TheBarbary lion was apopulation of the lion subspeciesPanthera leo leo. It was also calledNorth African lion,Atlas lion, andEgyptian lion. It lived in the mountains and deserts of theMaghreb ofNorth Africa fromMorocco toEgypt. It was eradicated following the spread offirearms andbounties for shooting lions. A comprehensive review of hunting and sighting records revealed that small groups of lions may have survived in Algeria until the early 1960s, and in Morocco until the mid-1960s. Today, it islocally extinct in this region.Fossils of the Barbary lion dating to between 100,000 and 110,000 years were found in the cave of Bizmoune nearEssaouira.
Until 2017, the Barbary lion was considered a distinctlion subspecies. Results ofmorphological andgenetic analyses of lion samples from North Africa showed that the Barbary lion does not differ significantly from theAsiatic lion and falls into the samesubclade. This North African/Asian subclade is closely related to lions fromWest Africa and northern parts ofCentral Africa, and therefore grouped into the northern lion subspeciesPanthera leo leo.
Barbary lionzoological specimens range in colour from light to dark tawny. Male lion skins had manes of varying colouration and length.[2]Head-to-tail length of stuffed males in zoological collections varies from 2.35 to 2.8 m (7 ft 9 in to 9 ft 2 in), and of females around 2.5 m (8 ft 2 in). Skull size varied from 30.85 to 37.23 cm (1 ft 0.15 in to 1 ft 2.66 in). Some manes extended over the shoulder and under the belly to the elbows. The mane hair was 8 to 22 cm (3.1 in to 8.7 in) long.[2][3][4]
In 19th-century hunter accounts, the Barbary lion was claimed to be the largest lion, with a weight of wild males ranging from 270 to 300 kg (600 to 660 lb).[5] Yet, the accuracy of such data measured in the field is questionable. Captive Barbary lions were much smaller but kept under such poor conditions that they might not have attained their full potential size and weight.[5]
The colour and size of lions' manes was long thought to be a sufficiently distinctmorphological characteristic to accord asubspecific status to lion populations.[6] Mane development varies with age and between individuals from different regions, and is therefore not a sufficient characteristic for subspecific identification.[7] The size of manes is not regarded as evidence for Barbary lions' ancestry. Instead, results ofmitochondrial DNA research support the genetic distinctness of Barbary lions in a uniquehaplotype found in museum specimens that is thought to be of Barbary lion descent. The presence of this haplotype is considered a reliablemolecular marker to identify captive Barbary lions.[8]Barbary lions may have developed long-haired manes, because of lower temperatures in the Atlas Mountains than in other African regions, particularly in winter.[5]Results of a long-term study on lions inSerengeti National Park indicate thatambient temperature, nutrition and the level oftestosterone influence the colour and size of lion manes.[9]
Taxonomy
Map shows range ofP. l. leo andP. l. melanochaita[10]
In the 20th and early 21st centuries, there has been much debate and controversy among zoologists on lionclassification and validity of proposed subspecies:
The Barbary lion was considered a distinctlion subspecies.[21][19]In 2017, the Cat Classification Task Force of the Cat Specialist Group subsumed the lion populations in North, West and Central Africa andAsia toP. l. leo.[22]
The Barbary lion was also called North African lion,[1] Atlas lion,[23] and Egyptian lion.[24]
While the historical Barbary lion wasmorphologically distinct, its genetic uniqueness remained questionable.[25]In a comprehensive study about the evolution of lions in 2008, 357 samples of wild and captive lions from Africa and India were examined. Results showed that four captive lions from Morocco did not exhibit any unique genetic characteristic, but sharedmitochondrialhaplotypes with lion samples fromWest andCentral Africa. They were all part of a majormtDNA grouping that also included Asiatic lion samples. Results provided evidence for the hypothesis that this group developed in East Africa, and about 118,000 years ago traveled north and west in the first wave of lion expansion. It broke up within Africa, and later inWest Asia. Lions in Africa probably constitute a single population that interbred during several waves of migration since theLate Pleistocene.[26]Genome-wide data of a wild-born historical lion specimen from Sudan clustered withP. l. leo in mtDNA-based phylogenies, but with a high affinity toP. l. melanochaita.[27]
A comprehensive genetic study published in 2016 confirmed the close relationship between the extinct Barbary lions from Northern Africa and lions from Central and West Africa and in addition showed that the former fall into the same subclade as the Asiatic lion.[28]
Former distribution and habitat
The last photograph of a wild lion in the Atlas Mountains, taken byMarcelin Flandrin in 1925[29]This detail of a map by Jan Janssonius (1588-1664) shows the former "Barbary Coast" of North Africa, known in the 17th century as Barbaria, now covered by Algeria.
