Lion
Panthera leo
M. Pittet
Description
Previously, the eleven former lion subspecies described were reclassified into two subspecies: the African lion (Panthera leo leo) and the Asiatic lion (Panthera leo persica). However, more recent studies indicate that lions from Asia and West and Central Africa are more closely related to each other than to lions from Eastern and Southern Africa. Based on these recent genetic studies, two subspecies of the lion are recognised:
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Panthera leo leo in Central and West Africa and India, formerly throughout North Africa, South-East Europe, the Middle East, Arabian Peninsula and South-West Asia
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Panthera leo melanochaita in Southern and Eastern Africa.
These two main divisions of lions are not homogeneous as there is genetic subdivision within each, with more genetic variation and deeper divergences within the Eastern and Southern branch than within the Asian and West and Central African one. In North-Eastern Africa, notably in Ethiopia, there is an overlap zone, where both subspecies exist.
The lion is very muscular and its body is designed for strength and short bursts of high speed. The shoulders and forepaws are particularly massive and powerful enabling lions to tackle and subdue large prey. The lion has a uniformly tawny coat, which blends into the grass cover of the savanna. There is some local variation in coat colour from pale yellowish or tawny to light grey or dark brown. Leucism, which is white fur but with pigmented eyes and skin, has been reported only from the vicinity of Timbavati in the Kruger National Park, South Africa. Melanistic lions have not been recorded. Male lions have tufted tails and manes, which vary from being quite sparsely furred in hot and humid areas to very impressive manes in milder climates. The mane's function is to attract females for mating, serving as an indicator of individual fitness. Young lions have pink noses that become progressively speckled with black spots as they mature. By five or six years of age, about sixty percent of the nose is black, becoming solidly black in some older individuals. This change in nose colour typically coincides with male lion's manes becoming darker and more complete at about five to six years of age. Lions in Africa have shorter fur and a more pronounced tail tassel than lions in Asia. The lions in Asia have a smaller, shorter mane than the lions in Africa and a more pronounced belly (lingual) fold of skin and a more pronounced hair tuft at the elbow. The belly fold is also seen in lions from West Africa.
Weight
110 - 272 kg
Body Length
137 - 250 cm
Tail Length
60 - 100 cm
Longevity
12 - 18 years
Litter Size
1 - 4 cubs
P. Meier
Status and Distribution
The lion is classified as Vulnerable in the IUCN Red List due to a population reduction in the past of over 30% based on direct observation, index of abundance, decline in habitat and exploitation. In some areas the lion shows stable populations or is even recovering but in others it is declining or even facing local extirpation. The lion in Africa is extinct in Algeria, Burundi, Congo, Djibouti, Egypt, Eritrea, Eswatini, Gambia, Lesotho, Libya, Mauritania, Morocco, Sierra Leone, Tunisia, and the Western Sahara. Furthermore, it is possibly extinct in Côte d'Ivoire, Ghana, Guinea, Guinea-Bissau, Mali, and Togo. In Asia, it only remains in India in a single isolated subpopulation in the 1,400 km² Gir Forest National Park and Wildlife Sanctuary and the surrounding landscape. There are indications of a general population decline across the lion’s range and the global population is highly fragmented. This is especially true in West Africa where all seven subpopulations are considered to be isolated from one another. In Central and East Africa, various populations are also fragmented. In 2023 the extant range of the lion was estimated at 6% of its historical range. This represents an overall decline of 36% in the species’ range over the time span from 2002 to 2023, resulting in a suspected population reduction in the same magnitude. In South Africa, the lion is listed as Least Concern but as Critically Endangered in West Africa. In parts of the lion's range, its current status is not well known, with key knowledge gaps in Angola, Central African Republic, Democratic Republic of Congo, Nigeria, Somalia, and South Sudan. Populations there are assumed to be rapidly declining, mainly due to armed conflicts.
In Southern Africa lions partly occur in fenced reserves and the lion populations in Southern Africa are increasing or stable. In West Africa, lions are largely confined to small populations numbering less than 50 individuals in Niokolo-Koba, and Yankari and Kainji Lake National Parks (Nigeria) with the largest population (estimated 187 lions) in the W-Arly-Penjari Complex (Benin, Burkina Faso and Niger). Throughout most of Africa, lions are becoming increasingly rare outside protected areas and a lot of lions live in isolated and small populations. The situation of the African lion is most alarming in West and Central Africa.
