The lion (Panthera leo) is a large cat of the genus Panthera, currently found only in Sub-Saharan Africa and 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.
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 literature and films. 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. The English word lion is derived via Anglo-Norman liun from Latin leōnem (nominative: leō), which in turn was a borrowing from Ancient Greek λέων léōn.
The African lion is both the most sexually dimorphic (with physical differences) of the big cats and the largest of African carnivores. Adult lions can be over 8 feet long and weigh between 250 and 500 pounds. The lion is the only feline with a tufted tail. They also have a small spine on the very tip of their tail, formed by fused terminal caudal vertebrae.
Lions are the most social of all the cats and appear to enjoy physical contact when resting or relaxing, which they do for approximately 20 hours of every day. In stark contrast, they are highly antagonistic to each other when feeding and there is a distinct hierarchy when it comes to dinnertime privileges. Male lions associated with a pride seldom hunt, instead providing protection and security. Females do the "lion's share" of securing prey but the larger males always eat first. Cubs are the last to eat and competition can be so fierce that mortality approaches 80% for lions under two years old.
A lion's pride consists of a few adult males, related females, and cubs. Groups of female lions usually hunt together, preying mostly on medium-sized and large ungulates. The lion inhabits grasslands, savannahs, and shrublands. It is usually more diurnal than other wild cats, but when persecuted, it adapts to being active at night and at twilight.
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During the Neolithic period, the lion ranged throughout Africa and Eurasia, from Southeast Europe to India, but it has been reduced to fragmented populations in sub-Saharan Africa and one population in western India.
In the 19th and 20th centuries, several lion type specimens were described and proposed as subspecies, with about a dozen recognised as valid taxa until 2017. Between 2008 and 2016, IUCN Red List assessors used only two subspecific names: P. l. leo for African lion populations, and P. l. melanochaita for lion populations in East and Southern African regions.
Evolutionary History
P. fossilis was larger than the modern lion and lived in the Middle Pleistocene. P. spelaea, or the cave lion, lived in Eurasia and Beringia during the Late Pleistocene. It became extinct due to climate warming or human expansion latest by 11,900 years ago. Bone fragments excavated in European, North Asian, Canadian and Alaskan caves indicate that it ranged from Europe across Siberia into western Alaska. It likely derived from P. fossilis.
Estimates for the divergence time of the modern and cave lion lineages range from 529,000 to 392,000 years ago based on mutation rate per generation time of the modern lion. There is no evidence for gene flow between the two lineages, indicating that they did not share the same geographic area.
The modern lion was probably widely distributed in Africa during the Middle Pleistocene and started to diverge in sub-Saharan Africa during the Late Pleistocene. Lion populations in East and Southern Africa became separated from populations in West and North Africa when the equatorial rainforest expanded 183,500 to 81,800 years ago. They shared a common ancestor probably between 98,000 and 52,000 years ago.
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Due to the expansion of the Sahara between 83,100 and 26,600 years ago, lion populations in West and North Africa became separated. As the rainforest decreased and thus gave rise to more open habitats, lions moved from West to Central Africa. 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. A whole genome-wide sequence of lion samples showed that samples from West Africa shared alleles with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia.
Physical Characteristics
The lion is a muscular, broad-chested cat with a short, rounded head, a reduced neck, and round ears; males have broader heads. The 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 may still be seen on the legs and underparts.
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" composed of dermal papillae. The functions of the spur are unknown.
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.
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The male lion's mane is the most recognisable feature of the species. It may have evolved around 320,000-190,000 years ago. It grows downwards and backwards, covering most of the head, neck, shoulders, and chest. This feature likely evolved to signal the fitness of males to females. Males with darker manes appear to have greater reproductive success and are more likely to remain in a pride for longer. They have longer and thicker hair and higher testosterone levels, but they are also more vulnerable to heat stress.
Unlike in other felid species, female lions consistently interact with multiple males at once. Another hypothesis suggests that the mane also serves to protect the neck in fights, but this is disputed. During fights, including those involving maneless females and adolescents, the neck is not targeted as much as the face, back, and hindquarters.
The white lion is a rare morph with a genetic condition called leucism, which is caused by a double recessive allele. It is not albino; it has normal pigmentation in the eyes and skin. White lions have occasionally been encountered in and around Kruger National Park and the adjacent Timbavati Private Game Reserve in eastern South Africa. They were removed from the wild in the 1970s, thus decreasing the white lion gene pool.
A majestic lion waiting in Namibia.
Habitat and Distribution
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 rarely enters closed forests. On Mount Elgon, the lion has been recorded up to an elevation of 3,600 m (11,800 ft) and close to the snow line on Mount Kenya.
Savannahs with an annual rainfall of 300 to 1,500 mm (12 to 59 in) make up the majority of lion habitat in Africa, estimated at 3,390,821 km2 (1,309,203 sq mi) at most, but remnant populations are also present in tropical moist forests in West Africa and montane forests in East Africa.
