Category Archives: P. leo leo

Learning from the history of lions in North Africa PART 2: Relevance of decline and re-emergence of the Barbary Lion

Special contribution

Lara J. Bazzu

Worcester Prize Winner 2021, Durrell Institute of Conservation & Ecology

The distinctiveness of Barbary lions

Figure 2. A Moroccan lion at Olomouc Zoo, Czech Republic in 2000. This individual has many of the 12 morphological traits, assumed to discriminate a “pure” Barbary lion (Yamaguchi and Haddane 2002).

North African lions were considered unique amongst lion populations because of their morphology (Figure 2) and behavioural ecology (Black 2016). They lived in a variety of habitats in the Maghreb (Black 2016), the area that extends from the Atlas Mountains to the Mediterranean (Lee et al. 2015) including lowland coastal plains, forests, mountains and semi-arid areas fringing the Sahara (Black 2016).

Notably, Barbary lions were adapted to a temperate climate with cold winters (Yamaguchi and Haddane 2002). The Barbary lion lived a more solitary existence, possibly as the result of lower prey densities in temperate habitats (Mazak 1970), but was also seen in family units comprising male, female and cubs (Black et al. 2013) which contrasts with the familiar, larger ‘prides’ observed in sub-Saharan Africa lions (Mazak 1970).

Historical range of the Barbary lion

Prior to the 18th century Barbary lions still roamed widely across the Maghreb region (Black et al. 2013) which along with coastal northern Libya, comprised the lion’s original range (Black 2016). By the 19th century, bounties issued by Turkish authorities contributed to the decrease of countless lions in western North Africa and later during French control of Algeria, rewards for lions were continued and many lions were killed between 1873 and 1883 (Yamaguchi and Haddane 2002).

Figure 3. A lion photographed by Flandrin in the Atlas Mountains in 1925 (Black et al. 2013)

In Morocco lions initially fared better since the country was ruled by the sultan (Yamaguchi and Haddane 2002) but continued widespread persecution in the 19th century left the animals isolated in separate remote areas in Morocco, Algeria and Tunisia (Black et al. 2013). The last lion in Tunisia was killed in 1891 (Yamaguchi and Haddane 2002).  Strikingly, the last visual proof of a Barbary lion in the wild is a 1925 aerial photograph (Figure 3) in Morocco from a Casablanca-Dakar flight (Black et al. 2013).

A lioness killed much later, in 1942 in the High Atlas mountains of Morocco, had been considered the last encountered in the wild (Lee et al. 2015). However, tiny populations seem to have persisted in Algeria and Morocco for years (Black et al. 2013) with sporadic sightings extending up to the 1960s (Figure 4). The Barbary lion’s final demise is thought to be the result of military conflict, when the forests north of Setif were destroyed in the French-Algerian War in 1958 (Black et al. 2013).

Figure 4. Last sightings of lions in North Africa (1900-1960). Grey shading depicts Mediterranean ecosystems. Triangles indicate sightings in Algeria and Tunisia and circles for sightings in Morocco. The dotted line is the Casablanca-Agadir-Dakar air route. Dashed lines are national borders and asterisks are towns (Black et al. 2013)

Relevance of sightings for lion conservation today

A later extinction date for Barbary lions provides lessons for conserving current lion populations in West and Central Africa (Black et al. 2013). The Barbary lion story illustrates how micro-populations can remain undetected for generations (Black et al. 2013) as recently observed in Gabon. Lions were declared extinct in Gabon in 2006 yet one was seen on a camera trap in 2017 in the Plateaux Batéké National Park and subsequent DNA sampling established that it belonged to the ancestral Batéké population (Barnett et al. 2018). A lack of sightings of lions may mean conservation effort ceases (Lee et al. 2015), yet research on past sightings suggests assumption of persistence is more sensible.

