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Writer's pictureMichael Constantine

Lion

Updated: May 2, 2020

All you need to know about the lions.



The lion (Panthera leo) is a species in the family Felidae; it is a muscular, deep-chested cat with a short, rounded head, a reduced neck and round ears, and a hairy tuft at the end of its tail.


It is sexually dimorphic; adult male lions have a prominent mane, which is the most recognisable feature of the species.


With a typical head-to-body length of 184–208 cm (72–82 in) they are larger than females at 160–184 cm (63–72 in). It is a social species, forming groups called prides.


A lion pride consists of a few adult males, related females and cubs. Groups of female lions usually hunt together, preying mostly on large ungulates.


The lion is an apex and keystone predator, although some lions scavenge when opportunities occur, and have been known to hunt humans, although the species typically does not. Typically, the lion inhabits grasslands and savannas, but is absent in dense forests. It is usually more diurnal than other big cats, but when persecuted it adapts to being active at night and at twilight.


In the Pleistocene, the lion ranged throughout Eurasia, Africa and North America, but today it has been reduced to fragmented populations in Sub-Saharan Africa and one critically endangered population in western India.


It has been listed as Vulnerable on the IUCN Red List since 1996 because populations in African countries have declined by about 43% since the early 1990s.


Lion populations are untenable outside designated protected areas.


Although the cause of the decline is not fully understood, habitat loss and conflicts with humans are the greatest causes for concern. One of the most widely recognised animal symbols in human culture, the lion has been extensively depicted in sculptures and paintings, on national flags, and in contemporary films and literature.


Lions have been kept in menageries since the time of the Roman Empire and have been a key species sought for exhibition in zoological gardens across the world since the late 18th century.


Cultural depictions of lions were prominent in the Upper Paleolithic period; carvings and paintings from the Lascaux and Chauvet Caves in France have been dated to 17,000 years ago, and depictions have occurred in virtually all ancient and medieval cultures that coincided with the lion's former and current ranges.


Lion Documentary Film



Evolution


The lion's closest relatives are the other species of the genus Panthera, namely tiger, snow leopard, jaguar, and leopard. The origin of the Panthera is most likely northern Central Asia during the Miocene. Results of phylogenetic studies published in 2006 and 2009 indicate that the jaguar and the lion belong to one sister group that diverged about 2.06 million years ago.

Results of later studies indicate that the leopard and the lion belong to the same sister group, which diverged 3.1–1.95 million years ago.

The earliest lions are thought to have originated in Asia or Africa, with the linages of modern and cave lions diverging in the Pliocene, around 1.89 million years ago. Hybridisation between ancestors of lion and snow leopard lineages may have continued until about 2.1 million years ago.

The Eurasian and American cave lions became extinct at the end of the last glacial period without mitochondrial descendants on other continents.

The modern lion was probably widely distributed in Africa during the Middle Pleistocene and started to diverge in sub-Saharan Africa during the Late Pleistocene.

Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183,500–81,800 years ago.

The expansion of the Sahara 83,100–26,600 years ago caused the separation of populations in West and North Africa.

As the rainforest decreased, thus giving rise to more open habitats, lions moved from West to Central Africa. Lions from North Africa dispersed to southern Europe and Asia 38,800–8,300 ago. Extinction of lions in southern Europe, North Africa and the Middle East interrupted gene flow between lion populations in Asia and Africa.

Genetic evidence revealed numerous mutations in lion samples from East and Southern Africa, which indicates that this group has a longer evolutionary history than genetically less diverse lion samples from Asia and West and Central Africa.

Results of phylogeographic research indicate that the two living lion subspecies diverged about 245,000 years ago.


Hybrids



In zoos, lions have been bred with tigers to create hybrids for the curiosity of visitors or for scientific purpose.


The liger is bigger than a lion and a tiger, whereas most tigons are relatively small compared to their parents because of reciprocal gene effects.


The leopon is a hybrid between a lion and leopard.


Description


The lion is a muscular, deep-chested cat with a short, rounded head, a reduced neck and round ears. Its fur varies in colour from light buff to silvery grey, yellowish red and dark brown. The colours of the underparts are generally lighter.

A new-born lion has dark spots, which fade as the cub reaches adulthood, although faint spots often may still be seen on the legs and underparts.

