Barbary Lion - Panthera leo leo
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Image: Barbary lion in a 1898 picture. This image is in the public domain because its copyright has expired. This applies to the European Union, Canada, the United States and those countries with a copyright term of life of the author plus 70 years.
|Subspecies||Panthera leo leo|
|TSEW Status||Extinct (EX), Year assessed: 2010|
|IUCN Status||Vulnerable (VU), Year assessed: 2008 (see Taxonomy)|
|English Name||Barbary Lion, Atlas Lion, Nubian Lion, North African Lion|
|Czech Name||Lev Berberský|
|Finnish Name||Atlasleijona, Barbarileijona|
|French Name||Lion de Barbarie, Lion de l'Atlas|
|Hebrew Name||ברבארי אריה|
|Hungarian Name||Berber Oroszlán|
|Italian Name||Leone di Barberia, Leone Berbero, Leone dell'Atlante|
|Portuguese Name||Leão do Atlas|
|Spanish Name||León del Atlas, León del Berberísco|
Probably due to its close geographical proximity to Europe, a lion from North Africa (Constantine, Algeria) was used as the type specimen when Linnaeus first gave the Latin name Felis leo to the species in 1758 (Harper, 1945).
Felis leo Linnaeus, 1758; Felis leo barbaricus Meyer 1826; Felis leo barbarus Fischer, 1829; Felis leo nubicus Blainville, 1843 (no type locality specified, but stated as ‘‘Nubia’’ by G. M. Allen, 1939); Felis leo nigra Loche, 1858.
Eight recent lions subspecies are commonly recognised, with one in Asia and seven in Africa. However, the validity of the African subspecies is debated and they could probably be grouped into one single African subspecies (Nowell & Jackson, 1996); in fact, the African lion is often considered as monotypic (e.g. Kingdon, 1997; Skinner & Smithers, 1990). The 2008 IUCN Red List of Threatened Species, based on genetic analysis (O'Brien et al. 1987, Dubach et al. 2005), recognised only two subspecies: the African lion (Panthera leo leo) and the Asiatic lion Panthera leo persica (Bauer, Nowell, & Packer 2008). The Asiatic lions (Panthera leo persica) are genetically distinct from the lions of sub-Saharan Africa, although the difference is not large, being smaller than the genetic distance between human racial groups. Based on genetic distance, the Asiatic lion is estimated to have separated from the African population as recently as 100,000 years ago, not long enough for reproductive incompatibilities to have evolved (O’Brien et al. 1987).
Recent genetic and morphological research showed that the Barbary lion (North African lion population) is slightly different from the Asiatic lion (Asian lion population), but clearly distinct from sub-Saharan lions (Hemmer and Burger 2005; Barnett et al. 2006; Burger and Hemmer 2006). Grouping all African lions into one single subspecies, including the North African Barbary lion, would therefore be incorrect. The Barbary lion (Panthera leo leo) can be considered a distinct valid lion subspecies.
There is much confusion between Barbary Lions and other long-maned captive lions. Early authors believed that the seemingly fixed external morphology of Barbary lions (male’s huge mane extending behind shoulders and covering belly) would justify their ‘‘distinct’’ subspecific status and be used to identify them. However, it is now known that the colour and size of a lion’s mane are influenced by various extrinsic factors, including the ambient temperature (West and Packer 2002). Therefore, a heavy mane developed in a cooler place (e.g. European or North American zoos) is an inappropriate marker for identifying Barbary lines, which need to be identified by molecular markers (Yamaguchi and Haddane 2002). (Barnett et al. 2006) Compound that with the many other variables that decide the extent and colour of a lion's mane and you begin to see just how inappropriate the following statement is: "This lion has a long mane and so must be a Barbary" (WildLink International, 2001b).
The possibility that some Barbary lions survive, and they may be the last remnants of a lost subspecies of lion, has become an extremely marketable concept. "It is not uncommon for zoos to advertise [that they possess a Barbary lion] when there is little or no evidence to back up the fact. (Walker, 2009) Lions in captivity today have been bred and cross-bred from lions captured in Africa long ago, with examples from all of these described 'subspecies'. Mixed together, hybridised, most of today's captive lions have a 'soup' of genes from many different lions. (WildLink International, 2001b) First DNA fingerprinting had to be produced, otherwise there was no definitive way to identify a lion as Barbary. In 2006, results of mitochondrial DNA research showed a mtDNA haplotype that is unique to the Barbary lion (Barnett et al. 2006). This can be a good molecular marker for identifying Barbary lions.