Connection between Moroccan lions and Barbary lions

Intriguingly, descendants of Barbary lions may be in captivity today, thanks to the sultans and kings of Morocco, whose lion collection was derived from animals obtained by Berber tribes in the Atlas mountains (Yamaguchi and Haddane 2002). Genetic links to Barbary lions are yet to be verified and Moroccan lions are not yet officially recognised as Barbary (Black et al. 2010), since historical mixing of Moroccan lions with sub-Saharan lions cannot be ruled out (Burger and Hemmer 2005). However, the precautionary principle favours conservation of the Moroccan lineage at least until science proves otherwise (Black et al. 2013). Captive Moroccan lions are found in zoos in Europe, Morocco and Israel (Black et al. 2010), with new cubs born in Neuwied, Pilsen, Hannover, Erfurt, Heidelberg, Plättli, Olomouc, Port Lympne and Rabat. Clearly, for breeding to be purposeful, the goal should be to return animals to the wild, to support survival of the northern subspecies P. l. leo (Black 2016).

Potential for reintroduction in the wild

The habitats of the Maghreb region have seen dramatic degradation in the 20th century as a result of human expansion, drought and primarily, overgrazing (Slimani and Aidoud 2004). Lions have been absent from the area for more than 60 years (Black 2016) which means their ecological role in the region is also lost, potentially influencing land impoverishment. In order to restore the North African ecosystem, the reintroduction of two lion types has been proposed: the Moroccan lion if/when its connection to the Barbary lion is substantiated, or from the same subspecies, the Asiatic lion, which today inhabits India (Black 2016) and is also in a perilous state. Any reintroduction would require careful planning, habitat development, prey population management, community involvement and monitoring to enable a shift from small scale pilot studies to a wider scale landscape recovery.

There is a chance that people may once again be able to see lions against the backdrop of the snow-covered Atlas mountains and hear the echo of roars as described by Ormsby in 1864 (Yamaguchi and Haddane 2002). It would be a fitting soundtrack for restored forests and valleys in North Africa.


Bazzu L.J. and B;ack S.A. Les lions d’Afrique du Nord : apprendre du passé pour façonner le futur. (translated C. Guy) Le Tarsier (Association Francophone des Soigneurs Animaliers) 26, 5-9.

Black, S. (2016). The Challenges and Relevance of Exploring the Genetics of North Africa’s “Barbary Lion” and the Conservation of Putative Descendants in Captivity. International Journal of Evolutionary Biology, 2016, 1-9.

Black, S., Fellous, A., Yamaguchi, N. and Roberts, D., 2013. Examining the Extinction of the Barbary Lion and Its Implications for Felid Conservation. PLoS ONE, 8(4), e60174.

Black, S., Yamaguchi, N., Harland, A. and Groombridge, J. (2010). Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions. European Journal of Wildlife Research, 56(1), 21-31.

Burger, J. and Hemmer, H. (2005). 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, 52(1), 54-58.

Lee, T., et al. (2015). Assessing uncertainty in sighting records: an example of the Barbary lion. PeerJ, 3, e1224.

Mazak V. (1970). The Barbary lion, Panthera leo leo (Linnaeus, 1758); some systematic notes, and an interim list of the specimens preserved in European museums. Z Saugetierkd 35,34-45.  

Slimani H. and Aidoud A. (2004) Desertification in the Maghreb: A Case Study of an Algerian High-Plain Steppe. In: Marquina A. (ed) Environmental Challenges in the Mediterranean 2000–2050. Vol. 37. Dordrecht: Springer, pp. 93-108.

Yamaguchi N., Haddane B. (2002). The North African Barbary lion and the Atlas lion project. Int. Zoo News, 49, 465–481.

Renewed importance of the Moroccan Royal Lions

A new study by Lehocká et al (2021) published right at the end of December 2021 in PLos ONE focuses attention back on the Moroccan Royal Lion population. Previous studies provided data on the Moroccan Royal lions which has now been examined further in this interesting analysis of the captive population of animals that are descendants of the King of Morocco’s collection.

As this blog has repeatedly stated, these lions, now mostly scattered across European zoos as well as a significant group still in Rabat Zoo, Morocco, are a potentially very important remnant of the North African lion population. In contemporary terms we should see these as being part of the wider Northern lion subspecies Panthera leo leo (range India, Middle East North Africa , West Africa, Central Africa).

