|A male Barbary lion photographed in Algeria by Alfred Edward Pease in 1893.|
|Lioness and cubs, New York Zoo, 1903.|
|Subspecies:||P. l. leo|
|Panthera leo leo
The Barbary lion (Panthera leo leo) is the nominate lion subspecies in North Africa. In Algeria, Egypt, Libya, and Morocco, lions are regionally extinct due to excessive hunting. The last recorded shooting of a wild Barbary lion took place in Morocco, near Tizi n'Tichka in 1942. Small groups of lions may have survived in Algeria until the early 1960s, and in Morocco until the mid-1960s.
Alfred Edward Pease referred to the Barbary lion as the North African lion, and claimed that the population had diminished since the mid-19th century, following the diffusion of firearms and bounties for shooting them. Some authors referred to it as the Berber lion occurring from the Atlas Mountains to Egypt. Since it inhabited the Atlas Mountains, it was also known as the Atlas lion.
Results of morphological and genetic analyses of lions warrant the designation of lion populations in North, West and northern parts of Central Africa to the subspecies P. l. leo, which differ genetically from P. l. melanochaita in Eastern and Southern Africa.
A lion from Constantine, Algeria was the type specimen for the specific name Felis leo used by Linnaeus in 1758. In the 19th century, several African lion specimens from North Africa were described and proposed as subspecies:
- In 1826, the Austrian zoologist Johann Nepomuk Meyer described a lion from the Barbary Coast under the trinomen Felis leo barbaricus.
- In 1843, the French zoologist Henri Marie Ducrotay de Blainville described a male lion from Nubia under the trinomen Felis leo nubicus that had been sent by Antoine Clot from Cairo to Paris and died in the Ménagerie du Jardin des Plantes in 1841.
- In 1891, the German zoologist Theodor Noack described two lion specimens from Somalia and proposed Felis leo somaliensis.
Reginald Innes Pocock subordinated lions to the genus Panthera, when he wrote about the Asiatic lion. In 1951, John Ellerman and Terence Morrison-Scott recognized only two lion subspecies in the Palearctic realm, namely the African Panthera leo leo and the Asiatic lion P. l. persica.
In 2005, the authors of Mammal Species of the World grouped P. l. barbarica, nubica and somaliensis under P. l. leo. In 2016, IUCN Red List assessors subsumed all African lion populations to P. l. leo. 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.
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 1936-1937. The skulls were radiocarbon-dated to 1280-1385 CE and 1420-1480 CE. The growth of civilizations along the Nile and in the Sinai Peninsula by the beginning of the second millennium BC stopped genetic flow by isolating lion populations. Desertification also prevented the Barbary lions from mixing with lions located further south in the continent.
In 2006, mtDNA research revealed that a lion kept in the German Zoo Neuwied originated from the collection of the King of Morocco and is very likely a descendant of a Barbary lion. Five tested samples of lions from the famous collection of the King of Morocco are not maternally Barbary lions. 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. Therefore, many lions in European and American zoos, which are managed without subspecies classification, are in fact partly descendants of Barbary lions.
While the historical Barbary lion was morphologically distinct, its genetic uniqueness remains questionable. Still unresolved is the taxonomic status of surviving lions frequently considered as Barbary lions, including those that originated from the collection of the King of Morocco.
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 lions from the Central African Republic and the northern part of the Democratic Republic of the Congo, and that of a lion from Ethiopia. A phylogeographic analysis of Pleistocene cave lions revealed that a lion sample from Sudan was distinct from lion samples that originated in the northeastern part of the Democratic Republic of the Congo.
In a comprehensive study about the evolution of lions, 357 samples of wild and captive lions from African countries and India were examined. Results indicate that four captive lions from Morocco did not exhibit any unique genetic characteristics, but shared mitochondrial haplotypes H5 and H6 with lions from West and Central Africa. They were all part of a major mtDNA grouping that also included Asiatic samples. Results provide evidence for the hypothesis that this group called 'lineage III' developed in East Africa, and about 118,000 years ago traveled north and west in the first wave of lion expansions. It broke up into haplotypes H5 and H6 within Africa, and then into H7 and H8 in Western Asia. African lions probably constitute a single population that interbred during several waves of migration since the Late Pleistocene.
Barbary lions were described as having greyish tawny coloured fur. Males had very dark and long-haired manes that extended over the shoulder and under the belly. Their elbow tufts were large and the brown tail tuft long. The mane hair was 8 to 22 cm (3.1 to 8.7 in) long. 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 females measure around 2.5 m (8 ft 2 in). Skull size varied from 30.85 to 37.23 cm (12.15 to 14.66 in).
It was often held that the Atlas lion was bigger than other lions. In some historic accounts, the weights of wild males was indicated as being very heavy, and reaching 270 to 300 kg (600 to 660 lb). But the accuracy of the measurements is questionable, and the sample size of captive Barbary lions was too small to conclude that it was the biggest.
Before it became possible to investigate the genetic diversity of lion populations, the colour and size of lions' manes was thought to be a sufficiently distinct morphological characteristic to accord a subspecific status to populations. Mane development varies with age and between individuals from different regions. It is therefore not a sufficient characteristic for subspecific identification. Sub-Saharan African lions kept in cool environments of European and North American zoos usually develop longer manes than wild lions. Barbary lions may have developed long-haired manes, because of temperatures in the Atlas Mountains that are much lower than in other African regions, particularly in winter. Therefore, the size of manes is not regarded as an appropriate evidence for identifying Barbary lions' ancestry. Instead, results of mitochondrial DNA research published in 2006 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 for the identification of Barbary lions surviving in captivity.