The genetic distinctiveness of the historical Barbary lion has not yet been fully established and the question over whether the Royal Lions are true Barbary lions remains unanswered (Dubach et al. 2005; Yamaguchi 2005; Barnett et al. 2006b; Burger and Hemmer 2006; Antunes et al. 2008). (Black et al. 2009)
The Sixth Extinction Website acknowledges that the subspecies is extinct and does not recognize the existence of a genetically pure living specimen of the Barbary lion until genetic testing identifies one. However, the morphology of the Barbary lion is quite distinct (ignoring the mane) and, more importantly, with details of a lion's parentage and background we can identify those most likely to be descended from the original Barbary lion. These lions are also the ones that people focus on in the Barbary Lion Project.
Although there were very few specimens (skins and skulls), photographs and paintings of true Barbary lions available, by examining them Hemmer and Leyhausen reconstructed the likely external characteristics of the Barbary lion to assess the royal lions.
Photo: A Barbary lion from Algeria. Photographed by Sir Alfred Edward Pease around 1893. This image is in the public domain because its copyright has expired. This applies to the European Union, Canada, the United States and those countries with a copyright term of life of the author plus 70 years.
These characteristics are (1) greyish pelage, (2) longer fur creating a shaggy look, (3) females and young males possessing long hairs around neck/throat, back of front legs and along belly, (4) males possessing a huge mane covering not only head, neck and shoulders, but also extending behind shoulders and covering belly, (5) colour of the mane varying amongst the parts of the body and becoming darker towards the posterior parts, (6) well-developed tail tuft, (7) higher occiput (back of head), with a more pointed crown, creating a straighter line between the tip of the nose and the back of the head, (8) rounded cheek and proportionally narrow muzzle, and (9) narrow postorbital constriction of the skull (Leyhausen, 1975; Hemmer, 1978). It is believed that Barbary lions possess the same belly fold (hidden under all that mane) that appears in the Asian lions (Panthera leo persica) today. It is the largest of the lion subspecies with males weighing between 230 to 270 kg and females 140 to 160 kg. Although, due to a small sample size available for study, we have to wait until more specimens may become available to be sure about this lion's size (Yamaguchi and Haddane, 2002).
The prey for the Barbary lion were mainly Barbary sheep (Ammotragus lervia), wild boar (Sus scrofa), Cuvier's gazelles (Gazella cuvieri) and Barbary stag (Cervus elaphus barbarus), but also Arab herds of cows and sheep, and even sometimes a horse. The method of hunting was never documented, but it is believed that they used the same death by strangulation method as do the other great cats of the world. (Yamaguchi and Haddane, 2002; Preservation Station, 2005)
Image (left-hand): Barbary sheep (Ammotragus lervia) in Tennōji Zoo in Tennōji-ku, Osaka, Japan. Photographed by Kuribo on 3 January 2008. This image has been released under the Creative Commons Attribution ShareAlike 3.0 Licence. See also Wikimedia Commons.
Image (middle): Barbary stag (Cervus elaphus barbarus) from the book Alfred Edmund Brehm Az állatok világa. This image comes from the Hungarian Electronic Library (abbr. MEK). The copyright and other privileges are owned by the author/owner of the document (if he/she is known). This image can be freely copied and distributed, but you can use it only for personal purposes and non-commercial applications, without modifying it, and with proper citation to the original source. Every other form of distribution and utilization requires a permission from the author/owner.
Image (right-hand): A Cuvier's Gazelle at the San Diego Zoo in San Diego, California, USA. Photographed by Colin M.L. Burnett on 29 May 2007. This image has been released under the Creative Commons Attribution ShareAlike 3.0 Licence. See also Wikimedia Commons.
Males and females only came together during the breeding season, which was thought to be January. Plains lions have no set breeding season, and can be found mating throughout the year (Preservation Station, 2005).
