Panthera leo leo is a lion subspecies present in West Africa, northern Central Africa and India. In West and Central Africa it is restricted to fragmented and isolated populations with a declining trajectory. It has been referred to as the northern lion.
Results of a phylogeographic study indicate that lion populations in West and Central African range countries are genetically close to populations in India, forming a major clade distinct from lion populations in Southern and East Africa. In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group subsumed lion populations according to the major clades into two subspecies, namely P. l. leo and P. l. melanochaita. Within P. l. leo three subclades are clearly distinguishable. One from Asia, which includes the extinct Barbary lions of North Africa, another one from West Africa and a third one from Central Africa, north of the rainforest belt.
P. l. leo is regionally extinct in North Africa, southern Europe, and West Asia. Asia's sole lion population lives in and around Gir National Park, India. The West African lion population is geographically isolated and numbers fewer than 250 mature individuals. It is listed as critically endangered on the IUCN Red List.
Taxonomy A lion from Constantine, Algeria, was the type specimen for the specific name Felis leo used by Carl Linnaeus in 1758. In the 19th and 20th centuries, several lion zoological specimens from Africa and Asia were described and proposed as subspecies: Felis leo persicus described in 1826 by Johann N. Meyer was a lion skin from Persia. Felis leo senegalensis also described by Meyer in 1826, but based on a lion skin from Senegal. Felis leo nubicus described in 1843 by Henri Marie Ducrotay de Blainville was a male lion from Nubia that had been sent by Antoine Clot from Cairo to Paris and died in the Ménagerie du Jardin des Plantes in 1841. Leo gambianus described in 1843 by John Edward Gray was a specimen from the Gambia in the collection of the British Museum of Natural History. Felis leo kamptzi described in 1900 by Paul Matschie was a lion skull from northern Cameroon. Leo leo azandicus described in 1924 by Joel Asaph Allen was a male lion that was killed in 1912 in northeastern Belgian Congo as part of a zoological collection comprising 588 carnivore specimens. Allen admitted a close relationship of this lion specimen to Leo leo massaicus from Kenya regarding cranial and dental characteristics, but argued that his type specimen differed in pelage colouration. In 1930, Reginald Innes Pocock subordinated the lion to the genus Panthera when he wrote about Asiatic lion specimens in the zoological collection of the British Museum of Natural History.
In the following decades, there has been much debate among zoologists on the validity of proposed subspecies: In 1939, Glover Morrill Allen recognized Felis leo kamptzi and F. l. azandicus as valid taxa among ten lion subspecies. Three decades later, John Ellerman and Terence Morrison-Scott recognized only two lion subspecies in the Palearctic realm, namely the African (P. l. leo) and Asiatic lions (P. l. persica). Some authors considered P. l. nubicus a valid subspecies and synonymous with P. l. massaica, a specimen from Kenya. Some authors considered P. l. azandicus synonymous with P. l. massaicus and P. l. somaliensis, and P. l. kamptzi synonymous with P. l. senegalensis. In 2005, Wallace Christopher Wozencraft recognized P. l. kamptzi, P. l. bleyenberghi and P. l. azandica as valid taxa. In 2016, IUCN Red List assessors used P. l. leo for all African lion populations. In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group subsumed lion populations in North, West and Central Africa and Asia to P. l. leo, based on results of genetic research on lion samples.
Phylogeny
Since the beginning of the 21st century, several phylogenetic studies were conducted to aid clarifying the taxonomic status of lion samples kept in museums and collected in the wild. Scientists analysed between 32 and 480 lion samples from up to 22 countries. They all agree that the lion comprises two evolutionary groups, one in the northern and eastern parts of its historical range, and the other in Southern and East Africa; they are estimated to have genetically diverged between 245,000 and 50,000 years ago. Tropical rainforest and the East African Rift possibly constituted major barriers between the two groups.
The two lion groups overlap in Ethiopia, as lion samples from Bale Mountains National Park clustered with lion samples from Central Africa, whereas other samples from this country clustered with samples from East Africa. Three clades can be distinguished within P. l. leo. Lion samples from North Africa and India clustered into a single clade, and the lions in West Africa and northern parts of Central Africa also form distinct clades. Analysis of phylogenetic data of 194 lion samples from 22 countries revealed that Central and West African lions diverged about 186,000–128,000 years ago from the melanochaita group in East and Southern Africa.
Samples from West Africa shared alleles with samples from Southern Africa, and samples from Central Africa shared alleles with samples from Asia. This indicates that Central Africa was a melting pot of lion populations after they had become isolated. They possibly migrated through corridors in the Nile Basin during the early Holocene. Genome-wide data of a historical lion sample from Sudan showed that it clustered with P. l. leo in mitochondrial DNA-based phylogenies, but with a high affinity with P. l. melanochaita. The taxonomic position of lions in Central Africa may therefore require revision.
