Barbary lion: Difference between revisions

Barbary lion
	Barbary lion in Algeria, 1893
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Carnivora
Suborder:	Feliformia
Family:	Felidae
Subfamily:	Pantherinae
Genus:	Panthera
Species:	P. leo;
Subspecies:	P. l. leo
Population:	†Barbary lion

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The Barbary lion was a population of the lion subspecies Panthera leo leo. It was also called North African lion, Atlas lion and Egyptian lion. It lived in the mountains and deserts of the Maghreb of North Africa from Morocco to Egypt. It was eradicated following the spread of firearms and bounties 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 is locally 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 near Essaouira.

Until 2017, the Barbary lion was considered a distinct lion subspecies. Results of morphological and genetic analyses of lion samples from North Africa showed that the Barbary lion does not differ significantly from the Asiatic lion and falls into the same subclade. This North African/Asian subclade is closely related to lions from West Africa and northern parts of Central Africa, and therefore grouped into the northern lion subspecies Panthera leo leo.

Characteristics

Barbary lion zoological 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 distinct morphological characteristic to accord a subspecific 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 of mitochondrial DNA research support the genetic distinctness of Barbary lions in a unique haplotype found in museum specimens that is thought to be of Barbary lion descent. The presence of this haplotype is considered a reliable molecular 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 in Serengeti National Park indicate that ambient temperature, nutrition and the level of testosterone influence the colour and size of lion manes.^[9]

Taxonomy

Felis leo was the scientific name proposed by Carl Linnaeus in 1758 for a type specimen from Constantine, Algeria.^[11] Following Linnaeus's description, several lion zoological specimens from North Africa were described and proposed as subspecies in the 19th century:

Felis leo barbaricus, described by the Austrian zoologist Johann Nepomuk Meyer in 1826, was a lion skin from the Barbary Coast.^[12]
Felis leo nubicus, described by Henri Marie Ducrotay de Blainville in 1843, 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.^[13]

In 1930, Reginald Innes Pocock subordinated the lion to the genus Panthera, when he wrote about the Asiatic lion.^[14]

In the 20th and early 21st centuries, there has been much debate and controversy among zoologists on lion classification and validity of proposed subspecies:

In 1939, Glover Morrill Allen considered F. l. barbaricus and nubicus synonymous with F. l. leo.^[15]
In 1951, John Ellerman and Terence Morrison-Scott recognized only two lion subspecies in the Palearctic realm, namely the African lion Panthera leo leo and the Asiatic lion P. l. persica.^[16]
Some authors considered P. l. nubicus a valid subspecies and synonymous with P. l. massaica.^[3]^[17]^[18]
In 2005, P. l. barbarica, nubica and somaliensis were subsumed under P. l. leo.^[19]
In 2016, IUCN Red List assessors used P. l. leo for all lion populations in Africa.^[20]^[19]

The Barbary lion was considered a distinct lion 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 and Asia to P. l. leo.^[22]

The Barbary lion was also called North African lion,^[1] Atlas lion,^[23] and Egyptian lion.^[24]

Genetic research

Results of a phylogeographic analysis using samples from African and Asiatic lions was published in 2006. One of the African samples was a vertebra from the National Museum of Natural History (France) that originated in the Nubian part of Sudan. In terms of mitochondrial DNA, it grouped with lion skull samples from the Central African Republic, Ethiopia and the northern part of the Democratic Republic of the Congo.^[8]

While the historical Barbary lion was morphologically 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 shared mitochondrial haplotypes with lion samples from West and Central Africa. They were all part of a major mtDNA 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 in West Asia. Lions in Africa probably constitute a single population that interbred during several waves of migration since the Late Pleistocene.^[26] Genome-wide data of a wild-born historical lion specimen from Sudan clustered with P. l. leo in mtDNA-based phylogenies, but with a high affinity to P. 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

Fossils of the Barbary lion dating to between 100,000 and 110,000 years were found in the cave of Bizmoune near Essaouira.^[30]^[31] The Barbary lion lived in the mountains and deserts of the Maghreb of North Africa from Morocco to Egypt. It was eradicated following the spread of firearms and bounties for shooting lions.^[1] Today, it is locally 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 inhabited Mediterranean forests, woodlands, and scrub. The westernmost sighting of a Barbary lion reportedly occurred in the Anti-Atlas in western Morocco. It ranged from the Atlas Mountains and the Rif in Morocco, the Ksour and Amour Ranges in Algeria to the Aurès Mountains in Tunisia.^[29] In Algeria, the Barbary lion was sighted in the forested hills and mountains between Ouarsenis in the west to the Chelif River plains in the north and the Pic de Taza in the east. It inhabited the forests and wooded hills of the Constantine Province southward into the Aurès Mountains.^[1]

In the 1830s, lions may have already been eliminated along the coast of the Mediterranean Sea and near human settlements.^[32] In Libya, 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, since bounties were paid for shooting lions. The cedar forests of Chelia and neighbouring mountains in Algeria harboured lions until about 1884.^[1] The Barbary lion disappeared in the Bône region by 1890, in the Khroumire and Souk Ahras regions by 1891, and in Batna Province by 1893.^[34] The last recorded shooting of a wild Barbary lion took place in 1942 near Tizi n'Tichka in the Moroccan part of the Atlas Mountains. A small relict population may have survived in remote montane areas into the early 1960s.^[29] The last known sighting of a lion in Algeria occurred in 1956 in Beni Ourtilane District.^[29]

Historical accounts indicate that in Egypt, lions occurred in the Sinai Peninsula, along the Nile, in the Eastern and Western Deserts, in the region of Wadi El Natrun and along the maritime coast of the Mediterranean.^[35] In the 14th century BC, Thutmose IV hunted lions in the hills near Memphis.^[36] The growth of civilizations along the Nile and in the Sinai Peninsula by the beginning of the second millennium BC and desertification 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]

When Barbary stag (Cervus elaphus barbarus) and gazelles became scarce in the Atlas Mountains, lions preyed on herds of livestock that were carefully tended.^[38] They also preyed on wild boar (Sus scrofa).^[39]

Sympatric predators in this region included the African leopard (P. pardus pardus) and Atlas bear (Ursus arctos crowtheri).^[21]^[40]

In captivity

The lions kept in the menagerie at the Tower of London in the Middle Ages were Barbary lions, as shown by DNA testing on two well-preserved skulls excavated at the Tower in 1936 and 1937. The skulls were radiocarbon-dated to around 1280–1385 and 1420−1480.^[37] In the 19th century and the early 20th century, lions were often kept in hotels and circus menageries. In 1835, the lions in the Tower of London were transferred to improved enclosures at the London Zoo on the orders of the Duke of Wellington.^[41]

The lions in the Rabat Zoo exhibited characteristics thought typical for the Barbary lion.^[42] Nobles and Berber 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 in Casablanca, with the rest being shifted to Meknès. 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 in Temara near Rabat.^[5] Results of a mtDNA research revealed in 2006 that a lion kept in the German Zoo 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, the Addis Ababa Zoo kept 16 adult lions. With their dark, brown manes extending through the front legs, they looked like Barbary or Cape lions. Their ancestors were caught in southwestern Ethiopia as part of a zoological collection for Emperor Haile Selassie of Ethiopia.^[43]

Since 2005, three Barbary lions were kept in Belfast Zoo obtained from Port Lympne Wild Animal Park, and a new Barbary lion enclosure was opened in 2023.^[44]

Cultural significance

The lion also appeared frequently in early Egyptian art and literature.^[45] Statues and statuettes of lions found at Hierakonpolis and Koptos in Upper Egypt date to the Early Dynastic Period.^[46] The early Egyptian deity Mehit was depicted with a lion head.^[47] In Ancient Egypt, the lion-headed deity Sekhmet 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 the Aegean islands of Crete, Euboea, Rhodes, Paros and Chios. They are associated with Sekhmet and date to the early Iron Age between the 9th and 6th centuries BC.^[49] The remains of seven mostly subadult lions were excavated at the necropolis Umm El Qa'ab in a tomb of Hor-Aha, dated to the 31st century BC.^[50] In 2001, the skeleton of a mummified lion was found in the tomb of Maïa in a necropolis dedicated to Tutankhamun at Saqqara.^[51] It had probably lived and died in the Ptolemaic 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 as Amun, Amesemi, Apedemak, Arensnuphis, Hathor, Bastet, Dedun, Mehit, Menhit, and Sebiumeker, were depicted as lion protectors in Kushite religion.^[53]

In Roman North Africa, lions were regularly captured by experienced hunters for venatio spectacles in amphitheatres.^[39]^[54]

The Morocco 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]

References

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External links

Marks, T. (2018). "'I find myself making growling noises while I'm painting' – an interview with Walton Ford, who painted Barbary lions". Apollo Magazine.
Babas, L. (2018). "History : When London's very first zoo housed Morocco's Atlas Lions". Yabiladi.
Moroccan 'Atlas' lion at Parc Sindibad, Casablanca (video). Archived from the original on 2021-12-22.
Black, S. (2014). "Moroccan lions in zoos today". University of Kent Blog.
"Barbary Lion Information". Being Lion.

[Pease1913-1] ^ ^a ^b ^c ^d ^e ^f Pease, A. E. (1913). "The Distribution of Lions". The Book of the Lion. London: John Murray. pp. 109−147.

[Mazak70-2] Mazák, V. (1970). "The Barbary lion, Panthera leo leo (Linnaeus, 1758); some systematic notes, and an interim list of the specimens preserved in European museums". Zeitschrift für Säugetierkunde. 35: 34−45.

[Hemmer1974-3] Hemmer, H. (1974). "Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758)". Veröffentlichungen der Zoologischen Staatssammlung. 17: 167–280.

[4] Mazák, J. H. (2010). "Geographical variation and phylogenetics of modern lions based on craniometric data". Journal of Zoology. 281 (3): 194–209. doi:10.1111/j.1469-7998.2010.00694.x.

[Yamaguchi2002-5] Yamaguchi, N.; Haddane, B. (2002). "The North African Barbary Lion and the Atlas Lion Project". International Zoo News. 49 (8): 465–481.

[BarnettYamaguchi2006-6] Barnett, R.; Yamaguchi, N.; Barnes, I.; Cooper, A. (2006). "Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation" (PDF). Conservation Genetics. 7 (4): 507–514. Bibcode:2006ConG....7..507B. doi:10.1007/s10592-005-9062-0. S2CID 24190889. Archived from the original (PDF) on 2006-08-24.

[7] O'Brien, S. J.; Joslin, P.; Smith, G. L.; Wolfe, R.; Schaffer, N.; Heath, E.; Ott-Joslin, J.; Rawal, P. P.; Bhattacharjee, K. K.; Martenson, J. S. (1987). "Evidence for African origins of founders of the Asiatic Lion Species Survival Plan". Zoo Biology. 6 (2): 99–116. doi:10.1002/zoo.1430060202.

[Barnett2006-8] Barnett, R.; Yamaguchi, N.; Barnes, I. & Cooper, A. (2006). "The origin, current diversity and future conservation of the modern lion (Panthera leo)" (PDF). Proceedings of the Royal Society B: Biological Sciences. 273 (1598): 2119–2125. doi:10.1098/rspb.2006.3555. PMC 1635511. PMID 16901830. Archived from the original (PDF) on 8 August 2007.

[West2002-9] West, P. M.; Packer, C. (2002). "Sexual selection, temperature, and the lion's mane". Science. 297 (5585): 1339–1343. Bibcode:2002Sci...297.1339W. doi:10.1126/science.1073257. PMID 12193785. S2CID 15893512.

[Bertola2016-10] Bertola, L.D.; Jongbloed, H.; Van Der Gaag K.J.; De Knijff, P.; Yamaguchi, N.; Hooghiemstra, H.; Bauer, H.; Henschel, P.; White, P.A.; Driscoll, C.A.; Tende, T.; Ottosson, U.; Saidu, Y.; Vrieling, K. & de Iongh, H. H. (2016). "Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (Panthera leo)". Scientific Reports. 6: 30807. Bibcode:2016NatSR...630807B. doi:10.1038/srep30807. PMC 4973251. PMID 27488946.

[Linn1758-11] Linnaeus, C. (1758). "Felis Leo". Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (in Latin). Vol. 1 (10th ed.). Holmiae (Laurentii Salvii). p. 41. Retrieved 8 September 2008.

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[DeManuel_al.2020-27] Manuel, M. d.; Ross, B.; Sandoval-Velasco, M.; Yamaguchi, N.; Vieira, F. G.; Mendoza, M. L. Z.; Liu, S.; Martin, M. D.; Sinding, M.-H. S.; Mak, S. S. T.; Carøe, C.; Liu, S.; Guo, C.; Zheng, J.; Zazula, G.; Baryshnikov, G.; Eizirik, E.; Koepfli, K.-P.; Johnson, W. E.; Antunes, A.; Sicheritz-Ponten, T.; Gopalakrishnan, S.; Larson, G.; Yang, H.; O’Brien, S. J.; Hansen, A. J.; Zhang, G.; Marques-Bonet, T. & Gilbert, M. T. P. (2020). "The evolutionary history of extinct and living lions". Proceedings of the National Academy of Sciences of the United States of America. 117 (20): 10927–10934. Bibcode:2020PNAS..11710927D. doi:10.1073/pnas.1919423117. PMC 7245068. PMID 32366643.

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@@ Line 1: / Line 1: @@
+{{Short description|Lion population}}
-{{Taxobox
+{{Population taxobox
-| color = pink
+|genus = Panthera
-| name = Barbary Lion
-| status = EW
+|species = leo
+|subspecies = leo
-| image = BarbaryLionB1898bw.jpg
+|population =  Barbary lion
-| regnum = [[Animal]]ia
+|extinct = yes
-| phylum = [[Chordate|Chordata]]
+|image = Barbary lion.jpg
-| classis = [[Mammal]]ia
+|image_caption = Barbary lion in Algeria, 1893<ref name=Pease1913/>
-| ordo = [[Carnivora]]
-| familia = [[Felidae]]
-| genus = ''[[Panthera]]''
-| species = ''[[Lion|P. leo]]''
-| subspecies = '''''P. l. leo'''''
-| trinomial = ''Panthera leo leo''
-| trinomial_authority = ([[Carolus Linnaeus|Linnaeus]], [[1758]])
-| synonyms = <center>''Felis leo'' <small>[[Carolus Linnaeus|Linnaeus]], [[1758]]</small><br>
-''Panthera leo berberisca''</center>
 }}
-The '''Barbary Lion''', '''Atlas lion''' or '''Nubian lion''' ''Panthera leo leo'' is a [[subspecies]] of lion that has become [[Extinction|extinct]] in the wild. It was believed to be extinct in captivity. However, possible Barbary lion individuals or descendants have been located in zoos and circus populations within the last three decades. It is often considered to be the largest of the lion subspecies with males weighing between 400-650 lbs (181 to 295 kg) and females 270-400 lbs (120 to 181 kg), approximately the size of [[Bengal tigers]]. However, more recent research suggests that it is only slightly larger than modern African lion, which weighs approximately 420 lbs on average. The Barbary Lion, also called the ''Atlas lion'' or ''Nubian lion'', formerly ranged in [[North Africa]] (from [[Morocco]] to [[Tunisia]]) and continuing until [[Egypt]].  The last known Barbary Lion in the wild was shot in the [[Atlas Mountains]] in [[1922]].
+The '''Barbary lion''' was a [[population]] of the lion subspecies ''[[Panthera leo leo]]''. It was also called '''North African lion''', '''Atlas lion''' and '''Egyptian lion'''. It lived in the mountains and deserts of the [[Maghreb]] of [[North Africa]] from [[Morocco]] to [[Egypt]]. It was eradicated following the spread of [[firearms]] and [[Bounty (reward)|bounties]] 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 is [[locally extinct]] in this region. [[Fossil]]s of the Barbary lion dating to between 100,000 and 110,000 years were found in the cave of Bizmoune near [[Essaouira]].
-Unlike most African [[lion]]s, the Barbary Lion was a mountain predator, preferring [[woodland]]s. The two other primary Atlas Mountain predators, the [[Atlas bear]] and [[Barbary leopard]] are now [[extinct]] or close to be, respectively.
+Until 2017, the Barbary lion was considered a distinct [[Lion#Subspecies|lion subspecies]]. Results of [[Morphology (biology)|morphological]] and [[Genetics|genetic]] analyses of lion samples from North Africa showed that the Barbary lion does not differ significantly from the [[Asiatic lion]] and falls into the same [[subclade]]. This North African/Asian subclade is closely related to lions from [[West Africa]] and northern parts of [[Central Africa]], and therefore grouped into the northern lion subspecies ''Panthera leo leo''.
-The mane of "[[Scar (The Lion King)|Scar]]," the [[villain]] of [[The Walt Disney Company|Disney]]'s ''[[The Lion King]]'', was based on a Barbary Lion.
+==Characteristics==
-==Possible surviving individuals==
+[[File:Sultan the Barbary Lion.jpg|thumb|A Barbary lion in the [[Bronx Zoo]], 1897]]
-There are several dozen individuals in captivity believed to be Barbary lions: [[Port Lympne Zoo|Port Lympne Wild Animal Park]] has twelve specimens [http://www.totallywild.net/animals.php?animal=Barbary/Atlas%20Lion] descended from animals owned by the [[King of Morocco]]. In addition, eleven animals believed to be Barbary lions were found in [[Addis Ababa]] zoo, descendants of animals owned by [[Haile Selassie I of Ethiopia|Emperor Haile Selassie]].
+Barbary lion [[zoological specimen]]s range in colour from light to dark tawny. Male lion skins had manes of varying colouration and length.<ref name=Mazak70>{{cite journal |author=Mazák, V. |year=1970 |title=The Barbary lion, ''Panthera leo leo'' (Linnaeus, 1758); some systematic notes, and an interim list of the specimens preserved in European museums |journal=Zeitschrift für Säugetierkunde |volume=35 |pages=34−45}}</ref>
+Head-to-tail length of stuffed males in zoological collections varies from {{cvt|2.35|to|2.8|m|ftin}}, and of females around {{cvt|2.5|m|ftin}}. Skull size varied from {{cvt|30.85|to|37.23|cm|ftin}}. Some manes extended over the shoulder and under the belly to the elbows. The mane hair was {{cvt|8|to|22|cm|ftin}} long.<ref name=Mazak70/><ref name=Hemmer1974>{{cite journal |author=Hemmer, H. |year=1974 |title=Untersuchungen zur Stammesgeschichte der Pantherkatzen (Pantherinae) Teil 3. Zur Artgeschichte des Löwen ''Panthera (Panthera) leo'' (Linnaeus, 1758) |journal=Veröffentlichungen der Zoologischen Staatssammlung |volume=17 |pages=167–280 |url=https://archive.org/stream/verfentlichungen171974zool#page/n215/mode/2up}}</ref><ref>{{cite journal |author=Mazák, J. H. |year=2010 |title=Geographical variation and phylogenetics of modern lions based on craniometric data |journal=Journal of Zoology |volume=281 |issue=3 |pages=194–209 |doi=10.1111/j.1469-7998.2010.00694.x}}</ref>
+In 19th-century hunter accounts, the Barbary lion was claimed to be the largest lion, with a weight of wild males ranging from {{cvt|270|to|300|kg}}.<ref name=Yamaguchi2002 /> 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.<ref name=Yamaguchi2002>{{cite journal |author1=Yamaguchi, N. |author2=Haddane, B. |year=2002 |title=The North African Barbary Lion and the Atlas Lion Project |journal=International Zoo News |volume=49 |issue=8 |pages=465–481 |url=https://www.researchgate.net/publication/266755974}}</ref>
-In the past scientists believed that the distinct sub-species status of the Barbary lion was established by its seemingly fixed external [[morphology (biology)|morphology]], particularly its heavier mane. However, it is now known that various [[extrinsic]] factors influence the colour and size of all lions' manes, such as [[ambient temperature]].{{ref|West2002}} As the cooler ambient temperature in European and North American zoos has been found to produce Barbary-like manes on ordinary lions, this characteristic is now considered an inappropriate marker for identifying Barbary ancestry.{{ref|Yamaguchi2002}}{{ref|Barnett2006}}
+The colour and size of lions' manes was long thought to be a sufficiently distinct [[morphology (biology)|morphological]] characteristic to accord a [[Subspecific name|subspecific]] status to lion populations.<ref name=BarnettYamaguchi2006>{{Cite journal |author1=Barnett, R. |author2=Yamaguchi, N. |author3=Barnes, I. |author4=Cooper, A. |year=2006 |title=Lost populations and preserving genetic diversity in the lion ''Panthera leo'': Implications for its ex situ conservation |journal=Conservation Genetics |volume=7 |issue=4|pages=507–514 |url=http://abc.zoo.ox.ac.uk/Papers/consgen06_lion.pdf |doi=10.1007/s10592-005-9062-0 |bibcode=2006ConG....7..507B |s2cid=24190889 |url-status=dead |archive-url=https://web.archive.org/web/20060824064412/http://abc.zoo.ox.ac.uk/Papers/consgen06_lion.pdf |archive-date=2006-08-24}}</ref> Mane development varies with age and between individuals from different regions, and is therefore not a sufficient characteristic for subspecific identification.<ref>{{cite journal |author=O'Brien, S. J. |author2=Joslin, P. |author3=Smith, G. L. |author4=Wolfe, R. |author5=Schaffer, N. |author6=Heath, E. |author7=Ott-Joslin, J. |author8=Rawal, P. P. |author9=Bhattacharjee, K. K. |author10=Martenson, J. S. |year=1987 |title=Evidence for African origins of founders of the Asiatic Lion Species Survival Plan |journal=Zoo Biology |volume=6 |issue=2 |pages=99–116 |doi=10.1002/zoo.1430060202 |url=https://zenodo.org/record/1229390}}</ref> The size of manes is not regarded as evidence for Barbary lions' ancestry. Instead, results of [[mitochondrial DNA]] research support the genetic distinctness of Barbary lions in a unique [[haplotype]] found in museum specimens that is thought to be of Barbary lion descent. The presence of this haplotype is considered a reliable [[molecular marker]] to identify captive Barbary lions.<ref name=Barnett2006/>
-Despite this, [[Mitochondrial DNA]] research published in 2006 does support the distinctness of the Barbary lions as a sub-species. The results found a unique [[Mitochondrial DNA|mtDNA]] [[haplotype]] to be present in some of those zoo specimens believed to be of Barbary descent. This may be a good molecular marker for identifying -- and excluding -- other potential Barbary lions. The mtDNA results revealed that five tested samples of lions from the famous collection of the King of Morocco are not, according to this criterion, maternally Barbary.{{ref|Barnett2006}}
+Barbary lions may have developed long-haired manes, because of lower temperatures in the Atlas Mountains than in other African regions, particularly in winter.<ref name=Yamaguchi2002/>
+Results of a long-term study on lions in [[Serengeti National Park]] indicate that [[ambient temperature]], nutrition and the level of [[testosterone]] influence the colour and size of lion manes.<ref name=West2002>{{Cite journal|author=West, P. M. |author2=Packer, C. |title=Sexual selection, temperature, and the lion's mane |journal=Science |volume=297 |issue=5585 |pages=1339–1343 |year=2002 |pmid=12193785 |doi=10.1126/science.1073257|bibcode=2002Sci...297.1339W |s2cid=15893512}}</ref>
+==Taxonomy==
-== The Barbary Lion Project ==
+[[File:Lion subspecies distribution3.png|thumb|Map shows range of ''P. l. leo'' and ''P. l. melanochaita''<ref name=Bertola2016>{{cite journal |author1=Bertola, L.D. |author2=Jongbloed, H. |author3=Van Der Gaag K.J. |author4=De Knijff, P. |author5=Yamaguchi, N. |author6=Hooghiemstra, H. |author7=Bauer, H. |author8=Henschel, P. |author9=White, P.A. |author10=Driscoll, C.A. |author11=Tende, T. |author12=Ottosson, U. |author13=Saidu, Y. |author14=Vrieling, K. |author15=de Iongh, H. H. |name-list-style=amp |year=2016 |title=Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (''Panthera leo'') |journal=Scientific Reports |volume=6 |page=30807 |doi=10.1038/srep30807|pmid=27488946 |pmc=4973251 |bibcode=2016NatSR...630807B}}</ref>]]
-The former popularity of the Barbary Lion as a zoo animal provides the only hope to ever see it again in the wild in [[North Africa]].
+''Felis leo'' was the [[scientific name]] proposed by [[Carl Linnaeus]] in 1758 for a [[type specimen]] from [[Constantine, Algeria]].<ref name=Linn1758>{{Cite book |last=Linnaeus |first=C. |title=Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis |publisher=Holmiae (Laurentii Salvii) |year=1758 |page=41 |chapter=''Felis Leo'' |language=la |volume=1 |edition=10th |chapter-url=https://www.biodiversitylibrary.org/page/726936 |access-date=8 September 2008}}</ref> Following Linnaeus's description, several lion [[zoological specimen]]s from North Africa were described and proposed as subspecies in the 19th century:
-After years of research into the science of the Barbary Lion and stories of surviving examples, WildLink International, in collaboration with [[Oxford University]], launched their ambitious International Barbary Lion Project. They are using the very latest [[DNA]] techniques to identify the DNA 'fingerprint' of the Barbary Lion [[subspecies]]. [[WildLink International]] has taken bone samples from remains of Barbary Lions in [[Museum]]s across [[Europe]], like those in Brussels, Paris, Turin and others. These samples are returned to Oxford University where the science team is extracting the DNA sequence that identifies the Barbary as a separate subspecies.
+*''Felis leo barbaricus'', described by the [[Austria]]n zoologist Johann Nepomuk Meyer in 1826, was a lion skin from the [[Barbary Coast]].<ref>{{cite book |author=Meyer, J. N. |year=1826 |title=Dissertatio inauguralis anatomico-medica de genere felium. Doctoral thesis |publisher=University of Vienna |location=Vienna}}</ref>
-Although the Barbary is officially extinct, WildLink International had identified a handful of lions in captivity around the world that are descended from the original Barbary Lion, like the royal lions in [[Temara Zoo]] in [[Rabat, Morocco|Rabat]], [[Morocco]]. These descendants will be tested against the DNA fingerprint and the degree of any hybridisation (from crossbreeding) can then be determined. The best candidates will then enter a selective breeding programme that will 'breed back' the Barbary Lion. The final phase of the project will see the lions released into a National Park in the [[Atlas Mountains]] of Morocco.
+*''Felis leo nubicus'', described by [[Henri Marie Ducrotay de Blainville]] in 1843, 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.<ref name="DeBlainville1843">{{cite book |author=Blainville, H. M. D. de |year=1843 |chapter=''F. leo nubicus'' |title=Ostéographie ou description iconographique comparée du squelette et du système dentaire des mammifères récents et fossils pour servir de base à la zoologie et la géologie. Vol 2 |location=Paris |publisher=J. B. Baillière et Fils |page=186}}</ref>
-WildLink International can not be reached anymore and their website is nowadays offline. Everyone is in the dark as to what happened to WildLink International. WildLink International and the University of Oxford had made the deal that WildLink International would raise money for the project and that the university would do the research. With the disappearance of WildLink International no money was raised.
+In 1930, [[Reginald Innes Pocock]] subordinated the lion to the genus ''[[Panthera]]'', when he wrote about the [[Asiatic lion]].<ref>{{cite journal |author=Pocock, R. I. |year=1930 |title=The lions of Asia |journal=Journal of the Bombay Natural Historical Society |volume=34 |pages=638–665}}</ref>
-Dr. Noboyuki Yamaguchi, a scientist from the University of Oxford, has used his own funding for as long as possible to further the scientific research on Barbary Lions and its genetics. The project is now indefinitely on hold until the funds can be raised.{{ref|Yamaguchi2002}}
+In the 20th and early 21st centuries, there has been much debate and controversy among zoologists on lion [[Taxonomy (biology)|classification]] and validity of proposed subspecies:
-== "Asiatic Lion", an Asian relative ==
+*In 1939, [[Glover Morrill Allen]] considered ''F. l. barbaricus'' and ''nubicus'' [[Synonym (taxonomy)|synonymous]] with ''F. l. leo''.<ref name=Allen1939>{{cite journal |author=Allen, G. M. |year=1939 |url=https://archive.org/stream/bulletinofmuseum83harv#page/242/mode/2up |title=A Checklist of African Mammals |journal=Bulletin of the Museum of Comparative Zoology at Harvard College |volume=83 |pages=1–763}}</ref>
-In [[1968]], a study on the skulls of the Barbary, extinct [[Cape Lion|Cape]], [[Asiatic Lion|Asiatic]], and other African lions showed that the same skull characteristics - the very narrow postorbital bar - existed in only the Barbary and the Asiatic lion skulls. This shows that there may have been a close relationship between the lions from Northernmost [[Africa]] and [[Asia]]. It is also believed that the South [[European]] lion that became extinct at the beginning in A.D. 80-100, could have represented the connecting link between the North African and Asiatic lions. It is believed that Barbary lions possess the same belly fold (hidden under all that mane) that appears in the Asian lions today.
+*In 1951, [[Sir John Ellerman, 2nd Baronet|John Ellerman]] and [[Terence Morrison-Scott]] recognized only two lion subspecies in the [[Palearctic realm]], namely the African lion ''Panthera leo leo'' and the Asiatic lion ''P. l. persica''.<ref name=ems66>{{cite book |author=Ellerman, J. R. |author2=Morrison-Scott, T. C. S. |year=1966 |url=https://archive.org/stream/checklistofindia00elle#page/312/mode/2up |title=Checklist of Palaearctic and Indian mammals 1758 to 1946 |edition=Second |publisher=British Museum of Natural History |location=London |pages=312–313}}</ref>
+*Some authors considered ''P. l. nubicus'' a [[Valid name (zoology)|valid]] subspecies and synonymous with ''P. l. massaica''.<ref name=Hemmer1974/><ref name=Haas2005>{{cite journal |author1=Haas, S.K. |author2=Hayssen, V. |author3=Krausman, P.R. |title=''Panthera leo'' |year=2005 |journal=Mammalian Species |volume=2005 |issue=762 |pages=1–11 |url=http://www.science.smith.edu/msi/pdf/762_Panthera_leo.pdf|doi=10.1644/1545-1410(2005)762[0001:PL]2.0.CO;2 |s2cid=198968757 |url-status=dead |archive-url=https://web.archive.org/web/20170728131140/http://www.science.smith.edu/msi/pdf/762_Panthera_leo.pdf |archive-date=28 July 2017}}</ref><ref name="Kingdonetal.2013">{{cite book|author1=West, P. M.|title=Mammals of Africa|author2=Packer, C.|date=2013|publisher=[[A & C Black]] |isbn=978-1-4081-8996-2 |editor1=Kingson, J. |editor-link=Jonathan Kingdon|volume=5|page=149−159|chapter=''Panthera leo'' Lion|editor2=Happold, D.|editor3=Butynski, T.|editor4=Hoffmann, M.|editor5=Happold, M. |editor6=Kalina, J. |chapter-url=https://books.google.com/books?id=B_07noCPc4kC&pg=RA4-PA149}}</ref>
+*In 2005, ''P. l. barbarica'', ''nubica'' and ''somaliensis'' were subsumed under ''P. l. leo''.<ref name=MSW3>{{MSW3 Carnivora |id=14000228 |page=546 |heading=Species ''Panthera leo''}}</ref>
+*In 2016, [[IUCN Red List]] assessors used ''P. l. leo'' for all lion populations in [[Africa]].<ref name=iucn/><ref name=MSW3>{{MSW3 Carnivora |id=14000228 |page=546 |heading=Species ''Panthera leo''}}</ref>
+The Barbary lion was considered a distinct [[Lion#Subspecies|lion subspecies]].<ref name=CAP>{{Cite book |last1=Nowell |first1=K. |last2=Jackson |first2=P. |name-list-style=amp |title=Wild Cats: Status Survey and Conservation Action Plan |year=1996 |publisher=IUCN/SSC Cat Specialist Group |location=Gland, Switzerland |isbn=978-2-8317-0045-8 |pages=17–21 |chapter=African lion, ''Panthera leo'' (Linnaeus, 1758) |url=http://carnivoractionplans1.free.fr/wildcats.pdf}}</ref><ref name=MSW3/>
+In 2017, the Cat Classification Task Force of the Cat Specialist Group subsumed the lion populations in North, West and Central Africa and [[Asia]] to ''P. l. leo''.<ref name=catsg>{{cite journal |author1=Kitchener, A. C. |author2=Breitenmoser-Würsten, C. |author3=Eizirik, E. |author4=Gentry, A. |author5=Werdelin, L. |author6=Wilting, A. |author7=Yamaguchi, N. |author8=Abramov, A. V. |author9=Christiansen, P. |author10=Driscoll, C. |author11=Duckworth, J. W. |author12=Johnson, W. |author13=Luo, S.-J. |author14=Meijaard, E. |author15=O’Donoghue, P. |author16=Sanderson, J. |author17=Seymour, K. |author18=Bruford, M. |author19=Groves, C. |author20=Hoffmann, M. |author21=Nowell, K. |author22=Timmons, Z. |author23=Tobe, S. |name-list-style=amp |year=2017 |title=A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group |journal=Cat News |issue=Special Issue 11 |pages=71–73 |url=https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf?sequence=1&isAllowed=y#page=71}}</ref>
+The Barbary lion was also called North African lion,<ref name=Pease1913/> Atlas lion,<ref name=Burger2006/> and Egyptian lion.<ref name=Geptner>{{cite book |author=Heptner, V. G. |author2=Sludskij, A. A. |orig-year=1972 |year=1992 |title=Mlekopitajuščie Sovetskogo Soiuza. Moskva: Vysšaia Škola |trans-title=Mammals of the Soviet Union. Volume II, Part 2. Carnivora (Hyaenas and Cats) |publisher=Smithsonian Institution and the National Science Foundation |location=Washington DC |chapter=Lion |chapter-url=https://archive.org/stream/mammalsofsov221992gept#page/82/mode/2up |pages=83–95 |isbn=978-90-04-08876-4}}</ref>
-== References ==
-* {{note|Barnett2006}}Barnett, R., N. Yamaguchi, I. Barnes & A. Cooper. 2006. Lost populations and preserving genetic diversity in the lion ''Panthera leo'': Implications for its ''ex situ'' conservation. Conservation Genetics. [http://abc.zoo.ox.ac.uk/Papers/consgen06_lion.pdf Online full-text pdf]
-* {{note|West2002}}West P.M., Packer C. 2002. Sexual selection, temperature, and the lion’s mane. Science, 297, 1339–1343.
-* {{note|Yamaguchi2002}}Yamaguchi, N. & Haddane, B. (2002). The North African Barbary lion and the Atlas Lion Project. International Zoo News 49: 465-481.
-== Port Lympne's lioness ==
-[[Image:Barbary_lioness.JPG|thumb|left]]
-This Barbary [[Lion]] comes from [[Port Lympne Wild Animal Park]]. Her name is Samira, when she was younger her dad accidently broke her leg when she was 4 days old. She was born on the 10th of [[July]] 2003. And on the 14th the accident happened, as a result Samira had to be hand reared by her [[kepper]]. Samira lived with her keeper until she was able to move around easily on her own. She was very vocal as a cub and earned the name Samira which means 'talks a lot' in Arabic. Samira's therapy for her injuries including electrotherapy and hydrotherapy has been shown around the world in various media. Samira is very friendly, but can be quite naughty. If she thinks she is not allowed to do something she will keep trying to do it.
-Samira has been joined in her enclosure by her two brothers Moonlight and Milo.
-== See also ==
+===Genetic research===
+Results of a [[phylogeographic]] analysis using samples from African and Asiatic lions was published in 2006. One of the African samples was a [[vertebra]] from the [[National Museum of Natural History (France)]] that originated in the Nubian part of [[Sudan]]. In terms of [[mitochondrial DNA]], it grouped with lion skull samples from the [[Central African Republic]], [[Ethiopia]] and the northern part of the [[Democratic Republic of the Congo]]<!-- nos. given in table do '''not''' refer to year of collection!! But to museum IDs of samples (ID no. 1996-2516 and 1996-2517 from Paris museum, and ID nos. A59:5062 and A59:5066 dating 1921 from Swedish Museum of Natural History, respectively) -->.<ref name=Barnett2006>{{Cite journal|last1=Barnett |first1=R. |last2=Yamaguchi |first2=N. |last3=Barnes |first3=I. |last4=Cooper |first4=A. |year=2006 |title=The origin, current diversity and future conservation of the modern lion (''Panthera leo'') |journal=Proceedings of the Royal Society B: Biological Sciences |pmid=16901830 |volume=273 |issue=1598 |pmc=1635511 |pages=2119–2125 |doi=10.1098/rspb.2006.3555 |name-list-style=amp |url=http://www.adelaide.edu.au/acad/publications/papers/Barnett%20PRS%20lions.pdf |url-status=dead |archive-url=https://web.archive.org/web/20070808182526/http://www.adelaide.edu.au/acad/publications/papers/Barnett%20PRS%20lions.pdf |archive-date=8 August 2007}}</ref>
-* [[Cape Lion]]
-* [[Asiatic Lion]] Barbary Lion's almost extinct nearest cousin
-* [[European lion]]
-* [[Ex-situ conservation]]
+While the historical Barbary lion was [[Morphology (biology)|morphologically]] distinct, its genetic uniqueness remained questionable.<ref name=Black2010>{{Cite journal |author=Black, S. |author2=Yamaguchi, N. |author3=Harland, A.|author4=Groombridge, J. |name-list-style=amp |title=Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions |journal=European Journal of Wildlife Research |volume=56 |issue=1 |pages=21–31 |year=2010 |doi=10.1007/s10344-009-0280-5 |s2cid=44941372 |url=http://kar.kent.ac.uk/27502/1/Black_et_al_%282009%29_Genetic_Health_of_putative_barbary_lions.pdf}}</ref>
-== External links ==
+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 shared [[mitochondrial]] [[haplotype]]s with lion samples from [[West Africa|West]] and [[Central Africa]]. They were all part of a major [[mtDNA]] 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 in [[West Asia]]. Lions in Africa probably constitute a single population that interbred during several waves of migration since the [[Late Pleistocene]].<ref name=Agostinho>{{Cite journal |author=Antunes, A. |author2=Troyer, J. L. |author3=Roelke, M. E. |author4=Pecon-Slattery, J. |author5=Packer, C. |author6=Winterbach, C. |author7=Winterbach, H. |author8=Johnson, W. E. |name-list-style=amp |title=The Evolutionary Dynamics of the Lion ''Panthera leo'' revealed by Host and Viral Population Genomics |journal=PLOS Genetics |volume=4 |issue=11 |pages=e1000251 |year=2008 |pmid=18989457 |pmc=2572142 |doi=10.1371/journal.pgen.1000251 |doi-access=free }}</ref> [[Genome]]-wide data of a wild-born historical lion specimen from Sudan clustered with ''P. l. leo'' in mtDNA-based phylogenies, but with a high affinity to ''P. l. melanochaita''.<ref name="DeManuel_al.2020">{{cite journal |author1=Manuel, M. d. |author2=Ross, B. |author3=Sandoval-Velasco, M. |author4=Yamaguchi, N. |author5=Vieira, F. G. |author6=Mendoza, M. L. Z. |author7=Liu, S. |author8=Martin, M. D. |author9=Sinding, M.-H. S. |author10=Mak, S. S. T. |author11=Carøe, C. |author12=Liu, S. |author13=Guo, C. |author14=Zheng, J. |author15=Zazula, G. |author16=Baryshnikov, G. |author17=Eizirik, E. |author18=Koepfli, K.-P. |author19=Johnson, W. E. |author20=Antunes, A. |author21=Sicheritz-Ponten, T. |author22=Gopalakrishnan, S. |author23=Larson, G. |author24=Yang, H. |author25=O’Brien, S. J. |author26=Hansen, A. J. |author27=Zhang, G. |author28=Marques-Bonet, T. |author29=Gilbert, M. T. P. |name-list-style=amp |title=The evolutionary history of extinct and living lions |journal=[[Proceedings of the National Academy of Sciences of the United States of America]] |doi=10.1073/pnas.1919423117 |date=2020|volume=117 |issue=20 |pages=10927–10934 |pmid=32366643 |pmc=7245068 |bibcode=2020PNAS..11710927D |doi-access=free}}</ref>
-* [http://www.barbarylion.com Preservation Station - Barbary Lions], the Barbary Lion preservation site.
-* [http://www.zionwildlifegardens.co.nz Zion WildLife Gardens - NZ], dedicated to the preservation of the Big Cats.
-* [http://www.petermaas.nl/extinct/speciesinfo/barbarylion.htm The Extinction Website - Species Info - Barbary Lion]
-* [http://www.springerlink.com/content/f638w0l4n181l0r7/ Urgent call for further breeding of the relic zoo population of the critically endangered Barbary lion (Panthera leo leo Linnaeus 1758)]
-* [http://www.arkive.org/species/GES/mammals/Panthera_leo/more_info.html Lion (Panthera leo) from “ARKive images of life on Earth” website]
-* [http://animaldiversity.ummz.umich.edu/site/accounts/information/Panthera_leo.html Panthera leo (lion) from “Animal Diversity Web”]
+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.<ref name="Bertola_al2016">{{cite journal |author1=Bertola, L. D. |author2=Jongbloed, H. |author3=Van Der Gaag, K. J. |author4=De Knijff, P. |author5=Yamaguchi, N. |author6=Hooghiemstra, H. |author7=Bauer, H. |author8=Henschel, P. |author9=White, P. A. |author10=Driscoll, C. A. |author11=Tende, T. |name-list-style=amp |year=2016 |title=Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (''Panthera leo'') |journal=Scientific Reports |volume=6 |page=30807 |doi=10.1038/srep30807|pmid=27488946 |pmc=4973251 |bibcode=2016NatSR...630807B}}</ref>
-[[Category:Extinct animals of Africa]]
-[[Category:Lions]]
-[[Category:Species extinct in the wild]]
-[[Category:Mammals of Africa]]
-[[Category:Fauna of Algeria]]
-[[Category:Fauna of Egypt]]
+==Former distribution and habitat==
-[[de:Berberlöwe]]
+[[File:'n Berberleeu in 1925.png|thumb|The last photograph of a wild lion in the Atlas Mountains, taken by [[Marcelin Flandrin]] in 1925<ref name="Black et al.2013"/>]]
-[[es:Panthera leo leo]]
+[[File:Atlas Van der Hagen - Barbary Coast.jpeg|thumb|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.]]
-[[fr:Lion de l'Atlas]]
+Fossils of the Barbary lion dating to between 100,000 and 110,000 years were found in the cave of Bizmoune near [[Essaouira]].<ref>{{Cite web |date=2022 |title=– بلاغ صحفي بقايا عظمية لأسد الأطلس – بلاغ صحفي |url=https://insap.ac.ma/?p=32654 |access-date=2022-12-09 |website=Insap |language=ar}}</ref><ref>{{Cite web |date=2022 |title=Près d'Essaouira, des chercheurs de l'INSAP découvrent des vestiges du lion de l'Atlas |url=https://ledesk.ma/encontinu/pres-dessaouira-des-chercheurs-de-linsap-decouvrent-des-vestiges-du-lion-de-latlas/ |access-date=2022-12-09 |website=Le Desk |language=fr-FR}}</ref>
-[[it:Panthera leo leo]]
+The Barbary lion lived in the mountains and deserts of the [[Maghreb]] of [[North Africa]] from [[Morocco]] to [[Egypt]]. It was eradicated following the spread of [[firearms]] and [[Bounty (reward)|bounties]] for shooting lions.<ref name=Pease1913>{{cite book |last=Pease |first=A. E. |author-link=Alfred Edward Pease |title=The Book of the Lion |publisher=[[John Murray (publishing house)|John Murray]] |chapter=The Distribution of Lions |pages=109−147 |location=London |year=1913 |chapter-url=https://archive.org/stream/bookoflion1913alfr#page/112/mode/2up}}</ref>
-[[he:אריה אפריקני]]
+Today, it is [[locally extinct]] in this region.<ref name=iucn>{{cite iucn |author=Bauer, H. |author2=Packer, C. |author3=Funston, P.F. |author4=Henschel, P. |author5=Nowell, K. |year=2016 |errata=2017 |title=''Panthera leo'' |page=e.T15951A115130419 |doi=10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en}}</ref>
-[[hu:Berber oroszlán]]
+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 inhabited [[Mediterranean forests, woodlands, and scrub]]. The westernmost sighting of a Barbary lion reportedly occurred in the [[Anti-Atlas]] in western Morocco. It ranged from the [[Atlas Mountains]] and the [[Rif]] in Morocco, the [[Ksour Range|Ksour]] and [[Amour Range]]s in Algeria to the [[Aurès Mountains]] in [[Tunisia]].<ref name="Black et al.2013">{{cite journal |last1=Black |first1=S. A. |last2=Fellous |first2=A. |last3=Yamaguchi |first3=N. |last4=Roberts |first4=D. L. |year=2013 |title=Examining the Extinction of the Barbary Lion and Its Implications for Felid Conservation |journal=PLOS ONE |volume=8 |issue=4 |page=e60174 |doi=10.1371/journal.pone.0060174 |pmid=23573239 |pmc=3616087|bibcode=2013PLoSO...860174B |doi-access=free}}</ref>
-[[ja:バーバリライオン]]
+In Algeria, the Barbary lion was sighted in the forested hills and mountains between [[Ouarsenis]] in the west to the [[Chelif River]] plains in the north and the Pic de [[Taza National Park|Taza]] in the east. It inhabited the forests and wooded hills of the [[Constantine Province]] southward into the Aurès Mountains.<ref name=Pease1913/>
-[[ko:바바리사자]]
-[[pt:Leão-do-atlas]]
+In the 1830s, lions may have already been eliminated along the coast of the [[Mediterranean Sea]] and near human settlements.<ref>{{cite book |author=Jardine, W. |year=1834 |title=The Naturalist's Library. Mammalia Vol. II: the Natural History of Felinae |chapter=The Lion |chapter-url=https://archive.org/stream/naturalistslibra16jardrich#page/n85/mode/2up |pages=87−123 |publisher=W. H. Lizars |location=Edinburgh}}</ref>