Record in captivity show that their gestation period is approximately 110 days, after which 1-6 cubs are born, with 3-4 being most common. The cubs are generally heavily spotted with very dark rosettes and weigh approximately 3.5 pounds at birth. They gain an average of 3.5 ounces per day, and their eyes open around the 6th day. They begin to walk at 13 days. Female Barbary lions start coming into oestrous around 2 years old, but do not generally conceive until 3-4 years. Males show an interest in females between 24-30 months, but do not tend to produce cubs before the age of 3, and more commonly until 4. (Preservation Station, 2005)
Barbary lions were solitary like the other cat species, or occasionally lived in pairs. This was because food was not abundant. Females raised their cubs until maturity (about 2 years) and then separated from them. (Preservation Station, 2005)
|Range & Habitat||
To judge by historical records, one contiguous lion population may once have been distributed from North Africa through the Middle East to India, and was probably connected to the sub-Saharan population through the present Egypt–Sudan–Ethiopia region. Latest genetic research has confirmed this (Barnett et al. 2006). The lions distributed in the whole of Africa north of the Sahara were called Barbary lions (Panthera leo leo).
Image: map showing the possible previous range of the Barbary lion (coloured red), the possible previous range of the Asiatic lion (Panthera leo persica) is coloured yellow. Based on distribution maps by Guggisberg (1961) and Barnett et al. (2006). Created by Peter Maas for The Sixth Extinction Website. This image has been released under the Creative Commons Attribution Non-commercial No Derivatives 3.0 Licence.
However, the eastern part of North Africa (modern Libya and Egypt) may not have supported a dense lion population even well before the time of major human persecution (Harper, 1945; Nowell and Jackson, 1996), possibly because of the aridity of the region. Probably by the early 18th century at the latest, lions had disappeared from that part of the North African Mediterranean littoral. This left an isolated population in the western part of North Africa (now called Morocco, Algeria and Tunisia). (Yamaguchi and Haddane, 2002) Unlike most African lions, the Barbary Lion was a mountain predator, preferring woodlands (Preservation Station, 2005).
|History & Population||
The earliest known lion ancestor is a form like Panthera gombaszoegensis from early Pleistocene (about 1.5 million years old) deposits at Olduvai Gorge in East Africa. It had both lion- and tiger-like characters. Primitive lions (Panthera leo fossilis) dispersed in the Old World about 500,000 years ago, in harmony with changing climate and the spread of steppe-like terrain, to which lions were well adapted. "Panthera youngi", with similarities to both cave and American lions, appeared in northeastern China (Choukoutien) some 350,000 years ago or less. Probably it links Panthera leo fossilis and the "spelaea" group (cave lions of Eurasia and America) the other category being the "leo" group including the modern lions of southern Asia and Africa. (Harington, 1996)
Molecular phylogenetic studies suggested that modern lions share a common ancestor in the very recent past, estimated at between 55,000 and 200,000 years ago (O'Brien et al., 1987). Based on genetic distance, the North African-Asiatic lion is estimated to have separated from the African population as recently as 100,000 years ago (O'Brien et al., 1987).
The first humans in the range of the North African or Barbary lion clung to the River Nile for protection against the harshness of the desert. These Egyptians were the first to challenge the Barbary lions with spears and arrows. The Berbers came to found small villages across the mountains of North Africa and eke out a living from small farms, about 3,000 years ago. They defended their homes against the lions, but they were no real threat for the Barbary lion population. (Williamson, 2009)
It was the ancient Roman Empire that first reduced the Barbary to small numbers. Roman Emperors sought to entertain the people and to reassure them that their civilization had control over nature. The ancient Romans imported lions from North Africa to use in the games of the Coliseum in Rome, and other such arenas. It is known that literally thousands were taken from their homes to other parts of the Roman empire to serve as gladiator's rivals. The Roman carnage ended after six centuries, but the Barbary’s troubles were not over. The Vandals and Byzantine Empire briefly held sway over the land until the Arabs came in the 600s. As the Arab presence grew, the lions retreated. They were branded a nuisance and a reward was offered for every lion destroyed. (Williamson, 2009)
The earliest written record of a lion in England dates from 1240, referring to the upkeep of "the King's lion". It is known that King John of England established a Royal Menagerie in the 12th and 13th centuries in Woodstock near Oxford, one of the world's oldest zoos. It was later relocated to the Tower of London. The Tower's moat had been excavated in the 1930's and revealed two medieval lion skulls. DNA research showed that the skulls were from male Barbary lion, one was from late 13th to late 14th century (1280-1385) and the other from the 15th century (1420-1480). (Highfield, 2008)