Lions are difficult to survey, and knowledge of lion numbers and distribution is limited. Especially estimating lion populations is a challenging and difficult task. In Africa, lion density can vary from 1.5 to 55 per 100 km². In the early 2000's, several initiatives, based on a compilation of recent estimates from scientific surveys, expert opinions and best guesses, estimated the total number of wild lions in Africa at 22,800 to 39,000 individuals. In 2013, the total number of lions in Africa was estimated at around 35,000 individuals, based on reviewing all available data and combining it with satellite imagery. In 2023 the lion population in Africa was estimated at approximately between 22,000 and 25,000 adult and subadult lions.
In 1880 the lion population in the Gir forests, India, was only estimated at around 12 animals. From 1968 onwards the lion population increased steadily and was estimated at ~670 adult and subadult lions in 2020. The population is now stable. In the Gir forest, lion density varies between management zones and depending on terrain, habitat, and human influences. Maximum lion density was estimated in the East of the Gir with 16 animals / 100 km². In the West and Central Gir, lion density was estimated at 12 and 6 individuals / 100 km² respectively.
Habitat
The African lion occurs south of the Sahara. It has a high habitat tolerance and inhabits different environments such as forested areas, dry forests, scrubs, and deserts, but is largely a species of the savannas. Lions have been recorded at elevations of up to 4,240 m in the Bale Mountains in Ethiopia and up to 3,600 m on Mount Elgon in Kenya. The lion has a preference for open areas such as scrub and grass complexes, and open and closed woodland, and requires areas with sufficient shelter and cover for hunting and denning, sufficient suitable prey, and minimal disturbance from humans.
In Asia, the lion is found predominantly in forested habitats. It shows a preference for moist mixed forests, followed by mixed forests, savannah habitats, and Teak-Acacia-Zizyphus dominated woodlands. During the day, lions in Asia seem to prefer densely vegetated areas but at night their movement is more widespread, and they may even move through agricultural fields and human habitations. Dense vegetation within forested areas thus allows lions to escape from the heat of the day and also provide cover at a time when human activity is likely to be high.
The Gir Forest, India, is a dry deciduous forest dominated by teak (Tectona grandis) in the west, and in the drier parts in the east predominantly vegetated with acacia thorn savannah. The Gir forest is surrounded by cultivated landscape.
M. Pittet
Threats
The main threats to lions include continued habitat loss, retaliatory or indiscriminate killing (using spears, guns, snares and poisoning), prey base depletion, bushmeat poaching, unsustainable or poorly monitored trophy hunting, targeted poaching, and violent extremism/warfare. Another major threat is human-lion conflict due to livestock predation and rare attacks on humans, resulting in indiscriminate killing of lions. This threat has increased dramatically in conjunction with increasing human populations and lions are persecuted intensely in livestock areas across Africa. Especially in India, the close proximity of predators, livestock and humans with the increasing expansion of settlements and agricultural areas outside protected areas give rise to a number of management problems which threaten the survival of the lion. In India, over 20% of the lion population now survives outside the boundaries of the protected area and move throughout the Greater Gir Landscape, causing higher livestock depredation and subsequent conflicts. Generally, increasing human populations steadily extirpate lions out of many areas lacking formal protection. It is not known how many lions are killed as problem animals by local people, although this is the major threat to lions outside of protected areas. Lions are very vulnerable to being killed in wire snares and leghold traps set for large ungulates. In some areas, this has the greatest impact on lion populations, and often occurs well within the boundaries of large protected areas. In India, until now the lions have been tolerated by local communities and cultural tolerance within the agro-pastoral economy appears to be conducive to lion conservation in the region. The management of conflict between lions and people has important implications for the conservation of the lion in India.
Prey species are increasingly under threat due to unsustainable and raising commercialized bushmeat trade. The bushmeat trade is an underestimated contributor to lion population declines and vast areas of suitable habitat are now completely devoid of prey, with lions from those areas either having been caught in snares, which are widespread over much of the lion’s range, or killed when resorting to hunting livestock. Herbivore populations have increased by 24% in Southern Africa but have declined by 52% in Eastern and by 85% in West Africa.
Trophy hunting is taking place in various countries in sub-Saharan Africa. Such hunting can be considered an important conservation tool providing it is done sustainably, and that money is directed to the communities that live alongside lions. However, in all the key lion hunting countries in Africa, this is not the case: communities see very little benefit from coexisting with lions and benefits from lion hunts are not directed to the communities. Offtake is largely limited by age-based regulations, the theory being that hunting lions only above the age of six years should be sustainable. Thus, lions continue to be unsustainably killed both legally for trophies and illegally for retribution, cultural purposes, and in traps set for their prey.
