Category Archives: Ecology

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


Fragmentation of lion populations: a global threat

Lions are one of the more numerous charismatic carnivores, merely ‘vulnerable’ according to the IUCN Red List.

However the real picture, aside from core populations in east Africa, is one of fragmentation. For most of their historic range lions are today only found either in isolated protected areas, the remnants of habitats from their former range (particularly in West Africa, Central Africa and India) or in fenced reserves (Southern Africa).

Their history of survival and decline elsewhere suggests this is a very poor situation. We see our surviving African and Indian populations suffering pressures for the last two or three decades but clinging on. This may not be sustainable. In North Africa (Tunisia, Algeria, Morocco) and the Middle East (Syria, Iraq, Iran, Turkey) the same situation held true from the 1880s through to the 1940s (even 1950s). Today few people would associate any of those countries with the lion.

Will we say the same about many of its current range states in the next ten or twenty years?


Black, SA (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, vol. 2016, Article ID 6901892, 9 pages, 2016.

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

Bauer, H,  Chapron, G,  Nowell, K. et al., (2015) Lion (Panthera leo) populations are declining rapidly across Africa, except in intensively managed areas. Proceedings of the National Academy of Sciences of the United States of America, 112 (48), pp. 14894–14899.

A. E. Schnitzler, AE (2011) Past and present distribution of the North
African-Asian lion subgroup: a review. Mammal Review, vol. 41, no. 3, pp. 220–243.






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

Food for lions: diet in the Atlas mountains

Wild boar were certainly a main prey species for lions in the Atlas Mountains, but also red deer (Barbary deer) and barbary sheep, plus oryx, addax and small gazelles(Cuvier’s gazelle, Dorcas gazelle, Dama gazelle).

As human inhabitants became more numerous the temptation for predating livestock grew.  Lions were a threat to the mixed flocks (sheep and goats) which were kept in mountainous areas by pastoralists. Flocks would not be left overnight  near wooded areas or places known to be inhabited by predators, and would be moved closer to villages or to open ground at least.

Flocks up in the higher ground that would stay out but if the herder was wary they would put up rough stockades of thorn bushes as protection from leopards, jackals and hyenas. This appears likely because the same type of temporary construction was used by local guides taking people travelling through the mountains if the had to stay overnight in a remote areas in the 1800s and early 1900s.

Cattle were also vulnerable and many accounts mention their demise at the jaws of lions. Additionally on the domestic front were pack animals, the camels and mules used by travellers and pastoralists.

It would not be unusual for a rough stone shed or shelter to be made near regular pastures (especially in cattle pastures) for farmers to stay in overnight – more common in lowlands hills though. If a flock had been attacked previously, the farmer would stay in the hope the visiting predator would venture in again and could be shot. There are certainly accounts of this ‘stake out’ approach in Tunisia and Algeria in the 1800s.


Black, SA (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 vol 2-16, Article ID 6901892 9 pages

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

Leopard and Tiger spatial use implications for conservation

Conservation of wildlife is often a question of trade-offs. In most cases, including the in conservation of large carnivores the trade-off is between human needs and big cat needs. In these cases a point of tolerance by both parties is reached, relating to economics, security, retaliatory action and translocation of dangerous animals.

However it has been noted that where tiger conservation has been successful and tiger numbers have increased, there has been a knock-on reduction in leopard presence. Leopards will generally retreat from areas where tigers dominate. This is a classic shift in niche breadth as a function of interspecific social dominance (Morse 1974). Clearly a tiger is an apex predator and will take out (and prey on) leopards.  These two big cats differ in the size of prey killed, use of vegetation types, and in activity periods (Seidensticker, 1976).

In the absence of the tiger, the leopards can exist on prey which is similar in structure and less abundant than required by tiger. In the presence of tiger, leopards have been shown to shift to areas where the tiger does not occur frequently.

Also some evidence suggests that leopards are forced into predation of small livestock (which tigers do not do) when habitats and prey bases are shared (Bhattarai & Kindlmann, 2012), which raises conflict between cats and humans once again.

This has implications in areas where leopards themselves are under threat, the amur leopard in the Russian Far East being a good example.

Dramatic footage captures a tiger and a leopard in a brutal battle to the death… just a few feet from shocked tourists.

