Category Archives: Zoo Research

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.

How to win another 10 years for the Moroccan lions

The lions which are direct descendents from the captive collection of the Sultans and later Kings of Morocco are all in zoos. There is a healthy group of lions in Temara Zoo in Rabat, plus even more related individuals spread across a dozebn or so zoos across Europe, with around 100 animals in captivity overall. Ten years ago there were about 80 of these lions and many of those were beyond breeding age, or had already contributed significantly. Just around then, old and alone, the last female with genetic representation from founder animal 7 died and with her 1/12th of the genetic basis of the captive population.

Unfortunately until the studbook was developed from detailed examination of handwritten zoo records and a sweep of various databases, websites and personal contacts acrtoss European zoos the precariousness of the captive population was unknown. However since then a number of zoos have joined the programe to breed the animals and there have been successful transfers that have made the population a lot more healthier. When the studbook was devised it was hoped that a reinvigorated programme would give the zoo population another 10-15 years breathing space as a viable captive group.

However a few animals are underutilised – males needing suitable females – and some breeding pairings have been completely unsuccessful. Greater cooperation is needed between zoos to maximise the strenght of the whole population, not just the small groups held in each zoo exhibit. The first ten years since the revival of the breeding program has since passed. Concerted effort and active partnership is needed now to get inactive males and females together to develop breeding pairs.


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

Black S, Yamaguchi N, Harland A, Groombridge J (2010). Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions. Eur J Wildl Res 56: 21–31. doi: 10.1007/s10344-009-0280-5

Yamaguchi N, Haddane B. (2002). The North African Barbary lion and the Atlas Lion Project. International Zoo News 49: 465-481.

Barbary lion breeding shows improvements

A recent paper on improving conservation decision making (Black, 2015) includes some of the data used in developing the Moroccan Royal Lion studbook (Black, Yamaguchi, Harland and Groombridge, 2010) as previously gathered by the author with Dr Nobuyuki Yamaguchi (University of Qatar) and Adrian Harland (Aspinall – Port Lympne Wild Animal Park).

Data on productivity (in this case, the number of cubs born) shows that since the initiative to revisit and re-catalogue the lions known to be direct descendents of the King of Morocco’s collection there has been a marked increase in the production of new cubs. The great news is that these are also form well-matched pairs with no further in-breeding.

However the analysis also shows that a greater level of breeding is still needed to bring the  natural regeneration of the population under control. Currently there are practical restrictions to achieving this since clearly there are limits to the capacity that zoos have for increasing lion group sizes and for controlling breeding behaviour.

Further Reading:

Black S.A. (2015) System behaviour charts inform an understanding of biodiversity recovery. International Journal of Ecology

Black S, Yamaguchi N, Harland A, Groombridge J (2010). Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions. Eur J Wildl Res 56: 21–31. doi: 10.1007/s10344-009-0280-5

Yamaguchi N, Haddane B. (2002). The North African Barbary lion and the Atlas Lion Project. International Zoo News 49: 465-481.

Was the lion ever native to India?

lion india ashokaIn 2013 a book was published by Valmik Thapar which presented the idea that both the cheetah and the lion were most probably non-native species in India, introduced as captive animals from Africa or Central Asia, trained or used for Royal entertainment in the many substantial parks across the subcontinent and, with the demise of the various imperial and local royal dynasties between the 1200s and the mid 20th century, feral animals had become established as wild populations, hence the species now being seen as native (and rare – the lion, or extinct  -the cheetah).

This is an intriguing idea. The basis for these ideas runs from the lack of early accounts of either lion or cheetah in the region, but the subsequent rise of the follwing occurences in the time since Alexander the Great:

  • An active series of royal hunting parks and hunitng as a royal passtime with the use of lions and cheetahs being particlualry culturally important
  • Animals were exported to India from Central Europe and the middle east and also from Africa
  • The genetics of Indian lions show inbreeding suggesting an originally tiny population (escapee captive animals)
  • The genetics of captive Asiatic Lions (in the USA) shows traits of African subspecies.
  • Indian lions are ‘tame’ relative to their African counterparts (including accounts form North Africa)

Thapar and his co-writers concede that they examine this as naturalists and hitorians, rather than from a deep scientific examination of evidence. But the proposal does raise testable questions:

What are the research implications?

Do we understand the genetics of Indian lions relative to (and as different from) African lions? See recent work by Barnett et al. (2014).

Are all Asiatic lions Asiatic-African hybrids? This was the case in American Zoo animals in the 1980s – but those zoos may have mismanaged Asiatic-African pairings in captivity earlier in the 20th century.

What are the conservation implications?

Should Asiatic lions still be conserved? – YES – even if they are non-native to India, they are the last remnants of the lions which once ranged from Egypt to India (i.e. to the banks of the Indus river).

Might Indian lions be close relatives of Barbary Lions? – This is an intriguing possibility (see Barnett et al. 2014).

What about an Indian – Moroccan Royal lions Hybrid? – if Indian lions are ‘tame’ (which is NOT the case with many captive Moroccan Royal lions), then you could out-breed ‘tameness’ and retain an authentic the asiatic (northern) subspecies of lion. Similalry Asiatic lioins could eb used to retain or ‘clean up’ the Moroccan lions if they are wshown to be Barbary/subSaharan hybrids.


There is little reason to accept Thapar’s hypotheses. Improvements in genetic analysis will enable us to better understand lion phylogeny in due course. In the meantime, precaution suggests continued efforts in Indian lion conservation are strongly recommended.



Anon (2014) New Genetic Study Reconstructs Distribution History of Lion.