The lion is the only member of the cat family that displays obvious sexual dimorphism. Males have broader heads and a prominent mane that grows downwards and backwards covering most of the head, neck, shoulders, and chest.

The mane is typically brownish and tinged with yellow, rust and black hairs. The tail of all lions ends in a dark, hairy tuft that in some lions conceals an approximately 5 mm (0.20 in)-long, hard "spine" or "spur" that is formed from the final, fused sections of tail bone.


The functions of the spur are unknown. The tuft is absent at birth and develops at around ​5 1⁄2 months of age. It is readily identifiable by the age of seven months.

Of the living felid species, the lion is rivaled only by the tiger in length, weight and height at the shoulder.

Its skull is very similar to that of the tiger, although the frontal region is usually more depressed and flattened, and has a slightly shorter postorbital region and broader nasal openings than those of the tiger.


Due to the amount of skull variation in the two species, usually only the structure of the lower jaw can be used as a reliable indicator of species. The size and weight of adult lions varies across global range and habitats. Accounts of a few individuals that were larger than average exist from Africa and India.



Color Variation



The white lion is a rare morph with a genetic condition called leucism that is caused by a double recessive allele. It is not albino; it has normal pigmentation in the eyes and skin.

White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool.

Nevertheless, 17 births have been recorded in five prides between 2007 and 2015. White lions are selected for breeding in captivity.

They have reportedly been bred in camps in South Africa for use as trophies to be killed during canned hunts.


Mane



The lion's mane is the most recognisable feature of the species.

It starts growing when lions are about a year old. Mane colour varies and darkens with age; research shows its colour and size are influenced by environmental factors such as average ambient temperature.

Mane length apparently signals fighting success in male–male relationships; darker-maned individuals may have longer reproductive lives and higher offspring survival, although they suffer in the hottest months of the year.

The presence, absence, colour and size of the mane are associated with genetic precondition, sexual maturity, climate and testosterone production; the rule of thumb is that a darker, fuller mane indicates a healthier animal.

In Serengeti National Park, female lions favour males with dense, dark manes as mates. The main purpose of the mane is thought be the protection of the neck and throat in territorial fights with rivals.

Cool ambient temperature in European and North American zoos may result in a heavier mane. Asiatic lions usually have sparser manes than average African lions. Almost all male West African lions in Pendjari National Park are either maneless or have very short manes.

Maneless lions have also been reported in Senegal, in Sudan's Dinder National Park and in Tsavo East National Park, Kenya.

The original male white lion from Timbavati in South Africa was also maneless. The hormone testosterone has been linked to mane growth; castrated lions often have little to no mane because the removal of the gonads inhibits testosterone production.

Increased testosterone may be the cause of maned lionesses reported in northern Botswana. The lion's mane may have evolved around 320,000–190,000 years ago.

Cave paintings of extinct Eurasian cave lions almost exclusively show hunting animals without a mane; some suggest that this is evidence that they were indeed maneless.


Distribution and Habitat


African lions live in scattered populations across Sub-Saharan Africa.

The lion prefers grassy plains and savannahs, scrub bordering rivers and open woodlands with bushes. It is absent from rainforest and rarely enters closed forest.

On Mount Elgon, the lion has been recorded up to an elevation of 3,600 m (11,800 ft) and close to the snow line on Mount Kenya.

Lions occur in savannah grasslands with scattered acacia trees, which serve as shade.

The Asiatic lion now only survives in and around Gir National Park in Gujarat, western India.

Its habitat is a mixture of dry savannah forest and very dry, deciduous scrub forest.


Behaviour and Ecology


Lions spend much of their time resting; they 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 continue until dawn, when hunting most often takes place. They spend an average of two hours a day walking and 50 minutes eating.


Group Organisation



The lion is the most social of all wild felid species, living in groups of related individuals with their offspring. Such a group is called a "pride". Groups of male lions are called "coalitions".

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 leave and become nomadic.

The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Large prides, consisting of up to 30 individuals, have been observed.

The sole exception to this pattern is the Tsavo lion pride that always has just one adult male.

Male cubs are excluded from their maternal pride when they reach maturity at around two or three years of age. Some lions are "nomads" that range widely and move around sporadically, either in pairs or alone. Pairs are more frequent among related males who have been excluded from their birth pride. A lion may switch lifestyles; nomads can become residents and vice versa.