With the advent of the European hunter in North Africa in the 19th and 20th century, remaining Barbary lion numbers plummeted. Local guides in the mountains of Tunisia and Morocco would track lions for European hunting for sport, live animals for zoos or museum collections. (WildLink International, 2001a)
Lions were extirpated from Tripolitania (western-Libya) as early as 1700. The last known Barbary lion in Tunisia was killed in 1891 near Babouch, between Tabarka and Ain-Draham. The last known lion in Algeria was killed in 1893 near Batna, 97 km south of Constantine, although the last Algerian lion may have been shot in an unknown location as recently as 1943. In nowadays Tunisia and Algeria the Turks had encouraged the killing of lions by paying well for the skins. After the French occupation the price went down: the French paid only 50 francs for a skin. On the other hand, many Frenchmen in North Africa became relentless Lion hunters. In Algeria over 200 Lions were killed between 1873 and 1883. Lions disappeared from the Moroccan coast by the mid-1800s. In Morocco, lions survived well into the 20th century, but finally ceased to exist in the wild in the 1940s. The last kill was recorded in 1942 on the northern side of the Tizi-n-Tichka pass in the Atlas Mountains, near the road between Marrakesh and Ouarzazat, two major tourist destinations today. (Harper, 1945; Guggisberg, 1961; Nowell and Jackson, 1996; Van den Hoek Ostende, 1999; Yamaguchi and Haddane, 2002)
Meanwhile it was said that sultans and kings of Morocco had been presented lions as the sign of obedience by the nobles, as well as by indigenous Berber people who had shared the Atlas Mountains with the last Barbary lions. Over the coming half-century the royal lions survived war and insurrection. In 1953, when Sultan Sidi Mohammed Ben Youssef (later King Mohammed V) was forced to abdicate and went into exile, the royal lions (21 in total) too lost their home at the palace. Three of them were sent to a zoo in Casablanca and the rest were sent to a zoo in Meknès. After the sultan came back to the palace in 1955, the Meknès lions moved back to Rabat again, but the Casablanca lions never came back. Meanwhile the rest of the world continued to assume the Barbary lion was extinct, a premature belief which nearly became fact when a respiratory disease hit the royal lions hard in the late 1960s. King Hassan II, then the owner, decided to reduce the risk and improve life for the lions. A new enclosure was completed in Temara, near Rabat, to house the royal lions, and in 1973, under the administration of the Ministry of Agriculture, this became Rabat Zoo. (Yamaguchi and Haddane, 2002)
Some lions in Rabat Zoo were identified in 1974 by Leyhausen and Hemmer (Leyhausen 1975) as having physical characteristics of the Barbary lion: very clear light iris, rather than brown; mane spreading behind the shoulders and covering the belly right to the groin, high occiput (back of the head), short legs and deep chest (W. York quoted in introduction to Leyhausen 1975) but none appeared absolutely flawless (Leyhausen 1975). The current royal lions in Rabat Zoo have not only the right Barbary looks, but also, very importantly, the right pedigree supported by circumstantial evidence. Although it may not be very solid, the existence of such evidence clearly separates the royal lions from all the other big-maned lions whose pedigrees are untraceable. (Yamaguchi and Haddane, 2002)
The Barbary lion's genetic distinctiveness has not yet fully established and it is still unknown whether the royal lions are true pure Barbary lions (Dubach et al. 2005; Yamaguchi 2005; Barnett et al. 2006; Burger and Hemmer 2006; Antunes et al. 2008; Black et al. 2009). However, the Barbary lion is morphologically more distinct than any of the African lion populations (Hemmer, 1978). Because of this, the guidelines of the precautionary principle (Foster et al. 2000) would suggest that reasonable action to conserve diversity is preferable. Until better data can clarify the Barbary lion's distinctiveness, maintaining the Moroccan royal lions and its decendants as a separate captive management unit distinct from other capitive lion populations is appropriate. (Black et al. 2009) These Moroccan royal lions and its decendants are now the focus of The Barbary Lion Project. For more information see 'Selective Breeding'.
Hunting no doubt contributed to the extinction of the Barbary Lion, but the ecological changes brought about by cultivation seem to have been the major cause. Forests were degraded as a result of cattle-grazing and the herds were well guarded. At the same time other food sources, such as deer and gazelle, became depleted. (Van den Hoek Ostende, 1999)
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.
Photo: a possible Barbary Lion or a possible descendent in Ljubljana Zoo, Slovenia. Photographed by Wikimedia Commons user 'Lacen' and released under the GNU Free Documentation License.