Fig 1 Diagram showing the structure of the pedigree file based on Wright’s fixation index (FST) from Lehocká et al (2021) showing relations between captive ‘Morocccan Royal lions’, defined by the location of individuals in zoological gardens, showing three main groups.

The PLoS ONE paper highlights that the effective population size of this wider group (of just under 100 animals) is only 14.

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 Pleo leo.

Further Reading

Lehocká, K., Black, S. A., Harland, A., Kadlečík, O., Kasarda, R., & Moravčíková, N. (2021). Genetic diversity, viability and conservation value of the global captive population of the Moroccan Royal lions. PloS one16(12), e0258714.

The Barbary Lion Podcast

The North African lion story is discussed in the excellent Cats of the Wild podcast series, Episode 14  ‘The Barbary Lion’.

The history of lion presence in countries across North Africa, and investigation into the final decline of the species in the region and the lessons which need to be learned for current lion declines in West and Central Africa are discussed.

There is also discussion on the tantalising possibility of reviving populations based on captive animals in zoos in Europe and Morocco.

The Barbary Lion (2021) Episode 14, Cats of the Wild


Why is the Barbary Lion still important?

Until recently lions were colloquially split into various geographic grouping, often supported by characterisations from natural historians and hunters of the time variously covering ‘biggest’ , fiercest’, ‘tamest’, ‘most cowardly’ and so on. Historically names followed a range of different local clades, the Cape lion, Indian lion, Senagalese lion, Persian lion (sometimes known as the Mesopotamian lion), Nubian lion, various groups across central and eastern Africa and of course the Barbary lion.

Significant recent genetic studies have enabled more accurate categorisation of lions sub-populations into two subspecies, based on several phylogenetic studies. The Cat Classification Task Force of the Cat Specialist Group revised lion taxonomy into two subspecies – the northern sub species Panthera leo leo and the Southern subspecies Panthera leo melanochaita.

The northern species is the nominate species – first encountered by scientific classification as the animals were present and encountered in the accessible landscapes of antiquity around the Mediterranean. However in today’s global population the northern subspecies only accounts for 10% or less of the global lion population (much less if we include captive lions).

The diminishing micro-populations of lions in West Africa today mimics the decline of the Barbary lion in North Africa – can lessons be learned? For example in Senegal, the lion is the national symbol and the ‘name’ of the famous national football team. How many lions live in the wild in Senegal – less than 50.

And what of the supposed captive lions from Morocco – the likely descendants of the last Barbary lions? Well they may represent nearly half of the captive collection of all northern lions. If we ignore these animals it will be to our peril.

Lion conservation in captivity has, perhaps run a full circle in 50 years since Hemmer and Leyhausen raised excitement about the lions being moved from the King of Morocco’s collection in the Royal Palace in Rabat into new enclosures in the city zoo.

Lion populations in the wild have certainly more than halved since then, possibly are at a quarter of the levels in the 1970s.

Are we ready to use what knowledge and resources we have to turn things around?


Bertola LD, Jongbloed H, Van Der Gaag KJ, De Knijff P, Yamaguchi N, Hooghiemstra H, et al. (2016) Phylogeographic patterns in Africa and High Resolution Delineation of genetic clades in the Lion (Panthera leo). Sci. Rep. Aug 4; 6: 30807.

Black, S.A. (2016) The Challenges and Relevance of Exploring the Genetics of North Africa’s Barbary Lion and the Conservation of Putative Descendants in Captivity. International Journal of  Evolutionary Biology Article ID 6901892,

Canine Distemper in Gir lions highlights restricutions of population size and range

See the source imageFrom September 12 and October 2 2018, a total of 23 lions have died in the Gir Wildlife Sanctuary and National Park due to an outbreak of Canine Distemper, carried by domestic dogs, but known to have previously caused a mass wipeout of lions in East Africa’s Serengeti-Masai Mara ecosystem.

Asiatic lion expert Ravi Chellum summed up the situation: “If you allow the entire population of a species to be concentrated in one place without any populations in other areas as a safeguard to mitigate risk, this kind of scenario is bound to happen.”