Interactions between prides and nomads tend to be hostile, although pride females in estrus allow nomadic males to approach them.

Males spend years in a nomadic phase before gaining residence in a pride.

A study undertaken in the Serengeti National Park revealed that nomadic coalitions gain residency at between 3.5 and 7.3 years of age.

In Kruger National Park, dispersing male lions move more than 25 km (16 mi) away from their natal pride in search of their own territory. Females lions stay closer to their natal pride.

Therefore, female lions in an area are more closely related to each other than male lions in the same area.

The area occupied by a pride is called a "pride area" whereas that occupied by a nomad is a "range".

Males associated with a pride tend to stay on the fringes, patrolling their territory. The reasons for the development of sociality in lionesses – the most pronounced in any cat species – are the subject of much debate.

Increased hunting success appears to be an obvious reason, but this is uncertain upon examination; coordinated hunting allows for more successful predation but also ensures non-hunting members reduce per capita calorific intake.

Some females, however, take a role raising cubs that may be left alone for extended periods. Members of the pride tend to regularly play the same role in hunts and hone their skills.

The health of the hunters is the primary need for the survival of the pride; hunters are the first to consume the prey at the site it is taken.

Other benefits include possible kin selection; sharing food within the family; protecting the young, maintaining territory and individual insurance against injury and hunger.

Both males and females defend the pride against intruders, but the male lion is better-suited for this purpose due to its stockier, more powerful build. Some individuals consistently lead the defence against intruders, while others lag behind.

Lions tend to assume specific roles in the pride; slower-moving individuals may provide other valuable services to the group.

Alternatively, there may be rewards associated with being a leader that fends off intruders; the rank of lionesses in the pride is reflected in these responses.

The male or males associated with the pride must defend their relationship with the pride from outside males who may attempt to usurp them.

Asiatic lion prides differ in group composition. Male Asiatic lions are solitary or associate with up to three males, forming a loose pride while females associate with up to 12 other females, forming a stronger pride together with their cubs.

Female and male lions associate only when mating.

Coalitions of males hold territory for a longer time than single lions. Males in coalitions of three or four individuals exhibit a pronounced hierarchy, in which one male dominates the others and mates more frequently.


Hunting and Diet



The lion is a generalist hypercarnivore and is considered to be both an apex and keystone predator due to its wide prey spectrum.

Its prey consists mainly of mammals – particularly ungulates – weighing 190–550 kg (420–1,210 lb) with a preference for blue wildebeest, plains zebra, African buffalo, gemsbok and giraffe.

Lions also hunt common warthog depending on availability, although the species is below the preferred weight range.

In India, sambar deer and chital are the most commonly recorded wild prey, while domestic livestock may contribute significantly to their diet.

They usually avoid fully grown adult elephants, rhinoceroses and hippopotamus, as well as small prey like dik-dik, hyrax, hare and monkey.

Unusual prey items include porcupines and small reptiles. Lions kill other predators such as leopard, cheetah and spotted hyena but seldom consume them.


Young lions first display stalking behaviour at around three months of age, although they do not participate in hunting until they are almost a year old and begin to hunt effectively when nearing the age of two.

Single lions are capable of bringing down zebra and wildebeest, while larger prey like buffalo and giraffe are riskier.

In Chobe National Park, large prides have been observed hunting African bush elephants up to around 15 years old.

In typical hunts, each lioness has a favoured position in the group, either stalking prey on the "wing", then attacking, or moving a smaller distance in the centre of the group and capturing prey fleeing from other lionesses. Males attached to prides do not usually participate in group hunting.

Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland.


Lions are not particularly known for their stamina; for instance, a lioness' heart comprises only 0.57% of her body weight and a male's is about 0.45% of his body weight, whereas a hyena's heart comprises almost 1% of its body weight.

Thus, lions only run quickly 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 lion's attack is short and powerful; they attempt to catch prey with a fast rush and final leap. They usually pull it down by the rump and kill by a strangling bite to the throat. They also kill prey by enclosing its mouth and nostrils in their jaws.

