AppearanceBehind 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. Lion colouration 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 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.
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.
The size of adult lions varies across their range with those from the southern African populations in Rhodesia, Kalahari and Kruger Park averaging around 189.6 kg and 126.9 kg in males and females respectively compared to 174.9 kg and 119.5 kg of male and female lions from East Africa.
Reported body measurements in males are head-body lengths ranging from 170 to 250 cm, tail lengths of 90–105 cm. In females reported head-body lengths range from 140 to 175 cm, tail lengths of 70–100 cm, however, the frequently cited maximum head and body length of 250 cm 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 for a male shot near Mucsso, southern Angola in October 1973 and a weight of 313 kg for a male shot outside Hectorspruit in eastern Transvaal, South Africa in 1936. Another notably outsized male lion, which was shot near Mount Kenya, weighed in at 272 kg.
NamingIt was one of the species originally described by Linnaeus, who gave it the name "Felis leo", in his eighteenth-century work, "Systema Naturae".
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 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. 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.
DistributionIn Africa, lions can be found in savanna grasslands with scattered "Acacia" trees, which serve as shade; their habitat in India is a mixture of dry savanna forest and very dry deciduous scrub forest. 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.
StatusMost 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. Estimates of the African lion population range between 16,500 and 47,000 living in the wild in 2002–2004, 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. Habitat loss and conflicts with humans are considered the most significant threats to the species. 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.
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. 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. Another population in northwestern Africa is found in Waza National Park, where approximately 14–21 animals persist.
BehaviorLions 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 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. 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.
Lions have an array of facial expressions and body postures that serve as visual gestures. 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.
They most often roar at night; the sound, which can be heard from a distance of 8 kilometres, is used to advertise the animal's presence. Lions have the loudest roar of any big cat.
ReproductionMost 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; 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, the female giving birth to a litter of one to four cubs in a secluded den 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 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 so that they cooperate in the raising and suckling of the young, 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 among the younger cubs.
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 associated with a pride, the conqueror often kill any existing young cubs, 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.
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 associated with another pride. They begin to age and weaken between 10 and 15 years of age at the latest, if they have not already been critically injured while defending the pride. 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.
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. 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. Lions are shown to involved in group homosexual and courtship activities. Male Lions will also head rub and roll around with each other before having sex together.
FoodLions prefer to scavenge when the opportunity presents itself, and scavenged food provides more than 50% of their diets. They scavenge animals either dead from natural causes 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.
The lioness is the one that does most of 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, which can cause cerebral ischemia or asphyxia. The prey also may be killed by the lion enclosing the animal's mouth and nostrils in its jaws.
Lions 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, 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. They sneak up to the victim until they reach a distance of approximately 30 metres or less.
The prey consists mainly of medium-sized 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 such as kudu, hartebeest, gemsbok, and eland. 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. Giraffes and buffaloes are almost invulnerable to a solitary lion as well.
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. There is evidence that male lions are just as successful at hunting as females; they are solo hunters who ambush prey in small bush. 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.
EvolutionThe 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. 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, 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.
CulturalThe 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 and 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, as well as a symbol of bravery; it is featured in several fables of the sixth century BC Greek storyteller Aesop.
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