Even before Rabat Zoo was created, a German naturalist, Wolfgang Frey, who had been intensively searching for information on the Barbary lion, clearly pointed out the importance of the Moroccan royal lions. Realising how precious this bloodline was, Yves Raymond, then Director of Rabat Zoo, started to make contact with zoologists worldwide. Among them were the renowned big-cat experts Professors Helmut Hemmer and Paul Leyhausen. After being unable to answer all questions about the genetic purity of these royal lions, the zoologists decided to move forward by assuming that gene contaminations) had happened. A selective breeding programme was proposed. (Yamaguchi and Haddane, 2002)
In association with Oxford University, an UK-based organisation, Wildlink International launched their ambitious Atlas Lion Project. Wildlink International aimed to identify the Barbary markers by testing the skeletal remains of known Barbary lions in museums and collections around the world, including Barbary bones from the Coliseum in Rome. WildLink International disappeared after a while. In June 2009, The Sixth Extinction Website received new information from Wildlink International. This resurrected non-profit organisation from Aldington (Kent, England, United Kingdom) builds directly on the work of the former WildLink International. An urgent scientific population assessment of Panthera leo leo is being conducted by Pete Thompson of Wildlink International in association with Professor Helmut Hemmer (IUCN Cat Specialist Group Member) and Dr Joachim Burger of the University of Mainz (Rhineland-Palatinate, Germany). The priority now is to complete the Urgent Population Assessment without further delay (Tefera, 2003; Burger & Hemmer, 2006, Yamaguchi, 2006). (Lauren, L., personal communication 2009; WildLink International, 2001a)
Meanwhile, Simon Black, of the Durrell Institute of Conservation and Ecology at the University of Kent in Canterbury (United Kingdom), and colleagues Nobuyuki Yamaguchi, Adrian Harland and Jim Groombridge have created a Barbary lion stud book. This studbook identifies the surviving individuals, their locations, their interrelatedness and their line of descent from the original Moroccan royal population. (Black et al., 2009; Walker, 2009) The current known captive Moroccan royal lion groups in Spain, Germany, UK, France, Morocco, Czech Republic, Austria and Israel are sourced from four zoo-based family lineages: Rabat, Olomouc, Washington and Madrid (Black et al. 2009). When genetically proven similar to the Moroccan royal lions, other zoo lions could be included in future breeding efforts (Black et al. 2009).
The results will be used to develop a scientifically justified conservation breeding programme that best meets the criteria of the International Union for the Conservation of Nature (IUCN). As a final goal, reintroduction within IUCN guidelines is planned as soon as possible in the Host Country of Morocco. (Lauren, L., personal communication 2010)
Zoos and animal parks that claim to have Barbary lions or descendents
Rabat Zoo (Rabat, Morocco); Port Lympne Wild Animal Park (Kent, England, UK); Longleat Safari Park (Wiltshire, England, UK); Belfast Zoo (Belfast, Northern Ireland, UK), Zoo de Madrid (Madrid, Spain); Parc de la tête d'Or ( Lyon, France); Zoo des Sables d'Olonne (Les Sables-d'Olonne, France); Zoo Neuwied (Neuwied, Germany); Zoo Olomouc (Olomouc, Czech Republic); Hai Kef Zoo (Rishon LeZion, Israel); Great Cats of Indiana, (Idaville, Indiana, U.S.A.); Big Cat Rescue, (Tampa, Florida, U.S.A.); Tiger Safari (Tuttle, Oklahoma, U.S.A.); G.W. Exotic Animal Memorial Park (Wynnewood, Oklahoma, U.S.A.); Zion Wildlife Gardens (Kamo, New Zealand). Do you know more zoos or animal parks that claim to have Barbary lions? Please contact us.
Only nine skins, one skeleton and several skulls of Barbary Lions are preserved in museums. The National Museum of Natural History 'Naturalis' in Leiden (the Netherlands) possesses two mounted skins of males. One is only labelled "Barbarie". The other was killed by a herdsman on 13 February 1823 near Beja, Tunisia. The Lion had just brought down a cow and was killed while carrying off its prey. (Van den Hoek Ostende, 1999) The Royal Belgian Institute of Natural Sciences in Brussels (Belgium) has also Barbary lions in her collection. These Barbary lions lived in the Brussels Zoo, then located in Leopold Park, right under the walls of the present-day Royal Belgian Institute of Natural Sciences. The Belgian captive Barbary lions died there and, fortunately, found their way into the collections and the public display of the museum. (Devillers, 1999) Other museums with Barbary lion remains are the Muséum National d'Histoire Naturelle in Paris (France), and the Museo Regionale di Scienze Naturali in Turin (Italy). Do you know more museum with Barbary lion remains? Please contact this website.