The Asiatic lion now survives only in and around Gir National Park in Gujarat, western India. In Asia the lion once ranged in regions where climatic conditions supported an abundance of prey. It was present in the Caucasus until the 10th century. It lived in the Levant until the Middle Ages and in Southwest Asia until the late 19th century.
Social Behavior and Hunting
A lion pride in Kenya.
Lions spend much of their time resting; they are inactive for about twenty 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.
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. The majority of females remain in their birth prides while all males and some females disperse.
The average pride consists of around 15 lions, including several adult females and up to four males and their cubs of both sexes. Nomadic lions range widely and move around sporadically, either in pairs or alone. Pairs are more frequent among related males. A lion may switch lifestyles; nomads can become residents and vice versa.
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 patrol the fringes. 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. 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.
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 medium-sized to large ungulates, particularly blue wildebeest, plains zebra, African buffalo, gemsbok and giraffe. 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 exceptional cases, with the victims being calves, juveniles, and even subadults.
In typical group 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. Some evidence suggests, however, that males are just as successful as females; they are typically solo hunters who ambush prey in small bushland. They may join in the hunting of large, slower-moving prey like buffalo; and even hunt them on their own.
The lion's attack is short and powerful; it attempts to catch prey with a fast rush and final leap, usually pulls it down by the rump, and kills with a clamping bite to the throat or muzzle. 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).
Lion vs. Wildebeest: How Lions Hunt as a Pride
Skull Anatomy and Distinguishing Features
A lion’s skull is made up of thick, strong bones that are heavy, which is important as they protect the brain. Lions have 30 permanent teeth that are specially designed for hunting prey and eating their meet. These include:
- Canines - long sharp teeth at either side of the incisors.
- Incisors
- Carnassial teeth
The lion will use the canines first, then the incisors and then the carnassial teeth.
The dentary bone is a strong bone that has lots of muscles attached to it which helps it to move. The jaw is crucial for eating, breathing and hunting. A lion’s jaw can open up to 28cm wide but cannot move side to side, which helps to keep the teeth in the correct position for cutting.
The muscles that help a lion to open and close its mouth are powerful and there are ridges and valleys on the skull’s surface which are formed by the movement of these muscles. A lion’s sense of smell is well developed and they possess a special organ, called the Jacobson’s Organ, in the roof of the mouth which allows them to locate kills made by other animals and also to find their own.
Skull Morphology and Identification
While lions and tigers are osteologically very similar, they are not identical.
For the purposes of use by customs officials, however, identifying character traits should ideally be user-friendly, easily distinguished at a glance, should not require equipment and complex measurements of character length and ratios, and also be semantically accessible to a person with no expertise in comparative anatomy.
Only two characters met our requirement of being a practical identification tool with an acceptable degree of certainty, namely:
- The position and alignment of the posterior projections of the nasal-frontal and maxilla-frontal sutures on the cranium
- The ventral profile of the horizontal ramus of the mandible, including the number of contact points the mandibles/jaws have on a flat surface.
The mean skull mass for wild-origin African Lions was 1.3 ± 0.4kg, with skulls ranging in weight from 0.7kg for a lioness specimen collected in South Africa in 1906 (DNHM 385), to 2.7kg for a 7.5 year old male that died in Hwange in May 2013. Known female skulls weighed less than 2.0kg, whereas male skulls will be heavier.
Skull mass can be used as a proxy for lion size and age, and heavier skulls from individuals of unknown sex and age are inferred to have originated from bigger lions and, typically, older males.
Conservation Status and Threats
Conservation efforts are crucial for the survival of African lions.
Lions are a vulnerable species of mammals found in the savannah and grasslands of Africa and in India. In the past lions also lived in Europe and Asia but because of habitat destruction and hunting they are now only found in Africa and India.
They are classed as a vulnerable species as there are less than 20,000 in the wild. Few male lions live beyond ten years in the wild; females may live up to 15 years.
Illegal Trade in Lion Bones
South Africa has legally exported substantial quantities of lion bones to Southeast Asia and China since 2008, apparently as part of the multinational trade substituting bones and body parts of other large cats for those of the tiger in wine and other health tonics. The legal sale of lion bones may mask an illegal trade, the size of which is only partially known.
An observed component of the illegal trade is that quantities of skeletons are sometimes declared falsely/fraudulently on CITES export permits. Furthermore, there are emerging concerns that bones from tigers reared in captivity in South Africa and elsewhere are being laundered as lion bones using CITES Appendix II permits.
In light of this need for a new set of tools to cope with the now established trade in lion bones and concerns that tiger bones are being laundered in South Africa, the aim of this paper is to provide a guide to the average mass of lion skeletons and skulls, and a means for distinguishing the skulls of lions from tigers.
Monitoring and Identification Techniques
Our research indicates that it is possible to use skeletons, skulls and cranial sutures to detect misdeclarations in the lion bone trade. It is also possible to use the average mass of a lion skeleton to corroborate the numbers of skeletons declared on CITES permits, relative to the weight of the consolidated consignments stated on the air waybills.