Characteristics
The lion's fur varies in colour from light buff to dark brown. It has rounded ears and a black tail tuft. Average head-to-body length of male lions is with a weight of . Females are smaller and less heavy. Zoological lion specimens range in colour from light to dark tawny. Male skins have short manes, light manes, dark manes or long manes. Taxonomists recognised that neither skin nor mane colour and length of lions can be adduced as distinct subspecific characteristics. Then they turned to measuring and comparing lion skulls and found that skull length of Barbary and Indian lion samples does not differ significantly, ranging from in females and in males. A few lion specimens from West Africa obtained by museums were described as having shorter manes than lions from other African regions. In general, the West African lion is similar in general appearance and size as lions in other parts of Africa and Asia. Skeletal muscles make up 58.8% of the lion's body weight.
Distribution and habitat
Today, P. l. leo occurs in West and Central Africa and India. It is regionally extinct in The Gambia, Mauritania, Sierra Leone, the Western Sahara, Morocco, Algeria, Tunisia, Libya, Egypt, Saudi Arabia, Kuwait, Jordan, Lebanon, Syria, Turkey, Palestine, Israel, Iraq, Iran and Pakistan. During the Holocene, Asiatic lions colonised southeast Europe and adjacent areas of the continent, spanning from Hungary to Greece to Ukraine, but became extinct likely in classical times. In 2005, a Lion Conservation Strategy was developed for West and Central Africa. Contemporary lion distribution and habitat quality in savannahs of West and Central Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped. Educated guesses for size of populations in these LCUs ranged from 3,274 to 3,909 individuals between 2002 and 2012.
West African clade
The last populations of the West African lion clade are surviving in a few protected areas from Senegal in the west to Nigeria in the east. This population has lost 99% of its former range. Between 246 and 466 lions live in the WAP-Complex, a large system of protected areas formed mainly by W, Arli, and Pendjari National Parks in Burkina Faso, Benin, and Niger. It is regionally extinct in Mauritania, Gambia, Guinea-Bissau, Mali, Sierra Leone, Ivory Coast, and Togo, and possibly extinct in Guinea and Ghana. The border between the West African and the Central African lion clade is following largely the lower Niger River, which seems to act as a permanent barrier for gene flow. Range countries Lion Conservation Units Area in km2 Senegal, Mali, Guinea-Bissau, Guinea Niokolo-Koba National Park 90,384Guinea National Park of Upper Niger 613Benin, Burkina Faso, Niger W-Arly-Pendjari Complex 29,403Benin three unprotected areas 6,833Nigeria Kainji National Park 5,340
Central African clade
The Central African lion population inhabits protected areas of: Cameroon, where lions are present in Bénoué National Park. In the North Province, Cameroon, lions were recorded during a survey between January 2008 and May 2010. The small lion population in Waza National Park is isolated, and by 2008 had declined to maximum 20 individuals. In the southern part of the country, 2 lions were discovered in Mpem and Djim National Park in April 2019. Central African Republic, where lions are present in Bamingui-Bangoran National Park and Biosphere Reserve, Manovo-Gounda St. Floris and Awakaba National Parks, Aouk Aoukale, Yata Ngaya, Nana Barya and Zemongo Faunal Reserves, and in several hunting reserves of the country. Estimated lion numbers in the country are generally thought to be unreliable. Chad, where lions inhabit Siniaka-Minia Faunal Reserve and Zakouma and Aouk National Parks, but have been extirpated in Manda National Park. Lions may still be present in pastoral rangelands and mountain ranges outside protected areas. In 2004, the lion population in the country was estimated at maximum of 225 individuals. northern parts of the Democratic Republic of the Congo, where lions permanently inhabit rainforests and clearings in rainforest mixed with savannah grassland. Sudan's Southern Darfur province, where lions were abundant in the 1950s; some caused damage to livestock and were poisoned; 76 lions were shot between 1947 and 1952. Lions were recorded in the Dinder–Alatash protected area complex during surveys between 2015 and 2018. South Sudan, where little is known about lion distribution and population sizes. Lions in Radom and Southern National Parks are probably connected to lions in the Central African Republic. Ethiopia has lions from the Northern as well as from the Southern lion subspecies and is considered an admixture zone. While lions from Gambella National Park belong to the Northern subspecies (P. l. leo), lions from other parts of the country belong to the Southern lion.
Range countries Lion Conservation Units Area in km2 Nigeria Yankari National Park2,250Cameroon Waza and Bénoué National Parks 16,134Central African Republic eastern part of the country; Bozoum and Nana Barya Faunal Reserves 339,481Chad southeastern part 133,408Democratic Republic of CongoGaramba-Bili Uere 115,671Sudan, South Sudan 331,834South Sudan, Ethiopia Boma-Gambella 106,941
Asian/North African clade The Asiatic lion is the last surviving population of this clade. Once also found in the Middle East, it is nowadays confined in the wild to Gujarat in India. Genetically, the extinct lions from Northern Africa, formally termed as Barbary lions, fall into the same clade as the Asiatic lion. Therefore, the range of this lion clade encompassed historically North Africa, southeastern Europe, the Arabian Peninsula, the Middle East and the Indian subcontinent. In these regions, lions occurred in: the Sahel, mountain ranges of the Sahara, Barbary Coast and Maghreb, the eastern Mediterranean Basin and the Black Sea region, reed swamps of Mesopotamia, wooded steppe vegetation and pistachio-almond woodlands in Iran, the northwestern part of the Indian subcontinent up to Rajasthan and Bengal in North India.
The Barbary lion population in North Africa is extinct since the mid-1960s.