Probably because of convergence in adaptation to temperate and Mediterranean habitats, it resembled the also extinct Cape Lion (Panthera leo melanochaitus), but was evolutionarily related to the lions that occupied in antiquity south-eastern Europe, the Middle East and western Asia (Panthera leo persica), now restricted to a relict population in the Gir Forest of India.
Photos: Asiatic Lions (Panthera leo persica). Taken by Adrian Pingstone at Bristol Zoo, England, in January 2005. These images have been released into the public domain by its author. This applies worldwide.
In 1968, a study on the skulls of the Asiatic, Barbary, Cape, 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-eastern European lion (sometimes seen as a seperate subspecies Panthera leo europeae) 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. (Preservation Station, 2005) There appears to be no record of contiguous populations of the two subspecies in historic times (Nowell, Jackson, 1996).
|References||Allen, G. M. (1939). A checklist of African mammals. Bulletin of the Museum of Comparative Zoology at Harvard College 83: 1–763.
Antunes A., Troyer J.L., Roelke M.E., Pecon-Slattery J., Packer C., Winterbach C., Winterbach H., Hemson G., Frank L., Stander P., Siefert L., Driciru M., Funston P.J., Alexander K.A., Prager K.C., Mills G., Wildt D., Bush M., O’Brien S.J., Johnson W.E. (2008) The evolutionary dynamics of the lion Panthera leo revealed by host and viral population genomics. PLoS Genet 4:e1000251.
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. Online full-text pdf
Bauer, H., Nowell, K. & Packer, C. (2008). Panthera leo. In: IUCN 2008. 2008 IUCN Red List of Threatened Species. <www.iucnredlist.org>. Downloaded on 18 December 2008.
Black, S., Yamaguchi, N., Harland A. and Groombridge J. (2009). Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions. European Journal of Wildlife Research.
Blainville, H. M. D. DE. (1843). Oste´ographie ou description iconographique compare´e du squelette et du syste`me dentaire des mammife`res re´cents et fossils pour servir de base a` la zoologie et la géologie. Volume 2. J. B. Baillie`re et Fils, Paris, France.
Burger, J. and Hemmer, H. (2006). Urgent call for further breeding of the relic zoo population of the critically endangered Barbary lion (Panthera leo leo Linnaeus 1758). European Journal of Wildlife Research. Volume 52, Number 1 / March, 2006. pp 54-58. ISSN 1612-4642 (Print) 1439-0574 (Online). [full text pdf online]
Day, D. (1981). The Doomsday Book of Animals, Ebury Press, London.
Devillers, P. (1999). Atlas Lions. 300 Pearls - Museum highlights of natural diversity. Downloaded on 27 September 2005.
Dubach, J; Patterson, B.D; Briggs, M.B; Venzke, K; Flamand, J; Stander, P; Scheepers, L; Kays, R.W. (2005). Molecular genetic variation across the southern and eastern geographic ranges of the African lion Panthera leo. Conserv. Genet. 2005; 6:15–24.
Fischer, J. B. (1829). Synopsis mammalium. Sumtibus J. G. Cottae, Stuttgart, Germany.
Foster K.R., Vecchia P., Repacholi M.H. (2000) Science and the precautionary principle. Science 288:979.
Guggisberg, C.A.W. (1961). Simba: the life of the lion. Howard Timmins, Cape Town.
Harington, C.R. (1996). American Lion, Yukon Beringia Interpretive Centre, Canada.
Harper, F. (1945): Extinct and Vanishing Mammals of the Old World . American Committee for International Wild Life Protection, New York Zoological Park, New York.
Hemmer, H. (1978): Grundlagen und derzeitiger Stand des Zuchtprogrammes zur Rückerhaltung des Berberlöwen. In International Tiger Studbook (eds. S. Seifert and P. Muller), pp. 65–73. Zoological Garden Leipzig, Germany.