At times, diseases such as rabies or canine distemper virus can also be a problem in some parts of their range, but generally diseases do not largely affect lion populations. The small size of many lion populations’ makes them particularly vulnerable to all of the threats listed above and as lions scavenge frequently, they are very vulnerable to poisoned carcasses.
Another threat to lions is use of their bones and body parts which as increased in recent years and emerged in several range countries. As wild living tiger populations are declining, the illegal trade for bones and body parts is shifting to lions as an alternative. Illegal trade in lion body parts for traditional medicinal purposes is a threat not only to the lions in India but also to African lion subpopulations.
Impact of violent extremism in several parts of lion ranges in Africa is likely to further accelerate the decline in lions in these regions. Especially the W-Arly-Pendjari complex, the last stronghold for lions in West and Central Africa, is affected by extremist groups, threatening the remaining lion population. In West Africa lions are largely restricted to protected areas, which have weak management leading to the decrease of prey populations and the persecution of lions. Further deterioration of the last protected areas containing lions in West Africa can lead to their regional extinction.
Climate change will likely have a negative impact on the species although its extent is unknown.
The lions in Asia, which are restricted to a relatively small population in a single site, face a variety of extinction threats, both genetic and environmental. This small population of lions is highly vulnerable to disease outbreaks, poaching, and environmental disasters. The isolation of the population can lead to inbreeding due to a reduction of genetic variation.
The above threats are typically underpinned by inadequate management resources to adequately safeguard protected populations. However, the ultimate factor influencing lion populations is the increasing human population, which leads to all of the above, eventually resulting in habitat fragmentation, and suppressed and small lion populations exposed to continuous human pressure. In the meantime, a lot of research is being done to find strategies where lions can co-exist with humans, even if this is only around the borders of larger protected areas. However, some scientists increasingly believe that the future for lions in Africa exists mainly in very large protected areas, and that smaller reserves will only be able to conserve lions if they are adequately fenced off from human activities.
P. Meier
Historically, lions were widely distributed and occurred in Africa, Europe, the Middle East and in south-west Asia. Lions disappeared from Europe during the first century CE and from North Africa, the Middle East, and most parts of Asia between 1800 and 1950. In North Africa the lion probably survived in the High Atlas Mountains up to the 1940s. In Asia, it previously inhabited most of south-west Asia ranging from Syria through Iraq, Iran, Pakistan, and most of Northern and central India to parts of eastern India. The lion became extinct in Eastern Europe at around 100 CE and in Palestine around the time of the Crusades. In all other areas, the lion in Asia remained widespread until the mid-1800s. The invention of firearms and widespread indiscriminate hunting led to its extinction over large areas. In Syria, the last report of lions dates back to 1891. By the late 1800s the lion had disappeared from Turkey while in Iran and Iraq it was last recorded in 1942 and 1918 respectively. In India, lions ranged east to the state of Bihar but they declined rapidly due to heavy hunting pressure and habitat loss. By the turn of the 19th century the lion in Asia remained only in the Gir Forest in Gujarat, India. Presently, lions are also found outside the Gir Protected Area such as in forests of Girnar, Mitiyala, coastal areas of Sutrapada-Kodinar, Jafarabad-Rajula up to the Savarkundla and Palitatna hills covering an area of around 10,400 km².
Ecology and Behaviour
The lion is mainly nocturnal and crepuscular killing over 95% of its prey at night. Lions spend large parts of the day and night resting and are typically active for four to six hours per day. The lion is the most social of all cat species, living in complex social systems called prides and exhibiting group territorial behaviour. Females engage in several cooperative behaviours unique among felids. Pride members often give birth in synchrony and the young are reared communally. There seems to be a complex division of labour among hunting lionesses. The pride structure varies between lions in Africa and lions in Asia and may even differ locally depending on climate and prey base conditions. In the Gir forest, most prides consist of only two females. The average size of lion prides in Africa is four to six adult lionesses but can range from 1-21. This is the core and most stable unit in the pride. Prides are typically defended by coalitions of two males, (though this can range from 1-9), and typically have a number of dependent cubs or subadults. Thus, prides typically average about 12-16 individuals, however prides of up to 30 individuals have been observed in larger ecosystems. In areas with high human hunting pressure and reduced prey base, pride sizes tend to be smaller, cub survival lower, and pride persistence is less secure.