Results of a long-term study of East African lions in Serengeti National Park indicate that various factors, such as ambient temperature, nutrition and the level of testosterone, influence the colour and size of lion manes.
Behaviour and ecology
Pease accounted in 1913 that in areas where lions were not very numerous, they were more frequently found in pairs or family parties comprising a lion, lioness and one or two cubs. He several times came across two old lions and a lioness living and hunting together. Observations of wild Barbary lions made between 1839 and 1942 involved solitary animals, pairs and family units. Analysis of these historical records suggests that Barbary lions retained living in prides even when under increasing persecution during the last decades, especially 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.
Extinction in the wild
Barbary lions inhabited the range countries of the Atlas Mountains including the Barbary Coast. Jardine remarked in 1834 that at the time lions may have already been eliminated from the coastlines, marking the border to human settlements. In Algeria, they lived in the forest-clad hills and mountains between Ouarsenis in the west, the Pic de Taza in the east, and the plains of the Chelif River in the north. There were also many lions among the forests and wooded hills of the Constantine Province eastwards into Tunisia and south into the Aurès Mountains. By the middle of the 19th century, their numbers had been greatly diminished. The cedar forests of Chelia and neighbouring mountains harboured lions until about 1884. The last survivors in Tunisia were extirpated by 1890.
In the 1970s, Barbary lions were assumed to have been extirpated in the wilderness by the early 20th century. However, a comprehensive review of hunting and sighting records indicates that a lion was shot in the Moroccan part of the Atlas Mountains in 1942. Additionally, lions were sighted in Morocco and Algeria into the 1950s, and small remnant populations may have survived in remote areas into the early 1960s.
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Historically, lions were offered in lieu of taxes and as gifts to royal families of Morocco and Ethiopia. The rulers of Morocco kept these 'royal lions' through war and insurrection, splitting the collection between zoos when the royal family went briefly into exile. After a respiratory disease nearly wiped out the royal lions in the late 1960s, the current ruler established enclosures in Temara near Rabat, Morocco, to house the lions and improve their quality of life. There are currently a small number of 'royal lions' that have the pedigree and physical characteristics to be considered as mostly pure Barbary descendants. Some were returned to the palace when the exiled ruler returned to the throne.
In the 19th century and the early 20th century, lions were often kept in hotels and circus menageries. The lions in the Tower of London were transferred to more humane conditions at the London Zoo in 1835 on the orders of the Duke of Wellington. One famous Barbary lion named "Sultan" was kept in the London Zoo in 1896.
Emperor Haile Selassie I of Ethiopia had a collection of lions at Addis Abeba's zoo. They looked like Barbary or Cape lions, because of their dark, brown manes that extended over the chest through the front legs.
In March 2010, two lion cubs were moved to The Texas Zoo in Victoria, Texas, where efforts were made to preserve the species under the WildLink International conservation programme. Whether the cubs are of Barbary lion descent is not yet determined. In 2011, the Port Lympne Animal Park in Kent received a Barbary lioness as a mate for the resident male. As of June 2016, Wisconsin Big Cat Rescue in Rock Springs, Wisconsin, has two female lions born in 2001 that have been shown by DNA testing to be Atlas lions. The Living Treasures Wild Animal Park in New Castle, Pennsylvania, claims to keep a pair of Barbary lions in the park's collection. The Zoo des Sables d'Olonne, Vendee, France, also claims to have a male and female Atlas lion.
More recently a number of researchers and zoos have supported the development of a studbook of lions directly descended from the King of Morocco's collection. This work has been conducted on the precautionary principle that this subpopulation of animals may hold unique Barbary lion genes as well as demonstrating the morphology of the ancestral Barbary lion.
The former popularity of the Barbary lion as a zoo animal provides the only hope to ever see it again in the wild. Many zoos provide mating programmes. After years of research into the science of the Barbary lion and stories of surviving examples, WildLink International, in collaboration with Oxford University, launched its ambitious International Barbary Lion Project. Oxford used DNA techniques to identify the DNA 'fingerprint' of the Barbary lion subspecies. Researchers took bone samples from remains of Barbary lions in museums across Europe. These samples were returned to Oxford University, where the science team extracted the DNA sequence to identify the Barbary as a separate subspecies.
Although the North African lion is certainly extinct in the wild, WildLink International looked to identify a handful of lions in captivity around the world that may be descended from the original Barbary lion. These descendants were to be tested against the DNA fingerprint, and the degree of any hybridization (from crossbreeding) could then be determined. The best candidates were to then enter a selective breeding programme slated to 'breed back' the Barbary lion. The final phase of the project intended to see the lions released into a national park in the Atlas Mountains of Morocco.
The lion is called by various names, depending on the region of its occurrence, such as Mauretania and Numidia. Omar Mukhtar, the Libyan revolutionary was called the Asad aṣ-Ṣaḥrā’ (Arabic: أَسَـد الـصَّـحْـرَاء, "Lion of the Desert"). The Morocco national football team is nicknamed "Atlas Lions."
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