extensive manes amongst wild lions. Research has suggested,
however, that environmental factors such as average ambient temperature influence the colour and size of a lion's mane. 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. The males of the Asiatic subspecies, however, are
characterised by sparser manes than average African lions.
In the area of Pendjari National Park, almost all West African males are
maneless or have very weak manes. Maneless, male African lions have also
been reported from Senegal, from Sudan (Dinder National Park), 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. In
addition, increased testosterone may be the cause of the maned lionesses of
northern Botswana.
Cave paintings of
extinct European cave lions almost exclusively show animals with no manes,
suggesting that either they were maneless, or that the paintings depict
lionesses, as seen hunting in a group.[citation needed] Nevertheless, in the Chauvet cave,
there is a drawing of two maneless lions, with one of them having a scrotum.
White lions
The white
lion is not a
distinct subspecies, but a special morph with a genetic condition, leucism, 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
caused by a recessive allele. Reportedly, they have been bred in camps in South
Africa for use as trophies to be killed during canned
hunts.
Behaviour
Lions spend much of their time resting and are
inactive for about 20 hours per day. 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 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.
Group organisation
Lions are the most socially inclined of
all wild felids, most of which remain quite solitary in nature, and have two
types of social organization. Some lions are residents, living in groups of related
lionesses, their mates, and offspring. Such a group is called a pride. Females form the stable social
unit in a pride
and do not tolerate outside females. Membership only changes with the
births and deaths of lionesses, although some females do leave and become
nomadic. Although extremely large prides, consisting of up over 30 individuals,
have been observed, the average pride consists of around fifteen lions;
including several adult females, up to four males (known as a coalition if more than one) and their cubs
of both sexes. The sole exception to this pattern is the Tsavo
lion pride which
always has just one adult male. Male cubs are excluded from their maternal
pride when they reach maturity at around 2–3 years of age. The second
organizational behaviour is labeled nomads, who range widely and move about sporadically, either
singularly or in pairs. 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. Interactions
between prides and nomads tend to be hostile, although pride females in estrous
may allow nomad males to approach them.
The area a pride occupies is called
a pride
area, whereas that by a
nomad is a range. 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 calorific 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.
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. Some individuals consistently lead the
defence against intruders, while others lag behind. Lions tend to assume
specific roles in the pride. Those lagging behind may provide other valuable
services to the group. 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. 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.
Hunting and diet
Lions are generalist hypercarnivores[92] and usually hunt in groups. Their prey consists mainly of mammals,
particularly ungulates, with a preference for wildebeest, zebras, buffalo, gemsbok, and giraffes in Africa and chital, sambar
deer, nilgai, wild
boar, chinkara and chousingha in India. African lions
prefer prey weighing 190–550 kg (420–1,210 lb). They generally
avoid fully grown adult elephants, hippopotamuses, rhinoceroses, as well as very small prey like dik-dik, hyrax, hare and vervet monkey. However, Thomson's gazelles may be hunted and warthogs are often taken depending on
availability, despite being below the preferred weight range. In most
areas, a small number of species may make up around three-fourths of the lion's
diet. In Serengeti National Park, wildebeest, zebras and gazelle are the
majority of prey. In Kruger National Park, giraffes are the most common
prey, and in Manyara Park, Cape buffaloes constitute as much as 62% of the
lion's diet, due to the high number density of buffaloes. At the Okavango Delta,
with its strong seasonal changes in prey, up to eight species may make up
third-fourths of lion's diet. Occasionally adult hippopotamus are taken
at Gorongosa National Parkand calves are commonly hunted at Virunga National Park. In addition to size, the aquatic
nature of hippos makes them normally unavailable as prey. The lions
of Savuti, Botswana, have adapted to hunting young elephants during
the dry season, and a pride of 30 lions has been recorded killing individuals
between the ages of four and eleven years. Lions also attack domestic
livestock and in India cattle contribute significantly to their
diet. Unusual prey items include porcupines and small reptiles. Lions will kill other predators
such as leopards, cheetahs, and hyenas, but they seldom devour them.
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. Single lions are capable of bringing down prey like zebra and
wildebeest, which can be twice their own weight, while hunting larger prey like
giraffes and buffalo alone are too much of a risk. Cooperative-hunting lions
are usually successful. In prides, lionesses do most of the hunting.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 in flight from other
lionesses. Males attached to prides do not usually participate in group
hunting. However, some evidence suggests that pride males are just as
successful as females; they are solo hunters who ambush prey in small
bush. Lions 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. Thus, they only run fast in short bursts, 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.
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, 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). Prey is typically eaten
at the location of the hunt, although large prey is sometimes dragged into
cover. Lions tend to squabble over a kill, particularly the males. When
food is scarce, cubs tend to suffer the most but otherwise all pride members
can eat their fill, including old and crippled ones which can live on
leftovers. There is more sharing of larger kills. An adult lioness
requires an average of about 5 kg (11 lb) of meat per day, a male
about 7 kg (15 lb). A lion may gorge itself and eat up to
30 kg (66 lb) in one sitting; 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. Lions
will defend their kills from scavengers like vultures and hyenas.