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

Thapar V., Thapar R., and Ansari Y. (2013) Exotic Aliens: the lion and cheetah in India. Aleph, India.


Moroccan Royal Lions: “Who’s Who”

Family tree lions symbolsBy the end of the 1990s efforts by several zoos to engage in a pan-European breeding programme for lions derived from the King of Morocco’s collection was beginning to fade. Only Port Lympne continued with an active breeding group, and a male from Rabat zoo (number 241 on the diagram opposite) was brought in to reinvigorate a pride which was developed from animals imported from Washington zoo in the 1980s. Up until that point they had reached a point of inbreeding within a family group.

Research led by Black and Yamaguchi identified all the remaining animals of known
Moroccan heritage in zoos worldwide. Aside from the animals in Rabat Zoo (Morocco), all the other descendents were is zoos in Europe, plus two animals in Israel. However there had been no transfers since the early 2000s and those which had occurred had not given rise to new cubs.

The family tree derived from this work allowed potentially suitable (unrelated) pairs to be identified and for breeding transfers to be arranged by interested zoos. This has enabled reinvigoration of the zoo stock.

PL-3 Suliman

In the early 200os nearly a quarter of Moroccan Royal lions in European zoos were related to Suliman, the male at Port Lympne (photo: N. Yamaguchi).

Further Reading:

Black S, Yamaguchi N, Harland A, Groombridge J (2010). Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions. Eur J Wildl Res 56: 21–31. doi: 10.1007/s10344-009-0280-5

Yamaguchi N, Haddane B. (2002). The North African Barbary lion and the Atlas Lion Project. International Zoo News 49: 465-481.




Barbary lions: an opportunity for significant zoo-based conservation

The barbary lion is extinct in the wild, most probably since the early 1960s.

Up until that fairly recent time the Sultans of Morocco, followed by the Kings of Morocco after constitutional changes in the 1950s, kept lions in private menageries. These animals had been bred from cubs which were presented as tributes by Berber Tribes of the Atlas Mountains.

In the late 1960s the remaining lions were still in the King of Morocco’s lion garden at the palace of Fez, then later Rabat. After an outbreak of respiratrory disease in the collection in the late 1960s, the lions were then moved to a new purpose built zoo in Rabat in 1973 (Yamaguchi and Haddane, 2002).

The importance of this is that since that first transfer of animals, it has been possible to trace all lions of pure ancestry alive today back to their ancestors – these animals originally in the Moroocan Royal palace collection. This means that zoos holding these descendents have a unique opportunity to keep the bloodline pure, and alive. Efforts by a number of zoos in recent years (Port Lympne, Olomouc, Belfast, Hannover, Madrid) to make breeding transfers and to increase the number of cubs has enabled the population to recover. As recently as 2008 it looked like breeding of these animals was likely to cease and at least one unique bloodline from the original 27 animals moved from the Royal Palace to Rabat zoo was lost at this time when an old non-breeding female in Germany died.

A rejuvinated zoo population with active, well managed transfers of animals between collectiosn will geive enough time for deeper scinetific and genetic analysis to determine the uniqueness and deeper ancestry of these Moroccan animals and their significance to lion conservation.


Black S, Yamaguchi N, Harland A, Groombridge J (2010). Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions. Eur J Wildl Res 56: 21–31. doi: 10.1007/s10344-009-0280-5

Yamaguchi N, Haddane B. (2002). The North African Barbary lion and the Atlas Lion Project. International Zoo News 49: 465-481.

Moroccan lions in zoos today

A number of zoos in Europe have lions descended directly (and exclusively) from animals which were in the King of Morocco’s royal collection in Rabat in 1969. A few examples on the animals in captivity are shown here:

PL-3 Suliman

Male at Port Lympne Zoo, UK (Photo N Yamaguchi)


Male lion (Milo) at Port Lympne, UK (Photo: S Black)

Rabat-2 females

Lionesses at Rabat, Morocco (Photo: N Yamaguchi)

During the 1990s Port Lympne in Kent (UK) was one of the few zoos with a breeding group outside Morocco. Suliman (left) was sourced from Rabat zoo and at one stage was the sire to about a quarter of the total population of animals outside Europe. He is now retired from the breeding programme.

Two of Suliman’s sons are important breeding animals. One has since been transferred to Zoo Hannover (Chalid) to join a group of females. The remaining brother Milo (left) is now intended as the prime breeding male at Port Lympne. His current partner is a lioness sourced from Madrid zoo.

Several females have been imported from Morocco into European zoos over the past decade. The focus of zoo-based breeding is to preserve important bloodlines represented by various animals across several zoos. Active arrangements for transfers of animals have been revived over recent years.

Further Reading:

Black S, Yamaguchi N, Harland A, Groombridge J (2010) Maintaining the genetic health of putative Barbary lions in captivity: an analysis of Moroccan Royal Lions. Eur J Wildl Res 56: 21–31. doi: 10.1007/s10344-009-0280-5

Research on the zoo population of lions from the Moroccan Royal Collection

The link below takes you to the 2010 article which identified the total population of animals derived from the collection of the King of Morocco, which was moved from the lion garden at the palace in Rabat to a then newly constructed Rabat zoo in 1969.

Since that time several animals and their progeny have moved to zoos around the globe. Their descendents are now only found in zoos in Europe and Israel, plus the remaining animals in Morocco, recently moved to a modern park, le Jardin Zoologuique de Rabat.

A number of other zoos across the globe claim to have animals descended from this lineage, or considered descendents of the Barbary lion, but have this provenance is yet to be confirmed.

The research article on Moroccan and European-based animals can be found here: