Northern lion

Northern lion
An Asiatic lion in Gir Forest National Park, India
Barbary lioness and cubs, New York Zoo, 1903
Scientific classification
Kingdom:Animalia
Phylum:Chordata
Class:Mammalia
Order:Carnivora
Suborder:Feliformia
Family:Felidae
Subfamily:Pantherinae
Genus:Panthera
Species: P. leo
Subspecies: P. l. leo
Trinomial name
Panthera leo leo
(Linnaeus, 1758)
Synonyms[1]

formerly

  • P. l. azandicus
  • P. l. gambianus
  • P. l. kamptzi
  • P. l. nubicus
  • P. l. persica
  • P. l. senegalensis

The Northern lion[2][3] (Panthera leo leo) is the nominate subspecies of the lion, which is present in the northern portion of Africa (particularly in West and Central Africa) and in India.[4][5] It is regionally extinct in southern Europe, West Asia and North Africa. In West and Central Africa, lions are restricted to fragmented and isolated populations, most of them declining. In India, there is a single surviving population in and around Gir Forest National Park.[6][7][8] The West African population is listed as Critically Endangered on the IUCN Red List; this population is isolated and comprises fewer than 250 mature individuals.[9]

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 clade distinct from lion populations in Southern and East Africa.[10] In 2017, the Cat Classification Task Force of the IUCN Cat Specialist Group subsumed lion populations to two subspecies, namely P. l. leo and P. l. melanochaita. The Northern lion comprises four historically recognized subspecies, the Barbary lion, the Asiatic lion, the West African lion and the Central African lion.[4]

Taxonomic history

Range map including proposed clades and the two subspecies (P. l. leo and P. l. melanochaita) according to genetic research

A lion from Constantine, Algeria was the type specimen for the specific name Felis leo used by Linnaeus in 1758.[11] Due to the location of Algeria in Africa, the term "Northern lion" was coined for lions living in the northern portion of the continent in 1865.[2][3] In the 19th and 20th centuries, several lion specimens from Africa and Asia were described and proposed as subspecies:

In the following decades, there has been much debate among zoologists on the validity of proposed subspecies:

Genetic research

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 197 lion samples from up to 22 countries. They all agree that the lion species comprises two evolutionary groups, one in the northern and eastern parts of its historical range, and the other in Southern and East Africa that diverged between 245,000 and 50,000 years ago. They assume that tropical rainforest and the East African Rift constituted major barriers between the two groups.[23][24][25][26][10] Based on this assessment, lion taxonomy has been revised as comprising two subspecies:[4]

  • P. l. leo in the northern and eastern regions of its historical and contemporary distribution;
  • P. l. melanochaita in the contemporary southern and East African range countries.

In a comprehensive study about the evolution of lions, 357 samples of 11 lion populations were examined, including some hybrid lions. The hybrids had descended from lions captured in Angola and Zimbabwe, and apparently West or Central Africa. Results indicated that four lions from Morocco did not exhibit any unique genetic characteristics and shared mitochondrial haplotypes H5 and H6 with lions from West Africa, and together with them were part of a major mtDNA grouping (lineage III) that also included Asiatic samples. This scenario was well in line with theories on lion evolution: lineage III developed in East Africa and traveled north and west in the first wave of lion expansions about 118,000 years ago. It apparently broke up into haplotypes H5 and H6 within Africa, and then into H7 and H8 in West Asia[24]

Results of genetic analyses indicate that lions in West Africa and northern parts of Central Africa form distinct lion clades, which are more closely related to North African and Asiatic lions than to lions in Southern Africa and southern parts of East Africa. Lions from North Africa and India however, do form one single clade.[26] Analysis of phylogenetic data of 194 lion samples from 22 different countries revealed that Central and West African lions form a phylogeographic group that probably diverged about 186,000–128,000 years ago from the melanochaita group in East and Southern Africa.[10]

Several lions kept in Ethiopia's Addis Ababa Zoo were found to be genetically similar to wild lions from Cameroon and Chad.[27]

Characteristics

A male from Nubia in New York Zoo, 1903

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 2.47–2.84 m (8.1–9.3 ft) with a weight of 148.2–190.9 kg (327–421 lb). Females are smaller and less heavy.[28]

