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The North African Barbary lion and the Atlas Lion Project

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... P. leo was originally ranged across Africa (except the central Saharan regions), the Middle East, south-eastern Europe (Greece and Bulgaria), central Asia (specifically the Caucasus) and in north-western regions of South Asia (Pakistan and India) [4,5]. The socalled 'Barbary lion' from North Africa was distinctive in being characterised by a massive mane that can be coloured from light to dark and can extend from the head, neck and down to the belly and under the belly to the elbows [6], as well as other morphological features [7]. ...
... In the 1990s, the IUCN Wild Cat Status Survey and Action Plan indicated that some lions in captivity could be 'Moroccan Royal lions', while the wild population was extinct [18,20]. The best provenance for likely 'Moroccan Royal lions' are animals derived from the Moroccan royal collection, originally established from animals captured in the wild by local tribes as tributes to the Sultan, perhaps up until the early 20 th century [7]. ...
... Despite the efforts of genetic studies, the evolutionary importance of lions from the Moroccan collection has not yet been fully established, but prudence suggests that it is still necessary to establish an official scientific breeding programme [15]. Yamaguchi and Haddane [7] reported that there are currently only a few lion specimens that can be considered predominantly pure putative representatives of Barbary lions. Black [32] pointed out that there are several lions in European zoos that can be considered partial 'Moroccan Royal lions' but cannot necessarily be described as pure-blooded Barbary lions. ...
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This study evaluates the diversity of the so-called ‘Moroccan Royal lions’ using genealogical information. Lions are no longer extant in North Africa, but the previous wild population was an important element of the now-recognised northern subspecies ( Panthera leo leo ) that ranged across West Africa, North Africa and the Middle East into India. The remaining captive population of ‘Moroccan Royal lions’ seems to be significantly endangered by the loss of diversity due to the effective population size decrease. The pedigree file of this captive lion population consisted of 454 individuals, while the reference population included 98 animals (47 males and 51 females). The completeness of the pedigree data significantly decreased with an increasing number of generations. The highest percentage of pedigree completeness (over 70%) was achieved in the first generation of the reference population. Pedigree-based parameters derived from the common ancestor and gene origin were used to estimate the state of diversity. In the reference population, the average inbreeding coefficient was 2.14%, while the individual increase in inbreeding over generations was 2.31%. Overall, the reference population showed lower average inbreeding and average relatedness compared with the pedigree file. The number of founders (47), the effective number of founders (24) and the effective number of ancestors (22) were estimated in the reference population. The effective population size of 14.02 individuals confirms the critically endangered status of the population and rapid loss of diversity in the future. Thus, continuous monitoring of the genetic diversity of the ‘Moroccan Royal lion’ group is required, especially for long-term conservation management purposes, as it would be an important captive group should further DNA studies establish an affinity to P . leo leo .
... Deprived of its main habitat and prey, the Barbary lion began to feed on domestic livestock, which thus contributed to its persecution. The subsequent introduction of firearms significantly accelerated its demographic decline (Guggisberg 1963): in Algeria, lions were numerous enough for a bounty to be issued by the French colonial government (see Yamaguchi and Haddane 2002). ...
... In fact, for centuries, lion cubs were offered by tribes from the Atlas mountains as tributes. In the late 1960s, to improve life for the lions, a new enclosure (which in 1973 will become the Rabat zoo) was built in Tamara (Yamaguchi and Haddane 2002). ...
... In Tunisia the last lion was shot in 1891 and rumors about the presence of the species in the Khmir mountains and near Feriana continued until the early 1900s (Guggisberg 1963). In Algeria, the last lion of the Saharan Atlas was killed around 1920 (Yamaguchi and Haddane 2002). ...
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The vanishing population of big cats is a global problem difficult to solve. In modern era several subspecies and populations of felids of the genus Panthera have become extinct, or locally extinct. The review of their story seems to show a common trend: a decrease of prey due to hunting and habitat destruction that leads to the decrease of predators, which switch to livestock creating conflicts with human. In many cases the micro populations of big cats are very difficult to detect, surviving for several years after their official extinction without it being possible to implement conservation strategies. To be effective, conservation measures must be as interdisciplinary as possible and include the active involvement of the locals.
... The colloquially named "Barbary" lion of North Africa was familiar to historical writers in Europe due to the proximity of wild populations. The transport of wild lions to gladiatorial events of the Roman period and later menageries of medieval Europe imprinted the Barbary lion in popular culture, art, and literature [15]. The morphology of those animals, the shaggy mane and forequarters, became the model for lion images in mosaics, heraldry, pottery, paintings, and sculpture. ...
... The morphology of those animals, the shaggy mane and forequarters, became the model for lion images in mosaics, heraldry, pottery, paintings, and sculpture. North African lions are intriguing to scientists, natural historians, and conservationists being perceived through morphology and behavioural ecology as the most distinctive of all lion populations [15,16], living in temperate, forested montane habitats of North Africa's Maghreb region [2]. ...
... Whilst the habitat in the Maghreb is now thought to be largely unable to support the needs of lions, leopards still appear to survive in the region [18]. The interest in a potential reintroduction of lions to North Africa carries a broad conservation [16,19,20] and scientific [8,17] and cultural significance [19,21], and the topic regularly surfaces both within conservation circles and the wider media [15,[22][23][24][25]. ...
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The lions of North Africa were unique in ecological terms as well as from a human cultural perspective and were the definitive lions of Roman and Medieval Europe. Labelled " Barbary " lions, they were once numerous in North Africa but were exterminated by the mid-20th century. Despite subsequent degeneration of the Atlas Mountain ecosystem through human pressures, the feasibility of lion reintroduction has been debated since the 1970s. Research on the long-established captive lion collection traditionally kept by the sultans and kings of Morocco has enabled selective breeding coordinated across Moroccan and European zoos involving a significant number of animals. Molecular genetic research has recently provided insights into lion phylogeny which, despite previous suggestions that all lions share recent common ancestry, now indicates clear distinctions between lions in North, West, and Central Africa, the Middle East, and India versus those in Southern and Eastern Africa. A review of the evolutionary relevance of North African lions highlights the important challenges and opportunities in understanding relationships between Moroccan lions, extinct North African lions, and extant lion populations in India and West and Central Africa and the potential role for lions in ecosystem recovery in those regions.
... Most likely due to the expansion of human populations, lions disappeared from Greece by c. 2000 years ago, and from Palestine around the time of the Crusades c. 11 th -12 th century. In the past 200 years, lions have disappeared from many parts of their former range, including the southern part of South Africa (by 1870), Turkey (1870), Tunisia (1891), Iraq (1918), Iran (1942), Morocco (1942) and Algeria (by 1960) (Yamaguchi and Haddane 2002; Patterson, 2004; K. Difalla, personal communication). Among the now extinct populations two are particularly famous: the North African Barbary lion P. l. leo (Linnaeus, 1758) and the South African Cape lion P. l. melanochaita (Smith, 1842) (Mazák 1975; Nowell and Jackson 1996). ...
... Most likely due to the expansion of human populations, lions disappeared from Greece by c. 2000 years ago, and from Palestine around the time of the Crusades c. 11 th -12 th century. In the past 200 years, lions have disappeared from many parts of their former range, including the southern part of South Africa (by 1870), Turkey (1870), Tunisia (1891), Iraq (1918), Iran (1942), Morocco (1942) and Algeria (by 1960) (Yamaguchi and Haddane 2002; Patterson, 2004; K. Difalla, personal communication). Among the now extinct populations two are particularly famous: the North African Barbary lion P. l. leo (Linnaeus, 1758 ) and the South African Cape lion P. l. melanochaita (Smith, 1842) (Mazák 1975; Nowell and Jackson 1996). ...
... However, it is now known that size and colour of a lion's mane are influenced by various factors , including ambient temperature, animal's age and testosterone level (Kays and Patterson, 2002; West and Packer, 2002; Patterson et al., 2006 ). In particular , the Asiatic lion from India, which is characterised by a very small mane in its natural habitat, grows an enormous mane with its greater part being dark or even black, which extends behind the shoulders and covers the belly, in European zoos (Yamaguchi and Haddane, 2002; Yamaguchi unpublished, seeFig. 3). ...
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In natural history collections throughout Europe, there are many old lion specimens of unknown origin. If these specimens can be shown to have originated from now-extinct populations their value would significantly increase, as would the value of the collections. Recently, a 200-year old mounted skeleton in the Zoological Museum Amsterdam has been identified as the extinct Cape lion Panthera leo melanochaita (Smith, 1842), based primarily on morphological information inferred from a painting of this specimen while it was still alive. To test this hypothesis, we used ancient DNA (aDNA) techniques to extract and sequence mitochondrial DNA (mtDNA) from this specimen, and compared the genetic results with previously published lion mtDNA sequences. Our results show that the specimen is not a Cape lion, but that it instead possesses the mtDNA haplotype of the Asiatic lions P. l. persica (Meyer, 1826) from India. This Indian origin hypothesis is further supported by an investigation of its cranial morphology. As the amount of genetic information available for lions increases, in particular data from across their historic distribution, the potential for aDNA techniques to identify the origins of previously unassigned museum specimens continues to grow.
... The restoration of the extinct North African Barbary lion has attracted the attention of conservationists both inside and outside North Africa [19,31,67,68]. Although circumstantial evidence suggested that the Barbary lion could have survived in captivity [67,68], the most likely descendants of wild Barbary lions from the Moroccan Royal Menagerie do not appear to be (maternally) Barbary [19, this study]. ...
... The restoration of the extinct North African Barbary lion has attracted the attention of conservationists both inside and outside North Africa [19,31,67,68]. Although circumstantial evidence suggested that the Barbary lion could have survived in captivity [67,68], the most likely descendants of wild Barbary lions from the Moroccan Royal Menagerie do not appear to be (maternally) Barbary [19, this study]. However, there is a close mitochondrial relationship between the Barbary lion and the extant Indian lion, and this has been tentatively (but independently) supported by non-molecular studies [26,30]. ...
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Article
Understanding the demographic history of a population is critical to conservation and to our broader understanding of evolutionary processes. For many tropical large mammals, however, this aim is confounded by the absence of fossil material and by the misleading signal obtained from genetic data of recently fragmented and isolated populations. This is particularly true for the lion which as a consequence of millennia of human persecution, has large gaps in its natural distribution and several recently extinct populations. We sequenced mitochondrial DNA from museum-preserved individuals, including the extinct Barbary lion (Panthera leo leo) and Iranian lion (P. l. persica), as well as lions from West and Central Africa. We added these to a broader sample of lion sequences, resulting in a data set spanning the historical range of lions. Our Bayesian phylogeographical analyses provide evidence for highly supported, reciprocally monophyletic lion clades. Using a molecular clock, we estimated that recent lion lineages began to diverge in the Late Pleistocene. Expanding equatorial rainforest probably separated lions in South and East Africa from other populations. West African lions then expanded into Central Africa during periods of rainforest contraction. Lastly, we found evidence of two separate incursions into Asia from North Africa, first into India and later into the Middle East. We have identified deep, well-supported splits within the mitochondrial phylogeny of African lions, arguing for recognition of some regional populations as worthy of independent conservation. More morphological and nuclear DNA data are now needed to test these subdivisions.
... Our genome-wide data support this finding, and place Cape lions within the genetic diversity found in South African lions (Fig. 1A). In addition, the restoration of the extinct North African Barbary lion has attracted the attention of conservationists, both inside and outside North Africa (45). Although circumstantial evidence suggested that North African lions could have survived in captivity, the most likely descendants of wild Barbary lions from the Moroccan Royal Menagerie have appeared to be of Central African maternal descent (44). ...