Lions typically consume prey at the location of the hunt but sometimes drag large prey into cover. They tend to squabble over kills, particularly the males. Cubs suffer most when food is scarce but otherwise all pride members eat their fill, including old and crippled lions, which can live on leftovers.Large kills are shared more widely among pride members.

An adult lioness requires an average of about 5 kg (11 lb) of meat per day while males require about 7 kg (15 lb).

Lions gorge themselves and eat up to 30 kg (66 lb) in one session; if it is unable to consume all of the kill, it rests for a few hours before continuing to eat. On hot days, the pride retreats to shade with one or two males standing guard.

Lions defend their kills from scavengers such as vultures and hyenas.

Lions scavenge on carrion when the opportunity arises; they scavenge animals dead from natural causes such as disease or those that were killed by other predators. Scavenging lions keep a constant lookout for circling vultures, which indicate the death or distress of an animal.

Most carrion on which both hyenas and lions feed upon are killed by hyenas rather than lions. Carrion is thought to provide a large part of lion diet.


Predator Competition



Lions and spotted hyenas occupy a similar ecological niche and where they coexist they compete for prey and carrion; a review of data across several studies indicates a dietary overlap of 58.6%.

Lions typically ignore spotted hyenas unless the lions are on a kill or are being harassed by the hyenas, while the latter tend to visibly react to the presence of lions, with or without the presence of food.

Lions seize the kills of spotted hyenas; in the Ngorongoro crater it is common for lions to subsist largely on kills stolen from hyenas, causing the hyenas to increase their kill rate.

In Botswana's Chobe National Park, the situation is reversed; hyenas frequently challenge lions and steal their kills, obtaining food from 63% of all lion kills.

When confronted on a kill by lions, spotted hyenas may either leave or wait patiently at a distance of 30–100 m (100–330 ft) until the lions have finished.

Hyenas are bold enough to feed alongside lions and to force the lions off a kill. The two species attack one another even when there is no food involved for no apparent reason.

Lion predation can account for up to 71% of hyena deaths in Etosha National Park. Spotted hyenas have adapted by frequently mobbing lions that enter their territories.

When the lion population in Kenya's Masai Mara National Reserve declined, the spotted hyena population increased rapidly.

Experiments on captive spotted hyenas show that specimens without prior experience with lions act indifferently to the sight of them, but will react fearfully to lion scent.

Lions tend to dominate cheetahs and leopards, steal their kills and kill their cubs and even adults when given the chance.

Cheetahs in particular often lose their kills to lions or other predators.

A study in the Serengeti ecosystem revealed that lions killed at least 17 of 125 cheetah cubs born between 1987 and 1990.

Cheetahs avoid their competitors by using different temporal and habitat niches.

Leopards are able to take refuge in trees; lionesses, however, occasionally attempt to retrieve leopard kills from trees.

Lions similarly dominate African wild dogs, taking their kills and preying on young and rarely adult dogs. Population densities of wild dogs are low in areas where lions are more abundant.

However, there are a few reported cases of old and wounded lions falling prey to wild dogs.

Lions also charge at Nile crocodiles; depending on the size of the crocodile and the lion, either can lose kills or carrion to the other.

Lions have been observed killing crocodiles that ventured onto land. Lions also enter waterways, evidenced by the occasional lion claw found in crocodile stomachs.


Reproduce and life cycle



Most lionesses reproduce by the time they are four years of age.

Lions do not mate at a specific time of year and the females are polyestrous.

Like those of other cats, the male lion's penis has spines that point backward. During withdrawal of the penis, the spines rake the walls of the female's vagina, which may cause ovulation.

A lioness may mate with more than one male when she is in heat.

Generation length of the lion is about seven years.

The average gestation period is around 110 days;the female gives birth to a litter of between one and four cubs in a secluded den, which may be a thicket, a reed-bed, a cave, or some other sheltered area, usually away from the pride. She will often hunt alone while the cubs are still helpless, staying relatively close to the den.

Lion cubs are born blind – their eyes open around seven days after birth. They weigh 1.2–2.1 kg (2.6–4.6 lb) at birth and are almost helpless, beginning to crawl a day or two after birth and walking around three weeks of age.

To avoid a buildup of scent attracting the attention of predators, the lioness moves her cubs to a new den site several times a month, carrying them one-by-one by the nape of the neck.

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, particularly if other lionesses have given birth at about the same time.