Whilst issues with a small founder population are real, in the case of Asiatic lions which dwindled to about 20 animals in the 1930s, the issue is unavoidable. Genetic variation is very low among Asiatic lions with animals to some degree related to any other since they all descended, and this lack of genetic variation makes them vulnerable to the effects of disease.

Chellum has long advocate that these lions should be housed in different geographical areas so that they can develop new adaptation to their geography which can then reflect in their genetic structure. Separated sub-populations also act as a potential buffer to the spread of disease. Despite significant planning and identification of potential areas outside Gujarat, no progress has been achieved.

In June, the government announced that 109 sq km of area in Amreli and Bhavnagar districts of Gujarat would be reserved as a new sanctuary for lions. State politics has dogged plans for relocation of a small sub-population of these animals. Perhaps now is a serious prompt for a rethink on this strategy. India’s lions are the last large wild group of lions of the newly designated subspecies (recommend to the IUCN in 2016), Panthera leo leo, the northern group which ranged from West Africa and northern Africa across the Middle East into South Asia.

No one seriously challenges the cultural and biological importance of lions in Indian ecosystems, so some effort is needed to enable their sustainable conservation as a nationally and regionally important species.


Anon (2018) Gujarat lion deaths: What killed 11 big cats? BBC News 25th September 2018

Barnett, R. et al. (2014) Revealing the maternal demographic history of Panthera leo using ancient DNA and a spatially explicit genealogical analysis. BMC Evolutionary Biology 14: 70; doi: 10.1186/1471-2148-14-70

Bertola, L. D.,et al. (2016) Phylogeographic Patterns in Africa and High Resolution Delineation of Genetic Clades in the Lion (Panthera leo). Scientific Reports 6:30807. DOI: 10.1038/srep30807

Chellam, R., Joshua, J., Williams, C.A. & Johnsingh, A.J.T. (1995) Survey of Potential Sites for Reintroduction of Asiatic Lions. Unpublished Report, Wildlife Institute of India, Dehra Dun, India.

DownToEarth (2018) Those who have endangered the Asiatic Lion’s future, have to be held accountable.–61789

Johnsingh, A.J.T. , Goyal, S.P.  and Qureshi, Q. (2007) Preparations for the reintroduction of Asiatic lion Panthera leo persica into Kuno Wildlife Sanctuary, Madhya Pradesh, India. Oryx , 41(1), 93–96.

Stop and think – what about lions today?

There are perhaps as few as 20,000 lions left in the wild. When I first started studying lions just over a decade ago we thought there were 40,000. Will this figure halve again over the coming 10 years? For a sobering comparison, when I was a child there were 40,000 tigers in the wild; now only around 4,000. The similarity in the lion figures is startling. Decline is real and fearsome.

A comprehensive set of studies of lion genetics has recommended the IUCN to consider the lion family tree as two subspecies. One is the North/West /Central subspecies, Panthera leo leo (which includes lions in West and Central Africa and India, and the previous, now extinct populations in the Middle East, and of course the extinct Barbary lion of North Africa). The second is Panthera leo melanochaita from South and East Africa (i.e. about 90% the world’s population). All remaining ‘northern’ lion populations are tiny and very vulnerable. The ‘southern’ populations are much larger but many animals are cut off from each other in fenced reserves or fragmented landscapes.

Zoos may have a significant part to play in managing, sustaining and recovering the ‘northern’ subspecies Panthera leo leo. There are around 100 Asiatic (Indian) lions in captivity, but I do not know of a single captive lion from West or Central Africa in any zoo, even in their home countries on the African continent. There are a further 80 captive lions which may be related to the extinct North African Barbary lion. maybe the captive Addis Ababa lions in Ethiopia are linked I some way to this group.

Addis Ababa lion

A lion in Addis Ababa Zoo, Ethiopia

Lions are quite long-lived and can persist undetected for decades. This happened in North Africa, where lions had become almost mythical beasts by the late 1890s, but actually survived in northern Algeria into the late 1950s at a time when Gerry Durrell was founding the zoo in Jersey. I often wonder what Durrell might have done had he known that Barbary lions were still out there in the wild… However, this is not a purely historical issue since in 2016 a lion was caught unexpectedly on a camera trap in Gabon, and later confirmed by DNA evidence to be a survivor of a population thought extinct for 20 years – one of the important Central African relic populations of the ‘northern’ lions Panthera leo leo.