A few lion specimens from West Africa obtained by museums have been described as having shorter manes than lions from other African regions.[18] In general, the West African lion is similar in general appearance and size as lions in other parts of Africa and Asia.[21]

Zoological specimens range in colour from light to dark tawny. Male skins have short manes, light manes, dark manes or long manes.[29] 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 280–311.7 mm (11.02–12.27 in) in females and 338–362 mm (13.3–14.3 in) in males.[29][18]

Distribution and habitat

Historically, the northern subspecies' range encompassed North Africa, southeastern Europe, the Arabian Peninsula and Middle East in Western Asia.[4] In these regions, lions occurred in:

Today, the subspecies occurs in West and Central Africa and India.[4] It is regionally extinct in Gambia, Mauritania, Sierra Leone, the Western Sahara, Morocco, Algeria, Tunisia, Libya, Egypt, Saudi Arabia, Kuwait, Jordan, Lebanon, Syria, Turkey, Palestine-Israel, Iraq, Iran, Afghanistan and Pakistan. Its range has declined to the:[6]

Asiatic population

Men with live lion, photographed in Iran by Antoin Sevruguin around the end of the 19th century. The Asiatic lion is now extinct in Iran.[41]

The Asiatic lion is the last wild lion population in Asia and restricted to Gir Forest National Park and surrounding areas in Gujarat, India. It is listed as Endangered on the IUCN Red List because of its small size and area of occupancy.[42] This population inhabits remnant forest habitats in the two hill systems of Gir and Girnar that comprise Gujarat's largest tracts of dry deciduous forest, thorny forest and savanna.[40]

Western and Central clades

The West African population is distributed south of the Sahara from Senegal in the west to Nigeria in the east. This population has lost 99% of its former range.[43] It is regionally extinct in Mauritania, Gambia, Guinea-Bissau, Mali, Sierra Leone, Ivory Coast, and Togo, and possibly extinct in Guinea.[9] The largest West African subpopulation of between 246 and 466 individuals survives 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.[43][44]

The range of the Central African lion clade reaches from the lower Niger river in West Africa to Ethiopia, encompassing Cameroon, Central African Republic, northern parts of the Democratic Republic of the Congo, southern parts of Chad and Sudan, and South Sudan.[10] Some Central African populations permanently inhabit rainforests and clearings in rainforest mixed with savannah grassland.[7] In Cameroon, lions are present in Bénoué National Park, and smaller lion groups also in Waza National Park.[5] Habitat in Waza National Park comprises foremost dry woodland that is partly flooded during the rainy season from July to December. In this protected area, two radio-collared male lions used home ranges of between 428 and 1,054 km2 (165 and 407 sq mi), both inside and outside the park. Three radio-collared females had home ranges of between 352 and 724 km2 (136 and 280 sq mi) and stayed inside the park during most of the survey period.[45] In the Central African Republic, 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.[39] Estimated lion numbers in the country are generally thought to be unreliable.[5][46] Lions in Virunga National Park form a contiguous population with lions in the East African country of Uganda.[5][46] In Chad, 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.[7] In 2004, the lion population in the country was estimated at maximum 225 individuals.[5] The following table shows estimates of lion population sizes between 2002 and 2012:

Range countriesArea used in km2Estimated no. of individuals
Bénoué National Park complex, Cameroon14,682200[37]
Waza National Park in Cameroon1,45217[38]
Central African Republic339,418about 1,297[39]
Garamba National Park and Domaine Chasse Bili Uere, Democratic Republic of the Congo115,671175−320[46][7]
Southeastern Chad133,408400[46]
Southwestern Sudan331,834375[46]
Boma and Gambela National Parks, South Sudan and Ethiopia106,941500[46]
Omo National Park, Sudan and Ethiopia22,483200[46]
other protected and non-protected areas, Ethiopia70,75996[46]
Total1,136,6483,260−3,305

The range of the Central African lion clade reaches from the lower Niger river in West Africa to Ethiopia, encompassing Cameroon, Central African Republic, northern parts of the Democratic Republic of the Congo, southern parts of Chad and Sudan, and South Sudan.[10] Some Central African populations permanently inhabit rainforests and clearings in rainforest mixed with savannah grassland.[7]