Hemmer, H., Burger, J. (2005). Genetic confirmation of the unique nature of the zoo population of the Barbary lion. International Zoo News 52(4): 218-223.
Highfield, R. (2008). Barbary lions were part of medieval Tower of London zoo. Telegraph.co.uk. Downloaded on 11 November 2009 from http://www.telegraph.co.uk/science/science-news/3337200/Barbary-lions-were-part-of-medieval-Tower-of-London-zoo.html.
ISIS (2009) International Species Information System. http://app.isis.org/abstracts/abs.asp. Cited 25 December 2009.
Kingdon J. (1997). The Kingdon field guide to African Mammals. Academic Press, London and New York: Natural World.
Lauren, L. (2009). Wildlink International, Kent, United Kingdom. Personal communication.
Lauren, L. (2010). Wildlink International, Kent, United Kingdom. Personal communication.
Leyhausen, P. (1975): Preliminary report on the possibility of a breeding programme for the Atlas lion at Temara Zoo, Morocco. International Zoo News 21: 22–23.
Linnaeus, C. (1758). Systema naturae per regna tria naturae, secundum classis, ordines, genera, sepecies cum characteribus, differentiis, synonymis, locis. Tenth edition. Volume 1. Laurentii Salvii, Stockholm, Sweden.
Loche, [unknown]. (1858). Catalog de mammals et oiseaux observes en Alge´rie. [publisher and location not known] (not seen, cited by Hemmer 1974:230).
Meyer, J. N. (1826). Dissertatio inauguralis anatomico–medica de genere felium. Dissertation, Doctor of Medicine, University of Vienna, Austria, 62 pp.
Nowell, K. and Jackson, P. (compilers and editors) (1996). Wild Cats. Status Survey and Conservation Action Plan. IUCN/SSC Cat Specialist Group. IUCN, Gland, Switzerland.
O'Brien, S.J., Martenson, J.S., Packer, C., Herbst, L., De Vos, V., Joslin, P., Ott-Joslin, J., Wildt, D.E., and Bush, M. (1987): Biochemical genetic variation in geographic isolates of African and Asiatic lions. National Geographic Research 3: 114–124.
Preservation Station. (2005). BarbaryLion.com. Downloaded on 27 September 2005.
Skinner J.D., Smithers R.H.N. (1990). The mammals of the Southern African subregion. University of Pretoria, Pretoria.
Tefera, M. (2003) Phenotypic and reproductive characteristics of lions (Panthera leo) at Addis Ababa Zoo. Biodiversity and Conservation, 12: 1629-1639.
Van den Hoek Ostende, L.W. (1999). Lion - Slowly ticking away. 300 Pearls - Museum highlights of natural diversity. Downloaded on 27 September 2005.
Walker, M. (2009). Return of the royal Barbary lion. BBC News. Downloaded on 11 November 2009 from http://news.bbc.co.uk/earth/hi/earth_news/newsid_8109000/8109945.stm.
West P.M., Packer C. (2002) Sexual selection, temperature, and the lion’s mane. Science, 297, 1339–1343.
WildLink International. (2001a). WildLink Projects - Barbary Lions - History. Downloaded on 11 November 2009 from http://web.archive.org/web/20030608191900/www.wildlink.org.uk/proj/bar02.htm.
WildLink International. (2001b). WildLink Projects - Barbary Lions - Identification. Downloaded on 14 April 2010 from http://web.archive.org/web/20030608191713/www.wildlink.org.uk/proj/bar03.htm.
Williamson, J. (2009) Kingdom of the Barbaries. Tiger Touch Educational Centre and Resort. Downloaded on 26 September 2009.
Yamaguchi, N. (2005) The identity of the Royal lion of Morocco: a recent development in molecular investigation. Afr Lion News 6:23–26.
Yamaguchi, N. (2006) Re-evaluation of generic zoo lions. International Zoo News, 53: 1.
Yamaguchi, N. & Haddane, B. (2002). The North African Barbary lion and the Atlas Lion Project. International Zoo News 49: 465-481.
Yamaguchi, N. (2000). The Barbary lion and the Cape lion: their phylogenetic places and conservation. African Lion Working Group News 1: 9-11.
|Citation:||Maas, P.H.J. (2010). Barbary Lion - Panthera leo leo. In: TSEW (). The Sixth Extinction Website. <http://www.petermaas.nl/extinct>. Downloaded on .|
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|Updated:||20 August 2010|