Lions in the Mountains

As lions in North Africa became increasingly persecuted, populations sin the lowlands and coastal forests were eliminated and the population retreated to remoter areas to avoid human contact. Although in parts of Morocco and Algeria lions were not be seen in previously commonly inhabited areas for many decades, the animals were still observed unexpectedly in remoter, less well-explored areas years later.

Lion behaviour in the mountains would be more solitary, somewhat like the Siberian Tiger (A Harland pers.comm) than communal (as per lions in East Africa). However, even under these circumstances  lions tend towards social behaviour (Black et al. 2013), so a pair of animals would more likely stay together than disperse. It also seems, from hunting accounts, that in mountainous areas they would use ‘lairs’ usually caves as a base, so that is quite a peculiarity related to the landscape.

Although snow can persist on the highest peaks, the snowfall in the Atlas Mountains tends to be limited to the January February period and the strength of the African sun means that in general although snow cover will occur, the evaporation of a snowfall is quite fast compared to equivalent altitudes in Europe or North America.

Lions were known to range up to 3500m and footprints were used by hunters to track animals in the snow. If an animal was being pursued it would keep to remoter areas and if that meant the snowline it would stay there (see my point about caves below)and would be less likely to drop into lowlands if there was human activity unless there was prey (e.g. livestock). The animals would use mountain passes to track prey and keep to areas clear of human presence. So a lion in the snow would be somewhat transient – due  to both the passing nature of snowy conditions in the lower altitudes and the lesser need for the lions themselves to be at higher altitudes other than for transit.

Due to lower density of prey in the North African ecosystem (compared to African savannah or open forest) lions tended to be single, in pairs, breeding pairs or breeding pair with juveniles, rather than the expansive prides of southern Africa (Asiatic lions in India have smaller prides but not as small as North African lions). Single animals would often range on their own, certainly single males (of which at least one photograph exists), but there are also records of single females being shot in high mountain passes. The range of a lion in North Africa would have been quite extensive – we see this today in desert dwelling lions in Namibia for example. In northern forests of Morocco, Tunisia and Algeria in the 19th century some ranges were predictable to the point that locals would know a path that a lion took – and could tell hunters (N Yamaguchi pers comm). However in the Atlas mountains and southern Morocco, certainly in the 20th century, this was much less predictable as the distances animals travelled was far greater.


Black SA, Fellous A, Yamaguchi N, Roberts DL. (2013) Examining the extinction of the Barbary lion and its implications for felid conservation. PLoS ONE 8(4):e60174

Yamaguchi, N. and B. Haddane, B. (2002) “The North African Barbary lion and the Atlas lion project,” in International Zoo News, vol. 49, pp. 465–481, 2002

Occasional sightings suggest lions cling on in West Africa

Currently the West African populations of lions are Critically Endangered (Henschel et al., 2014) and currently occupy only 1% of their historical range. The presence of lion in Ghana is therefore very important, although it is known that many west african sub-populations cling on in single figure numbers.

A recent paper (Angelici and Rissi, 2017) suggests a recent sighting 11 years since the last confirmed lion presence in Mole National Park. If present the small number of individuals (perhaps as few as two or three) suggested by the sightings reported in the paper, might appear insignificant. Neverthless many populations in West africa are of this size, totalling perhaps less than 200 across the entire region – the ast wild representatives of the IUCN’s newly designated northern subspecies of lion Panthera leo leo (the rest are in India).

The few individuals in Mole are a sub population of a tiny population across Senegal, Burkina Faso, Benin, Niger and Nigeria (Henschel et al., 2014), which itself is a tiny population of Panthera leo leo globally (probably less than 2000 animals including in captivity).

Although we rarely consider individual animals in conservation, with lions it has reached that stage. If a few individuals can be conserved in Ghana, they might provide hope for continuation across the region and for the northern sub-species Panthera leo leo as a whole.