Contemporary lion distribution and habitat quality in savannahs of West and Central Africa was assessed in 2005, and Lion Conservation Units (LCU) mapped.[47] Educated guesses for size of populations in these LCUs ranged from 3,274 to 3,909 individuals between 2002 and 2012.[7][8]

Range countriesLion Conservation UnitsArea in km2
Senegal, Mali, Guinea-Bissau, GuineaNiokolo-Koba National Park90,384[8]
GuineaNational Park of Upper Niger613[8]
Benin, Burkina Faso, NigerW-Arly-Pendjari Complex29,403[8]
Beninthree unprotected areas6,833[8]
NigeriaYankari National Park and Kainji National Park6,551[8]
CameroonWaza and Bénoué National Parks16,134[38]
Central African Republiceastern part of the country; Bozoum and Nana Barya Faunal Reserves339,481[39]
Chadsoutheastern part133,408[8]
Democratic Republic of CongoGaramba-Bili Uere115,671[47]
South Sudan, Sudan331,834[47]
South Sudan, EthiopiaBoma-Gambella106,941
EthiopiaSouth Omo, Nechisar, Bale, Welmel-Genale, Awash National Parks, Ogaden93,274[8]

Former range

Men with live lion, photographed in Iran by Antoin Sevruguin around the end of the 19th century

Lion populations in southern Europe and Transcaucasia were eradicated between the first and 10th centuries.[48][49][31]

Populations of the Asiatic lion became fragmented during the 19th century and regionally extinct in Turkey and the Middle East between the early and mid-20th century.[50][51][52]

North African populations also declined since the mid-19th century and were eradicated by the early 1960s.[53][30]

Behaviour and ecology

In Benin's Pendjari National Park, groups of lions range from 1–8 individuals. Outside the national park, groups are smaller and more single male lions occur.[54]

In general, lions prefer large prey species within a weight range of 190 to 550 kg (420 to 1,210 lb). They hunt large ungulates in the range of 40.0 to 270.0 kg (88.2 to 595.2 pounds) including zebra, warthog, blue wildebeest, impala, gemsbok, Thomson's gazelle, kob, giraffe and Cape Buffalo.[55] In India's Gir Forest National Park, lions predominantly kill chital, sambar, nilgai, cattle, buffalo and less frequently also wild boar. Outside the protected area where wild prey species do not occur, lions prey on buffalo and cattle, rarely also on camel. They kill most prey less than 100 m (330 ft) away from water bodies, charge prey from close range and drag carcasses into dense cover.[56]

The African lion and spotted hyena 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.[57]

Threats

In Africa, lions are killed pre-emptively or in retaliation for preying on livestock. Populations are also threatened by depletion of prey base, loss and conversion of habitat. Since 1996, African lion populations have been listed as Vulnerable on the IUCN Red List.[6]

The West African lion is fragmented and isolated, comprising fewer than 250 mature individuals.[9] It is threatened by poaching and illegal trade of body parts. Lion body parts from Benin are smuggled to Niger, Nigeria, Gabon, Ivory Coast, Senegal and Guinea, and from Burkina Faso to Benin, Ivory Coast, Senegal and Guinea.[58] The Central African lion is threatened by loss of habitat and prey base and trophy hunting.[7][38][37]

Conservation

Two captive cubs at the Olusegun Obasanjo Library, Nigeria

In India, the lion is protected, and included in CITES Appendix I.[59] African lions are included in CITES Appendix II.[6] In 2004, it was proposed in 2004 to list all lion populations in CITES Appendix I to reduce exports of lion trophies and implement a stricter permission process, due to the negative impact of trophy hunting.[60]

In 2006, a Lion Conservation Strategy for West and Central Africa was developed in cooperation between IUCN regional offices and several wildlife conservation organisations. The strategy envisages to maintain sufficient habitat, ensure a sufficient wild prey base, make lion-human coexistence sustainable and reduce factors that lead to further fragmentation of populations.[47]

In captivity

In 2006, 1258 captive lions were registered in the International Species Information System, including 13 individuals originating from Senegal to Cameroon, 115 from India and 970 with uncertain origin.[23]

Cultural significance

Coat of arms of Sierra Leone

The title "Lion of Mali" was given to Marijata of the Mali Empire.[61][62]

The Cameroon national football team is nicknamed "The Indomitable Lions."[63]

See also

References

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