Lions prefer to scavenge when the opportunity presents
itself with carrion providing more than 50% of their diet. They
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. In fact, most dead prey on which both hyenas and lions feed upon
are killed by the hyenas instead of the lions.
Predator competition
Lions and spotted hyenas occupy
a similar ecological niche and compete for prey and carrion in the areas where
they coexist. A review of data across several studies indicates a dietary
overlap of 58.6%. 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 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. On the other hand, in Northern
Botswana's Chobe National Park, the situation is reversed: hyenas frequently
challenge lions and steal their kills: they obtain food from 63% of all lion
kills. When confronted on a kill by lions, spotted hyenas may either leave
or wait patiently at a distance of 30–100 m (98–328 ft) until the
lions have finished, but they are also bold enough to feed alongside
lions, and even force the lions off a kill. The two species may attack one
another even when there is no food involved for no apparent reason. 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. 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. 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 felids
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 in particular has a 50% chance of losing its kill to
lions or other predators. Lions are major killers of cheetah cubs, accounting
for up to 78.2% of predator-killed juveniles in one study. Cheetahs avoid
their competitors using different temporal (time) and spatial (habitat)
niches. Leopards are able to take refuge in trees; however, lionesses will
occasionally be successful in climbing to retrieve leopard
kills. 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. However, there are a few reported cases of old and wounded lions
falling prey to wild dogs. African lions may also conflict with Nile crocodiles.
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, while the reverse is true for lions entering waterways, as evidenced
by the occasional lion claw found in crocodile stomachs.
Man-eating
While lions do not usually hunt people,
some (usually males) seem to seek out human prey; one well-publicised case
includes 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. The hunter who killed the lions wrote a book detailing the animals'
predatory behaviour. The lions were larger than normal, lacked manes, and one
seemed to suffer from tooth decay. The infirmity theory, including tooth decay,
is not favoured by all researchers; an analysis of teeth and jaws of man-eating
lions in museum collections suggests that while tooth decay may explain some
incidents, prey depletion in human-dominated areas is a more likely cause of
lion predation on humans.
In their analysis of Tsavo and general man-eating,
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
among other pantherines and primates in the paleontological record.
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. 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.
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.
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.
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. 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 that
was cracked in several places. He further commented that "This lion
probably experienced a lot of pain, particularly when it was
chewing." 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.
Sometimes, even Asiatic lions may become
man-eaters. Their rather large number for a place like Gir Forest National Park means that a number of lions have
moved outside the protected area, making them a potential threat to humans even
outside the park. Two such incidents happened in 2012, including in area about
50–60 km (31–37 mi) from Gir Park.
Reproduction and life cycle
Mating
Most lionesses will have reproduced by
the time they are four years of age. Lions do not mate at any specific time of year, and
the females are polyestrous. 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. A lioness may mate with
more than one male when she is in heat.
The average gestation period is around
110 days, 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. 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. 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.
Usually, the mother does not integrate herself and her
cubs back into the pride until the cubs are six to eight weeks
old. 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 and
there is communal 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. 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, and older cubs do not dominate the
sucklings.
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 among 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.
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, 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, perhaps because females do not become
fertile and receptive until their cubs mature or die. A lioness often will
attempt to defend her cubs fiercely from a usurping male, but such actions are
rarely successful. Success is more likely when a group of three or four mothers
within a pride join forces against one male. Other sources of mortality
for cubs include starvation and abandonment as well such as predation by
leopards, hyenas and wild dogs. All in all, as many as 80% of the cubs
will die before the age of two.
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.
Both males and females may
interact homosexually. Lions are shown to be involved in group homosexual and
courtship activities. Male lions will also head rub and roll around with each
other before simulating sex together.
Health
Although adult lions have no natural predators,
evidence suggests that the majority die violently from humans or other
lions. 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. 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.
Various species of tick commonly infest the ears, neck and
groin regions of most lions. Adult forms of several species of the
tapeworm genus Taenia have been isolated from intestines, the lions having
ingested larval forms from antelope meat. 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. A more
recent outbreak in 2001 killed six lions.Lions, especially in captivity, are
vulnerable to the canine distemper virus
(CDV), feline immunodeficiency virus (FIV), and feline infectious peritonitis (FIP). 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. 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.
Communication
When resting, lion socialisation 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, which have been compared with grooming in
primates.Head rubbing – nuzzling one's forehead, face and neck against another
lion – appears to be a form of greeting, 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. 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.
Lions have an array of facial
expressions and body postures that serve as visual gestures. 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. Lions also use chemical and visual marking; male lions will spray and scrape both plots of ground and objects within their
territory.
Their repertoire of vocalisations is
also large; variations in intensity and pitch, rather than discrete signals,
appear central to communication. Most lion vocals are variations of
growling/snarling, miaowing and roaring. Other sounds produced include purring,
puffing, bleating and humming. 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. 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. Lions have the loudest roar of any big
cat