Angelicic, F.M. and Rossi, L. (2017) Further lion, Panthera leo senegalensis Meyer, 1826, sightings in Mole National Park, Ghana, and possible first serval Leptailurus serval Schreber, 1776 record after 39 years (Mammalia Felidae). Biodiversity Journal, 8 (2): 749-752

Henschel, P. et al. (2014) The lion in West Africa is Critically Endangered. PLoS ONE, 9: e83500. Doi: 10.1371/journal.pone.0083500



So what’s the big deal about a bunch of zoo lion cubs?

Five lion cubs have recently been shown to the public for the first time with great fanfare at Neuwied zoo in southwestern Germany.

The litter of cubs — three males Baz, Chaka and Sab and two females Jumina and Lin  — were born at the zoo in April 2017,  to six-year-old female Zari (previously from Hannover zoo) and the incumbent male, Schroeder (who was born in Olomouc Zoo in the Czech Republic), who is nearly ten.

So what is the big deal? The zoo presents them as barbary lions – certainly Schroeder and Zari are established on the European zoo studbook for lions descended from the King of Morocco’s collection. Around 300 animals are lsited on the studbook going back to the origninal animals taken form the King’s Palace in the late 1960s and placed in Rabat zoo.

Since there are only a few hundred lions in India and a few hundred in West and Central Africa which represent the northenr subspecies of lion, Panthera leo leo (Bertola et al, 2016; Black 2016). the north african population is no loinger present in the wild, and its most likely remnant are the 100 or so animals in european Collections and in the Mrooccan collection at Rabat. So these five new cubs may yet have an important role to play in lion conservation.


Asian Age (2017) German zoo displays rare barbary lion cubs

Associated Press (2017) 5 rare barbary lion cubs go on show in a zoo in Germany

Bertola, L. D.,et al. (2016) Phylogeographic Patterns in Africa and High Resolution Delineation of Genetic Clades in the Lion (Panthera leo). Scientific Reports 6:30807. DOI: 10.1038/srep30807

Black S.A. (2016) The Challenges of Exploring the Genetic Distinctiveness of the Barbary Lion and the Identification of Putative Descendants in Captivity, International Journal of Evolutionary Biology. vol. 2016, Article ID 6901892, 9 pages, 2016. doi:10.1155/2016/6901892

Spot the lion habitat

Typical savannah landscape in southern Africa

Lions are commonly associated with the savannah grasslands of southern and southeastern africa. They live in large prides, hunting herds of antelope and other ungulates of the grasslands. However these expansive grasslands are specific habitats to sub-saharan lions of eastern and southeastern Africa Panthera leo melanochaita. These habitats are the heartlands for lions in the modern world.

Unexpected sight of lions in desert dunes, southwest Africa

Dry forest in the Gir, India

Yet lions also survive in quite doverse habitats, such as the desert dunes of Namibia.

This would have been true for the northen subspecies Panthera leo leo in areas of the middle east and North Africa. Camels were known to be tracked by lions on isolated trails in the late 1800s. Animals were reported in the desert firnges of southern Morocco up to the 1930s and were also known in isolated forests and oases in western Algeria,

For  today’s remnant population in India mostly live in the dry forests of the Gir. These forests are less productive in dry years with prey numbers falling under challnging conditions. Some lions have chosen to venture beyond the Gor,, across agirculaural land and into new but limited habotats in the coastal forests and dunes of Gujarat. Smaller populations survive in the forests of central and west Africa, in the deserts of southwestern Africa.

Northern lions also previously roamed the marshes of the middle east, the mountains of North Africa occasionally up above the snowline, down to the coastal forests of the Mediterranean.

Image result for cork forest algeria

Mediterranean cork oak forests would seem to be an unfamiliar habitat for lions.

High Atlas plateau, North Africa. The last lions in Morocco were seen at high altitudes in the 1930s and 1940s.


Black, S. A. (2016). The Challenges and Relevance of Exploring the Genetics of North Africa’s “Barbary Lion” and the Conservation of Putative Descendants in Captivity. International Journal of Evolutionary Biology, 2016.

Yamaguchi, N. and B. Haddane, B. (2002) “The North African Barbary lion and the Atlas lion project,” inInternational Zoo News, vol. 49, pp. 465–481, 2002