-[[fi:Atlasleijona]]
+In [[Libya]], the Barbary lion persisted along the coast until the beginning of the 18th century, and was extirpated in Tunisia by 1890.<ref>{{cite book |author=Guggisberg, C. A. W. |year=1961 |title=Simba: the life of the lion |publisher=Howard Timmins |location=Cape Town}}</ref> By the mid-19th century, the Barbary lion population had massively declined, since [[Bounty (reward)|bounties]] were paid for shooting lions. The cedar forests of [[Chelia]] and neighbouring mountains in Algeria harboured lions until about 1884.<ref name=Pease1913/> The Barbary lion disappeared in the [[Bône (département)|Bône]] region by 1890, in the [[Khroumire]] and [[Souk Ahras Province|Souk Ahras]] regions by 1891, and in [[Batna Province]] by 1893.<ref>{{cite book |author=Joleaud, L. |year=1936 |chapter=Zoogéographie mammalogique |page=174 |title=Étude géologique de la région de Bône et de La Calle |publisher=Bulletin du Service de la Carte Géologique de l’Algérie |location=Alger}}</ref>
+The last recorded shooting of a wild Barbary lion took place in 1942 near [[Tizi n'Tichka]] in the Moroccan part of the Atlas Mountains. A small [[Relict (biology)|relict]] population may have survived in remote montane areas into the early 1960s.<ref name="Black et al.2013"/> The last known sighting of a lion in [[Algeria]] occurred in 1956 in [[Beni Ourtilane District]].<ref name="Black et al.2013"/>
+Historical accounts indicate that in Egypt, lions occurred in the [[Sinai Peninsula]], along the [[Nile]], in the [[Eastern Desert|Eastern]] and [[Western Desert (Egypt)|Western Desert]]s, in the region of [[Wadi El Natrun]] and along the maritime coast of the Mediterranean.<ref>{{cite book |author=Planhol, X. |year=2004 |title=Le Paysage Animal. L'homme et la Grande Faune: Une Zoogéographie Historique |location=Paris |publisher=Fayard}}</ref> In the 14th century [[Before Christ|BC]], [[Thutmose IV]] hunted lions in the hills near [[Memphis, Egypt|Memphis]].<ref>{{cite book |author=Wilkinson, J. G. |year=1878 |title=The manners and customs of the ancient Egyptians. Volume III |edition=revised |publisher=Dodd, Mead and Co. |location=New York}}</ref> The growth of civilizations along the Nile and in the Sinai Peninsula by the beginning of the second millennium BC and [[desertification]] contributed to isolating lion populations in North Africa.<ref name=Barnett_al2008/>
+==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.<ref name=Pease1913/> 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.<ref name="Black et al.2013"/>
+When [[Barbary stag]] (''Cervus elaphus barbarus'') and [[gazelle]]s became scarce in the Atlas Mountains, lions preyed on herds of livestock that were carefully tended.<ref name="Bryden1">{{cite book |author=Johnston, H. H. |year=1899 |chapter=The lion in Tunisia |pages=562–564 |title=Great and small game of Africa |location=London |chapter-url=https://archive.org/stream/greatsmallgameof00majo#page/562/mode/2up |editor1-last=Bryden |editor1-first=H. A. |publisher=Rowland Ward Ltd.}}</ref> They also preyed on [[wild boar]] (''Sus scrofa'').<ref name="Bryden2">{{cite book |author=Pease, A. E. |year=1899 |title=Great and small game of Africa |editor1-last=Bryden |editor1-first=H. A. |publisher=Rowland Ward Ltd. |location=London |chapter=The lion in Algeria |pages=564–568 |chapter-url=https://archive.org/stream/greatsmallgameof00majo#page/564/mode/2up}}</ref>
+[[Sympatry|Sympatric]] predators in this region included the [[African leopard]] (''P. pardus pardus'') and [[Atlas bear]] (''Ursus arctos crowtheri'').<ref name=CAP/><ref name="Bryden3">{{cite book |author=Johnston, H. H. |year=1899 |chapter=African bear |title=Great and small game of Africa |editor1-last=Bryden |editor1-first=H. A. |publisher=Rowland Ward Ltd. |location=London |pages=607–608 |chapter-url=https://archive.org/stream/greatsmallgameof00majo#page/607/mode/2up}}</ref>
+==In captivity==
+{{See also|Damnatio ad bestias}}
+The lions kept in the [[menagerie]] at the [[Tower of London]] in the [[Middle Ages]] were Barbary lions, as shown by [[DNA]] testing on two well-preserved skulls excavated at the Tower in 1936 and 1937. The skulls were [[radiocarbon]]-dated to around 1280–1385 and 1420−1480.<ref name=Barnett_al2008>{{cite journal |author=Barnett, R. |author2=Yamaguchi, N. |author3=Shapiro, B. |author4=Sabin, R. |year=2008 |title=Ancient DNA analysis indicates the first English lions originated from North Africa |url=http://dpc.uba.uva.nl/ctz/vol77/nr01/art02 |journal=Contributions to Zoology |volume=77 |issue=1 |pages=7–16 |doi=10.1163/18759866-07701002 |s2cid=7925316 |access-date=2013-11-28 |archive-date=2013-12-03 |archive-url=https://web.archive.org/web/20131203013823/http://dpc.uba.uva.nl/ctz/vol77/nr01/art02 |url-status=dead }}</ref>
+In the 19th century and the early 20th century, lions were often kept in hotels and [[circus]] [[menagerie]]s. In 1835, the lions in the Tower of London were transferred to improved enclosures at the [[London Zoo]] on the orders of the [[Arthur Wellesley, 1st Duke of Wellington|Duke of Wellington]].<ref name=Edwards1996>{{cite book |author=Edwards, J. |year=1996 |title=London Zoo from Old Photographs 1852–1914 |publisher=John Edwards |location=London}}</ref>
+The lions in the [[Rabat Zoo]] exhibited characteristics thought typical for the Barbary lion.<ref name="NowellJacksonImages">{{cite book |author1=Nowell, K. |author2=Jackson, P. |year=1996 |title=Wild Cats: status survey and conservation action plan |publisher=IUCN/SSC Cat Specialist Group, Gland, Switzerland |chapter=Wild Cats of Africa |pages=Plate I |chapter-url=http://carnivoractionplans1.free.fr/wildcats.pdf}}</ref> Nobles and [[Berber 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 in [[Casablanca]], with the rest being shifted to [[Meknès]]. 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 in [[Temara]] near [[Rabat]].<ref name=Yamaguchi2002/> Results of a [[mtDNA]] research revealed in 2006 that a lion kept in the German [[:de: Zoo Neuwied|Zoo Neuwied]] originated from this collection and is very likely a descendant of a Barbary lion.<ref name=Burger2006>{{Cite journal |author=Burger, J. |author2=Hemmer, H. |title=Urgent call for further breeding of the relic zoo population of the critically endangered Barbary lion (''Panthera leo leo'' Linnaeus 1758) |year=2006 |doi=10.1007/s10344-005-0009-z |journal=European Journal of Wildlife Research |volume=52 |issue=1 |pages=54–58 |s2cid=30407194 |url=http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Burger%20Hemmer%202006.pdf |access-date=2007-04-04 |archive-url=https://web.archive.org/web/20070703152826/http://www.uni-mainz.de/FB/Biologie/Anthropologie/MolA/Download/Burger%20Hemmer%202006.pdf |archive-date=2007-07-03 |url-status=dead}}</ref>
+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.<ref name=BarnettYamaguchi2006/> Several researchers and zoos supported the development of a studbook of lions directly descended from the King of Morocco's collection.<ref name=Black2010/>
+At the beginning of the 21st century, the [[Addis Ababa Zoo]] kept 16 adult lions. With their dark, brown manes extending through the front legs, they looked like Barbary or [[Cape lion]]s. Their ancestors were caught in southwestern Ethiopia as part of a zoological collection for Emperor [[Haile Selassie of Ethiopia]].<ref>{{cite journal |last=Tefera |first=M. |title=Phenotypic and reproductive characteristics of lions (''Panthera leo'') at Addis Ababa Zoo |journal=Biodiversity and Conservation |volume=12 |issue=8 |pages=1629–1639 |date=2003 |doi=10.1023/A:1023641629538 |s2cid=24543070}}</ref>