When first introduced to the rest of the pride, lion cubs lack confidence when confronted with adults other than their mother.

They soon begin to immerse themselves in the pride life, however, playing among themselves or attempting to initiate play with the adults.

Lionesses with cubs of their own are more likely to be tolerant of another lioness's cubs than lionesses without cubs. Male tolerance of the cubs varies – sometimes a male will patiently let the cubs play with his tail or his mane, whereas another may snarl and bat the cubs away.


Pride lionesses often synchronise their reproductive cycles and communal rearing and suckling of the young, which suckle indiscriminately from any or all of the nursing females in the pride.

The synchronisation of births is advantageous because the cubs grow to being roughly the same size and have an equal chance of survival, and sucklings are not dominated by older cubs.

Weaning occurs after six or seven months. Male lions reach maturity at about three years of age and at four to five years are capable of challenging and displacing adult males associated with another pride. They begin to age and weaken at between 10 and 15 years of age at the latest.

When one or more new males oust the previous males associated with a pride, the victors often kill any existing young cubs, perhaps because females do not become fertile and receptive until their cubs mature or die.

Females often fiercely defend their cubs from a usurping male but are rarely successful unless a group of three or four mothers within a pride join forces against the male.

Cubs also die from starvation and abandonment, and predation by leopards, hyenas and wild dogs.Up to 80% of lion cubs will die before the age of two.

Both male and female lions may be ousted from prides to become nomads, although most females usually remain with their birth pride. When a pride becomes too large, however, the youngest generation of female cubs may be forced to leave to find their own territory.

When a new male lion takes over a pride, adolescent lions – both male and female – may be evicted. Lions of both sexes may be involved in group homosexual and courtship activities; males will also head-rub and roll around with each other before simulating sex together.


Health


Although adult lions have no natural predators, evidence suggests most die violently from attacks by humans or other lions.

Lions often inflict serious injuries on members of other prides they encounter in territorial disputes or members of the home pride when fighting at a kill.

Crippled lions and cubs may fall victim to hyenas and leopards or be trampled by buffalo or elephants. Careless lions may be maimed when hunting prey.

Ticks commonly infest the ears, neck and groin regions of lions.

Adult forms of several species of the tapeworm genus Taenia have been isolated from lion intestines, having been ingested as larvae in antelope meat.

Lions in the Ngorongoro Crater were afflicted by an outbreak of stable fly (Stomoxys calcitrans) in 1962; this resulted in lions becoming emaciated and covered in bloody, bare patches.

Lions sought unsuccessfully to evade the biting flies by climbing trees or crawling into hyena burrows; many died or migrated and the local population dropped from 70 to 15 individuals.

A more recent outbreak in 2001 killed six lions.

Captive lions have been infected with canine distemper virus (CDV) since at least the mid 1970s.

CDV is spread by domestic dogs and other carnivores; a 1994 outbreak in Serengeti National Park resulted in many lions developing neurological symptoms such as seizures. During the outbreak, several lions died from pneumonia and encephalitis.

Feline immunodeficiency virus and lentivirus also affect captive lions.


Communication



When resting, lion socialisation occurs through a number of behaviours; the animal's expressive movements are highly developed.

The most common peaceful, tactile gestures are head rubbing and social licking, which have been compared with the role of allogrooming among primates.

Head rubbing – the nuzzling of the forehead, face and neck against another lion – appears to be a form of greeting and is seen often after an animal has been apart from others or after a fight or confrontation. Males tend to rub other males, while cubs and females rub females.

Social licking often occurs in tandem with head rubbing; it is generally mutual and the recipient appears to express pleasure.

The head and neck are the most common parts of the body licked; this behaviour may have arisen out of utility because lions cannot lick these areas themselves.

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; males will spray and scrape plots of ground and objects within the territory.

The lion's repertoire of vocalisations is large; variations in intensity and pitch appear to be central to communication. Most lion vocalisations are variations of growling, snarling, meowing and roaring.

Other sounds produced include purring, puffing, bleating and humming. Roaring is used to advertise its presence. Lions most often roar at night, a sound that can be heard from a distance of 8 kilometres (5.0 mi).

They tend to roar in a very characteristic manner starting with a few deep, long roars that subside into a series of shorter ones.

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