Fossils of the Barbary lion dating to between 100,000 and 110,000 years were found in the cave of Bizmoune nearEssaouira.[30][31]The Barbary lion lived in the mountains and deserts of theMaghreb ofNorth Africa fromMorocco toEgypt. It was eradicated following the spread offirearms andbounties for shooting lions.[1]Today, it islocally extinct in this region.[20]Historical sighting and hunting records from the 19th and 20th centuries show that the Barbary lion survived in Algeria until the early 1960s, and in Morocco until the mid-1960s. It inhabitedMediterranean forests, woodlands, and scrub. The westernmost sighting of a Barbary lion reportedly occurred in theAnti-Atlas in western Morocco. It ranged from theAtlas Mountains and theRif in Morocco, theKsour andAmour Ranges in Algeria to theAurès Mountains inTunisia.[29]In Algeria, the Barbary lion was sighted in the forested hills and mountains betweenOuarsenis in the west to theChelif River plains in the north and the Pic deTaza in the east. It inhabited the forests and wooded hills of theConstantine Province southward into the Aurès Mountains.[1]
In the 1830s, lions may have already been eliminated along the coast of theMediterranean Sea and near human settlements.[32]InLibya, the Barbary lion persisted along the coast until the beginning of the 18th century, and was extirpated in Tunisia by 1890.[33] By the mid-19th century, the Barbary lion population had massively declined, sincebounties were paid for shooting lions. The cedar forests ofChelia and neighbouring mountains in Algeria harboured lions until about 1884.[1] The Barbary lion disappeared in theBône region by 1890, in theKhroumire andSouk Ahras regions by 1891, and inBatna Province by 1893.[34]The last recorded shooting of a wild Barbary lion took place in 1942 nearTizi n'Tichka in the Moroccan part of the Atlas Mountains. A smallrelict population may have survived in remote montane areas into the early 1960s.[29] The last known sighting of a lion inAlgeria occurred in 1956 inBeni Ourtilane District.[29]
Historical accounts indicate that in Egypt, lions occurred in theSinai Peninsula, along theNile, in theEastern andWestern Deserts, in the region ofWadi El Natrun and along the maritime coast of the Mediterranean.[35] In the 14th centuryBC,Thutmose IV hunted lions in the hills nearMemphis.[36] The growth of civilizations along the Nile and in the Sinai Peninsula by the beginning of the second millennium BC anddesertification contributed to isolating lion populations in North Africa.[37]
Behaviour and ecology
In the early 20th century, when Barbary lions were rare, they were sighted in pairs or in small family groups comprising a male and female lion with one or two cubs.[1] Between 1839 and 1942, sightings of wild lions involved solitary animals, pairs and family units. Analysis of these sightings indicate that lions retained living in prides even when under increasing persecution, particularly in the eastern Maghreb. The size of prides was likely similar to prides living in sub-Saharan habitats, whereas the density of the Barbary lion population is considered to have been lower than in moister habitats.[29]
WhenBarbary stag (Cervus elaphus barbarus) andgazelles became scarce in the Atlas Mountains, lions preyed on herds of livestock that were carefully tended.[38] They also preyed onwild boar (Sus scrofa).[39]
The lions kept in themenagerie at theTower of London in theMiddle Ages were Barbary lions, as shown byDNA testing on two well-preserved skulls excavated at the Tower in 1936 and 1937. The skulls wereradiocarbon-dated to around 1280–1385 and 1420−1480.[37]In the 19th century and the early 20th century, lions were often kept in hotels andcircusmenageries. In 1835, the lions in the Tower of London were transferred to improved enclosures at theLondon Zoo on the orders of theDuke of Wellington.[41]
The lions in theRabat Zoo exhibited characteristics thought typical for the Barbary lion.[42] Nobles andBerber people presented lions as gifts to the royal family of Morocco. When the family was forced into exile in 1953, the lions in Rabat, numbering 21 altogether, were transferred to two zoos in the region. Three of these were shifted to a zoo inCasablanca, with the rest being shifted toMeknès. The lions at Meknès were moved back to the palace in 1955, but those at Casablanca never came back. In the late 1960s, new lion enclosures were built inTemara nearRabat.[5] Results of amtDNA research revealed in 2006 that a lion kept in the GermanZoo Neuwied originated from this collection and is very likely a descendant of a Barbary lion.[23]Five lion samples from this collection were not Barbary lions maternally. Nonetheless, genes of the Barbary lion are likely to be present in common European zoo lions, since this was one of the most frequently introduced subspecies. Many lions in European and American zoos, which are managed without subspecies classification, are most likely descendants of Barbary lions.[6] Several researchers and zoos supported the development of a studbook of lions directly descended from the King of Morocco's collection.[25]
At the beginning of the 21st century, theAddis Ababa Zoo kept 16 adult lions. With their dark, brown manes extending through the front legs, they looked like Barbary orCape lions. Their ancestors were caught in southwestern Ethiopia as part of a zoological collection for EmperorHaile Selassie of Ethiopia.[43]
The lion also appeared frequently in earlyEgyptian art andliterature.[45] Statues and statuettes of lions found atHierakonpolis andKoptos inUpper Egypt date to theEarly Dynastic Period.[46] The early Egyptian deityMehit was depicted with a lion head.[47] InAncient Egypt, the lion-headed deitySekhmet was venerated as protector of the country.[48] She represented destructive power, but was also regarded as protector against famine and disease. Lion-headed figures and amulets were excavated in tombs in theAegean islands ofCrete,Euboea,Rhodes,Paros andChios. They are associated with Sekhmet and date to the earlyIron Age between the 9th and 6th centuries BC.[49] The remains of seven mostly subadult lions were excavated at the necropolisUmm El Qa'ab in a tomb ofHor-Aha, dated to the 31st century BC.[50] In 2001, the skeleton of amummified lion was found in the tomb ofMaïa in anecropolis dedicated toTutankhamun atSaqqara.[51] It had probably lived and died in thePtolemaic period, showed signs of malnutrition and had probably lived in captivity for many years.[52]The Barbary lion is a symbol in Nubian culture and was often depicted in art and architecture. Nubian deities, such asAmun,Amesemi,Apedemak,Arensnuphis,Hathor,Bastet,Dedun, Mehit,Menhit, andSebiumeker, were depicted as lion protectors inKushite religion.[53]
TheMorocco national football team is called the "Atlas Lions", and the supporters are usually seen wearing T-shirts with a lion's face or wearing a lion suit.[55]
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