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Lions are one of the world’s most iconic megafauna, yet little is known about their temporal and spatial demographic history and population differentiation. We analyzed a genomic dataset of 20 specimens: two ca. 30,000-y-old cave lions ( Panthera leo spelaea ), 12 historic lions ( Panthera leo leo/Panthera leo melanochaita ) that lived between the 15th and 20th centuries outside the current geographic distribution of lions, and 6 present-day lions from Africa and India. We found that cave and modern lions shared an ancestor ca. 500,000 y ago and that the 2 lineages likely did not hybridize following their divergence. Within modern lions, we found 2 main lineages that diverged ca. 70,000 y ago, with clear evidence of subsequent gene flow. Our data also reveal a nearly complete absence of genetic diversity within Indian lions, probably due to well-documented extremely low effective population sizes in the recent past. Our results contribute toward the understanding of the evolutionary history of lions and complement conservation efforts to protect the diversity of this vulnerable species.
... We obtained 42 additional cyt-b sequences for Recent Panthera leo from GenBank (GU131164-GU131185, AY781195-AY781210, DQ018993-DQ018996), representing nine additional haplotypes, many of them produced by Bertola et al. (2011). Samples obtained in Morocco, from the Rabat Zoo and supposedly descended from Barbary lions, have a complicated history that includes the possibility that they originated in West or Central Africa instead (Yamaguchi and Haddane 2002;Barnett et al. 2006a). The subroutine CLUSTALW was implemented using default parameters to insure correct alignment; no gaps, insertions or deletions were encountered. ...
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Article
Current understanding of genetic variation in lions (Panthera leo) is inadequate to guide many management decisions necessary for conservation of the species. We studied sequence variation in the mitochondrial cytochrome-b (cyt-b) gene of 75 lions and nuclear variation at 11 microsatellite loci of 480 lions across 8 range states (Cameroon, Uganda, Kenya, Zambia, Zimbabwe, South Africa, Botswana, and Namibia) and 13 Lion Conservation Units (LCUs) plus two other unassigned sites (Cameroon and Zimbabwe). A total of 11 cyt-b haplotypes were found, whose variation follows an isolation-by-distance model. In combination with previously known sequences, the haplotypes document the close relationship, derived position, and limited variability of Asian and West and Central African lions relative to other extant lions. Both phylogenetic analyses and substitution networks identify two clades in Eastern and Southern Africa—one restricted to Namibia and South Africa and the other more widespread across the region. However, these analyses are equivocal on which of these is closest to the ancestor of modern lions. Microsatellite analyses showed high levels of variation within and among populations, subdivision among most LCUs, and evidence of isolation by distance. While rates of gene flow are generally low, admixture among lions in northern Botswana, Caprivi Strip (Namibia) and Zambia is apparent from STRUCTURE analyses. Conservation management plans should incorporate information on genetic variability and gene flow in delimiting management units and in guiding translocations of lions to minimize inbreeding and to control problem animals.
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Article
Estimations of species extinction dates are rarely definitive, yet declarations of extinction or extirpation are important as they define when conservation efforts may cease. Erroneous declarations of extinctions not only destabilize conservation efforts but also corrode local community support. Mismatches in perceptions by the scientific and local communities risk undermining sensitive, but important partnerships. We examine observations relating to the decline and extinction of Barbary lions in North Africa. Whilst the extinction predates the era of the scientific conservation movement, the decline is relatively well documented in historical records. Recently unearthed accounts suggest Barbary lions survived later than previously assumed. We use probabilistic methods to estimate a more recent extinction date for the subspecies. The evidence presented for a much later persistence of lions in North Africa, including generations when sightings were nil, suggests caution when considering felid populations as extinct in the wild. The case raises the possibility that captive animals descended from the Moroccan royal collection are closer contemporaries to wild Barbary lions. Furthermore, our results highlight the vulnerability of very small lion populations and the significance of continued conservation of remnant lion populations in Central and West Africa.
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