Related or unrelated males form coalitions to compete for tenure in a pride, which typically lasts for one breeding cycle, about 24-36 months. A single male or a male coalition holds tenure over one or more prides. Single males seldom maintain tenure over prides except in low-density lion populations, as competition for pride tenure is very high. Only very large coalitions, typically 3-5 individuals, tend to regularly have tenure over multiple prides, although paired coalitions often take over tenure of an adjacent pride once the cubs in their current pride are over 18 months old. When a pride is taken over by new males, infanticide is common if there are young cubs (less than 12 months old). Typically, then, most females lose their cubs within a period of about a month. Thereafter, lionesses come into oestrous quite soon, and the new males can reproduce earlier.
The size of the pride and its territory is strongly influenced by prey base. In high-density lion populations, density may range from 10-40 lions per 100 km², with territories ranging from 50-200 km². Such high densities are only possible in areas of high prey biomass, typically with resident prey and no limitation of other key resources. Moderate density lion populations, ranging from 4 to 10 lions/100 km² occur in areas of moderate prey biomass, which often include nutrient poor woodlands with limited water resources and thus support lower prey densities. In these areas pride ranges typically cover 200-500 km². When prey is far less abundant, typically limited by low rainfall or very poor soils, low density lion populations occur at densities of 1-3 lions per 100 km², with home ranges increasing exponentially in size from 500 to 5000 km². Lions occupying larger home ranges than this are probably not resident and may wander widely before settling down in a territory.
In the Gir forest, home ranges of male lions vary between 11 to 174 km² and the home ranges of females between 26 and 43 km². Intensively used core areas, are much smaller with around 10 km² for males and 5 km² for females. A radio-telemetry study in the Gir forest estimated an average home range of 85 ± 54 km² for males and a home range of 48.2 ± 10.6 km² and 35 ± 7 km² for females. The average dispersal distance of Asiatic lions was estimated at 26 km.
Lions are territorial and both sexes defend the territory against same sex intruders. While male lions holding a territory scent mark and roar very frequently to demarcate and announce their presence, lionesses seldom do so when they have young cubs, other than in the presence of the pride male. Lionesses start to scent mark more frequently when their cubs are about 30 months old or have dispersed. The lions roar can be heard by other lions 5-10 kilometres away and serves to warn other males not to enter their territory, to assure their pride that they are still defending the territory, and to allow subadult lions to avoid them.
Lions drink regularly when water is available but are also capable of obtaining most of their moisture requirements from their prey and even from plants. This allows lions to survive in very dry environments and go for very long periods without drinking water. However, waterpoints play a pivotal role in the movements and hunting behaviour of lions in most environments. Lions thus spend a large amount of time close to water and in many environments make most of their kills within about two kilometres of a waterhole or riverbank.
Lions are not seasonal breeders having cubs at any time of the year. However, in Asia, mating peaks during winter based on sightings of cubs, there seems to be a birthing peak from late winter to early summer. In some ecosystems, cub survival can be influenced by either seasonal or cyclical climatic events. When conditions are suitable, lion populations can respond quickly to excess prey, and prides can increase quickly in size, typically also shrinking the home range, allowing new prides to establish. The onset of spermatogenesis begins at 30 months, but, although sexually mature, few male lions can hold a territory, and thus breed successfully, before about five years of age. Females may begin mating at 24 months, but successful reproduction generally occurs when lionesses are older than three years of age. Typically, oestrus lasts for 4 days, and the inter-oestrus interval may be as short as 16 days. The gestation period lasts 100-119 days. The mortality of cubs is rather high in populations that experience extended periods of prey scarcity but is quite low in populations with moderate to high resident prey densities. Cub survival is higher in prides with three to seven lionesses. In prides with fewer lionesses, territorial skirmishes often result in cub mortality, and in large prides, intra-pride competition often results in the death or starvation of cubs. Infanticide can be quite high, causing up to 60% of cub mortality in lions in Asia and seems to occur mostly in the first year after birth.
The interbirth interval varies largely according to survival of cubs and the environment. When cubs survive, the interbirth interval is typically about two to three years, with longer interbirth intervals in ecosystems with relatively high resident prey biomass and thus high lion densities. Males generally leave their natal pride of their own volition at 3-4 years, but young males may be forced out much earlier during a pride takeover. Male lions that disperse from their natal pride at less than 30 months of age seldom survive, often being killed in retaliation for livestock depredation. Most young females are incorporated into their natal prides, but about 30% disperse at approximately 2.5 years. Females can reproduce up to the age of about 12-13 years. Although male lions of up to 16 years old can still produce viable sperm, very few males above 10-11 years old can maintain a territory, often dying quite soon after finally becoming solitary without a pride or coalition partner.