+Since 2005, three Barbary lions were kept in [[Belfast Zoo]] obtained from [[Port Lympne Wild Animal Park]], and a new Barbary lion enclosure was opened in 2023.<ref>{{Cite web |title=Lord Mayor opens New Kingdom at Belfast Zoo! |url=https://www.belfastcity.gov.uk/zoo/news-and-events/news/lord-mayor-opens-new-kingdom-at-belfast-zoo! |access-date=2023-09-07 |website=Belfast Zoo}}</ref>
+==Cultural significance==
+{{Further|Cultural depictions of lions|Lion#Cultural significance}}
+[[File:Eugène Delacroix - Lion Hunt in Morocco - WGA6228.jpg|thumb|Painting of a lion hunt in Morocco by [[Eugène Delacroix]], 1855, in the [[Hermitage Museum]]]]
+The lion also appeared frequently in early [[Egyptian art]] and [[Egyptian literature|literature]].<ref name=Porter1894>{{cite book |title=Wild beasts; a study of the characters and habits of the elephant, lion, leopard, panther, jaguar, tiger, puma, wolf, and grizzly bear |author=Porter, J. H. |year=1894 |chapter=The Lion |pages=76–134 |chapter-url=https://archive.org/stream/wildbeastsstud00port#page/76 |publisher=C. Scribner's sons |location=New York}}</ref> Statues and statuettes of lions found at [[Hierakonpolis]] and [[Koptos]] in [[Upper Egypt]] date to the [[Early Dynastic Period (Egypt)|Early Dynastic Period]].<ref>{{cite book |author=Adams, B. |year=1992 |chapter=Two more lions from Upper Egypt: Hierakonpolis and Koptos |title=The Followers of Horus. Studies Dedicated to Michael Allen Hoffman |publisher=Oxbow Press |location=Oxford |pages=69–76 |editor=Friedmann, R. |editor2=Adams, B.}}</ref> The early Egyptian deity [[Mehit]] was depicted with a lion head.<ref>{{cite book |author=Wilkinson, T. A. H. |year=1999 |title=Early Dynastic Egypt |publisher=Routledge |location=London, New York |isbn=0415260116}}</ref> In [[Ancient Egypt]], the lion-headed deity [[Sekhmet]] was venerated as protector of the country.<ref>{{cite book |author=Engels, D.W. |year=2001 |title=Classical Cats. The Rise and Fall of the Sacred Cat |publisher=Routledge |location=London, New York |isbn=0415261627 |url-access=registration |url=https://archive.org/details/classicalcats00dona }}</ref> She represented destructive power, but was also regarded as protector against famine and disease. Lion-headed figures and amulets were excavated in tombs in the [[Aegean islands]] of [[Crete]], [[Euboea]], [[Rhodes]], [[Paros]] and [[Chios]]. They are associated with Sekhmet and date to the early [[Iron Age]] between the 9th and 6th centuries BC.<ref>{{cite book |author=Apostola, E. |year=2014 |chapter=Cross-cultural Relations between Egypt and Greece during the Early Iron Age: Representations of Egyptian Lion-Headed Deities in the Aegean |title=Current Research in Egyptology: Proceedings of the Fifteenth Annual Symposium |publisher=Oxbow Books |location=Oxford |pages=100–112 |editor=Pinarello, M.S. |editor2=Yoo, J. |editor3=Lundock, J. |editor4=Walsh, C.}}</ref> The remains of seven mostly subadult lions were excavated at the necropolis [[Umm El Qa'ab]] in a tomb of [[Hor-Aha]], dated to the 31st century BC.<ref>{{cite book |author=Boessneck, J., von den Driesch, A. |chapter=Die Tierknochenfunde |year=1990 |title=Umm el-Qaab: Nachuntersuchungen im frühzeitlichen Königsfriedhof. 3./4. Vorbericht. Abteilung Kairo |publisher=46. Mitteilungen des Deutschen Archäologischen Instituts |location=Berlin | editor=Dreyer, G.}}</ref> In 2001, the skeleton of a [[Mummy|mummified]] lion was found in the tomb of [[Maia (nurse)|Maïa]] in a [[necropolis]] dedicated to [[Tutankhamun]] at [[Saqqara]].<ref name=Calou_al2004>{{cite journal |title=A lion found in the Egyptian tomb of Maïa |last1=Callou |first1=C. |last2=Samzun |first2=A. |last3=Zivie |first3=A. |journal=Nature |date=2004 |volume=427 |issue=6971 |pages=211–212 |doi=10.1038/427211a|pmid=14724625 |s2cid=4422033 }}</ref> It had probably lived and died in the [[Ptolemaic Kingdom|Ptolemaic]] period, showed signs of malnutrition and had probably lived in captivity for many years.<ref>{{cite journal |title=Le lion du Bubasteion à Saqqara (Égypte) |last1=Samzun |first1=A. |last2=Hennet |first2=P. |last3=Lichtenberg |first3=R. |last4=Callou |first4=C. |last5=Zivie |first5=A. |journal=Anthropozoologica |date=2011 |volume=46 |issue=2 |pages=63–84 |doi=10.5252/az2011n2a4|s2cid=129186181 |url=https://hal-mnhn.archives-ouvertes.fr/mnhn-02103375/file/az2011n2a4.pdf }}</ref>
+The Barbary lion is a symbol in Nubian culture and was often depicted in art and architecture. Nubian deities, such as [[Amun]], [[Amesemi]], [[Apedemak]], [[Arensnuphis]], [[Hathor]], [[Bastet]], [[Dedun]], Mehit, [[Menhit]], and [[Sebiumeker]], were depicted as lion protectors in [[Kushite religion]].<ref>{{cite book |author=Fisher, M. M. |author2=Lacovara, P. |author3=Ikram, S. |author4=D'Auria, S. |name-list-style=amp |author5=Yellin, J. W. |author6=Knoblauch, C. |year=2012 |title=Ancient Nubia: African Kingdoms on the Nile |location=Cairo, Egypt |publisher=American University in Cairo Press |isbn=9789774164781 |url=https://books.google.com/books?id=Lu6nZwEACAAJ}}</ref>
+In [[Roman North Africa]], lions were regularly captured by experienced hunters for [[venatio]] spectacles in [[amphitheatre]]s.<ref name="Bryden2"/><ref>{{cite book |title=Venationes Africanae: Hunting spectacles in Roman North Africa: cultural significance and social function |chapter=Chapter 2: Procuring beasts for hunting spectacles |pages=67–98 |publisher=Amsterdam School of Historical Studies |author=Sparreboom, A. |year=2016 |location=Amsterdam |isbn=9789463320238}}</ref>
+The [[Morocco 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.<ref>[https://www.africanews.com/2021/02/09/football-planet-atlas-lions-of-morocco-win-second-chan-title-in-a-row/ Atlas Lions of Morocco win second CHAN title in a row], retrieved: February 10th, 2021</ref>
+==See also==
+*{{hlist|Wild cats in Africa: | [[Cheetah]] | [[African golden cat]] | [[Caracal]] | [[Serval]] | [[African wildcat]] | [[Sand cat]] | [[Black-footed cat]]}}
+{{colbegin|colwidth=20em}}
+*[[History of lions in Europe]]
+*[[Holocene extinction]]
+*[[Cats in Ancient Egypt]]
+*[[Lion-baiting]]
+*[[Lake Akfadou]]
+{{colend}}
+==References==
+{{Reflist}}
+==External links==
+*{{cite news |url=https://www.apollo-magazine.com/i-find-myself-making-growling-noises-while-im-painting-an-interview-with-walton-ford/ |title='I find myself making growling noises while I'm painting' – an interview with Walton Ford, who painted Barbary lions |newspaper=Apollo Magazine |author=Marks, T. |year=2018}}
+*{{cite news |url=https://en.yabiladi.com/articles/details/72562/history-when-london-s-very-first.html |title=History : When London's very first zoo housed Morocco's Atlas Lions |author=Babas, L. |year=2018 |newspaper=Yabiladi}}
+*{{cite AV media |url=https://www.youtube.com/watch?v=pyNQDMEw6Iw  |archive-url=https://ghostarchive.org/varchive/youtube/20211222/pyNQDMEw6Iw |archive-date=2021-12-22 |url-status=live|type=video |title=Moroccan 'Atlas' lion at Parc Sindibad, Casablanca}}{{cbignore}}
+*{{cite web |url=https://blogs.kent.ac.uk/barbarylion/2014/07/10/moroccan-royal-lions-in-zoos-today |website=University of Kent Blog |title=Moroccan lions in zoos today |author=Black, S. |year=2014}}
+*{{cite web |url=http://beinglion.com/barbary-lions.php |title=Barbary Lion Information |website=Being Lion}}
+{{Taxonbar|from=Q221094}}
+{{DEFAULTSORT:Barbary Lion}}
+[[Category:Panthera leo leo]]
+[[Category:Mammals described in 1758]]
+[[Category:Taxa named by Carl Linnaeus]]
+[[Category:Extinct mammals of Africa]]
+[[Category:Mammal extinctions since 1500]]
+[[Category:Mammals of North Africa]]