K. Varma
Prey
The lion is a generalist hunter and will take almost any animal, but they have a marked preference for medium to large sized ungulates (100-300 kg) such as wildebeest, zebra, buffalo, giraffe, kudu, and waterbuck. They also catch smaller antelopes such as gazelles and impala quite frequently, and only prey on the lost or weakened young, or severely weakened adults, of very large prey such as rhinos, hippos, and elephants. Although they will go for long periods without food and ignore easy prey, at times lions also kill unattended livestock, and very rarely, humans. Lions also frequently steal the kills made by other carnivores and will scavenge from any carcass they find. Lions hunt mostly at night and in groups but have a relatively low hunting success rate with about 30% of attempts being successful. Although it is widely believed that lionesses do most of the hunting, when on their own, male lions are efficient hunters typically tackling large prey such as buffalo and giraffe. Male lions have also been found to prey successfully on warthogs by digging them out of their burrows, and on impala, particularly when hunting in thick bush. Lionesses prefer hunting medium-large ungulates such as wildebeest and zebra in more open areas where their co-ordinated hunting strategy is most effective.
In Asia, the lion preys mainly on sambar deer (Rusa unicolor), chital (Axis axis), chosingha (Tetracerus quadricornis), chinkara (Gazella bennetii), wild boar (Sus scrofa), and nilgai (Boselaphus tragocamelus). They also prey on porcupine (Hystrix indica), common langur (Semnopithecus entellus), rufus tailed hare (Lepus nigricollis ruficaudata), and peafowl (Pavo cristatus) and take livestock. Nilgai and wild pig are common wild prey species found outside of the Gir forest protected area. The proportion of livestock taken by lions in Asia seems to be influenced by the abundance of wild prey and varies seasonally. Significantly more wild animals are taken during summer when prey are concentrated around water sources. Within protected areas, lions predominantly consume wild prey in proportion to their availability. Outside protected areas, however, livestock seems to be the major prey and lions seem to depend considerably on domestic animals.
P. Meier
P. Meier
Conservation Efforts and Protection Status
The lion is protected under Appendix II of CITES. The lion in Asia is included under Appendix I of CITES and fully protected in India under Schedule I of the Wildlife Protection Act (1972). The IUCN SSC Cat Specialist Group’s African Lion Database is used to provide a continually updated assessment of the abundance and distribution of lions in Africa.
In Sub-Saharan Africa, the lion conservation community works in four broad regions: West, Central, Eastern, and Southern. Several workshops for the conservation of lions have been conducted throughout Africa. Regional Conservation strategies have been developed for West and Central as well as for Eastern and Southern Africa. These need to be reviewed and updated. The African lion conservation strategy in West and Central Africa has three main objectives:
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to reduce lion-human conflict
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to conserve habitat and wild prey
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to increase lion habitat and wild prey
The lion conservation strategy of Eastern and Southern Africa additionally includes policy and land use aspects as well as socio-economic issues, trade, and conservation politics. Both strategies have set priorities for lion conservation including the improvement of lion management and the creation of incentive through benefits for communities living alongside lions. These regional strategies help guide Action plans and conservation projects for lions and have the potential for broad and significant improvement of the lion status.
Key conservation efforts in Africa are effective protected area management including funding, mitigating anthropogenic causes of mortality such as lion-human conflict and snaring, and reducing or mitigating negative effects of armed conflicts. Human-lion conflict can be minimized through appropriate livestock management measures which include measures for problem animal control and mechanisms for compensating livestock losses. Urgent conservation action is especially needed for the lion in West Africa, where it is Critically Endangered and declining within the region, to prevent local extinctions of this subspecies within the region.
In India, the lion population has been steadily increasing in response to successful management and conservation initiatives spanning over the last five decades. Management interventions, such as reduction in livestock grazing and control of fire, have led to vegetation recovery and increase in wild ungulate populations. The lion population is now expanding and moving throughout the Greater Gir Landscape which is of outstanding importance for the long-term conservation of this species. The lions in India live in a landscape where people have a high tolerance and consider lions as part of their natural heritage. However, to maintain this positive attitude of local communities in the long-term, husbandry practices and economic incentives as well as human safety must be improved. Of concern is the human-lion conflict mainly outside the protected area, where human and livestock density are higher. To foster coexistence of humans and lions, conflict mitigation and the promotion of positive public perceptions of lions is essential for the continuation of successful Asiatic lion conservation. It may also be necessary to create corridors to enable free movement of lions between reserves or isolated forest patches. In order to maximize the genetic diversity and reduce the risk of extinction through an epidemic outbreak, there is a need to establish at least one more wild population of lions in Asia.
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P. Meier
African Lion Database. Unpublished Data. 2023 2024. Panthera leo. The IUCN Red List of Threatened Species. Version 2024-2