Further information:

Bhattarai, B. P., & Kindlmann, P. (2012). Interactions between Bengal tiger (Panthera tigris) and leopard (Panthera pardus): implications for their conservation. Biodiversity and conservation, 21(8), 2075-2094.
Carter et al (2015) Impacts of people and tigers on leopard spatiotemporal activity patterns in a global biodiversity hotspot. Global Ecology and Conservation, 3 910: 149-162.
Morse, D. H. (1974) Niche breadth as a function of social dominance. Am. Nat. 108: 818-830.
Seidensticker, J. (1976) On the ecological separation between tigers and leopards. Biotropica, 225-234.

Nicols, S. (2015) Conservation targeting tigers pushes leopards to change. Michigan State University. centre for Systems Integration and Sustainability

Escalation of Human-Wildlife Conflict is never the answer

This dramatic, duisturbing photograph sees local people in Bengal trying to dissuade wild elephants with firecrackers and burning balls of tar. Whilst elephants are highly destructive, dangerous, unpredictable, this type of conflict is unlikely to be productive.

An adult elephant will not be stopped, even by elephant proof fencing. As human land-use encroaches on traditional habitats and pathways of wild animals these conflict increase in frequency. Deterrents such as fencing, chilli ropes have an effect, but also the education of local people on how to respond to elephant encroachment.

A further challenge is to get officials, developers and local people to understand how to reduce the disruption of habitats by building in unsuitable areas.

An adult elephant and a calf on fire flee a crowd of people: The picture was taken by Biplab Hazra, a wildlife photographer from West Bengal.

© Biplab Hazra/Sanctuary Asia/Facebook The picture was taken by Biplab Hazra, a photographer from West Bengal.

Conflict causes escalation of consequences, for people, for wildlife or both.

Elsewhere in Assam, efforts by conservationists to give early warning to local crop producers when elephant herds are drawing near allows families and communities to remove crops from fields before the mammals encroach on the agricultural landscape. Whilst this arrangement might not be optimal, it is an example of the type of tolerance and human cooperation which reduces conflict and removes incentives of elephants to visit landscapes which would otherwise be attractive for foraging.

Other challenges occur with big cats, which can easily pass unnoticed in urban landscapes let alone villages and agricultural areas. this means living with a number of challenges discussed elsewhere in this blog.

Local people chase a leopard in Guwahati, Assam, 2009 (northeast India). The animal was tranquilized by a local wildlife official and relocated. However three people in the village had already been mauled by the big cat.

Further information these images:

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








Lions in Pakistan – wild or released – implications?

The last of lion killed in the regions of Pakistan was shot in 1842 near Kot Diji in Sindh.

However, with conserved population of the species (hosted by the Gir Forest in Gujarat, India) now expanding to the south and west, the possibility of lions leaving their protected areas and eventually expanding their range north or west into Pakistan is a possibility.


Recent social media alludes to active trade in lion cubs

There are even hints that lions have already moved into the Pakistani countryside bordering India (Naqaush, 2014).

Is another explanation possible? Might captive animals be  released in remote areas and then been seen and considered ‘wild’ specimens?

Recent social media posts allude to an active trade in cubs. When cubs become adults they become a new proposition. What if a collector or breeder ends up with too many males or an incompatible pair? Where do these captive lions come from? India is unlikley, africa, or african-origin captive history is more likely. WHat if those ex-captives hybridise with the wild asiatic lion?

The local countryside might, to an unwise but disgruntled owner, seem a good place to make unwanted animals ‘disappear’…for lion conservation it could be a whole new problem.


Mulki , M.A. (2012)A Walk on the Wild Side. The Express Tribune, Sunday Magazine, January 29th.

Naqaush, T. (2014) Asiatic lion spotted in AJK national park, DAWN Febraury 5th

Lions in the Rif of northern Morocco

Leopards were commonly hunted in Morocco well into the 20th century.

A colleague recently visited the national park of Talassemtane in the Rif mountains touring the area with a local guide. The guide told them that these mountains, near Chefchaouen, still retained dense fir forests up until after the second World War and that only shepherds visited the summits because people who lived in the towns and villages of the valleys were afraid of the wild landscape and the possible presence of lions. According to the guide, researchers from the Ceuta, believed that the lion was still present during the 20th century up until the time when mountains of the area had been deforested. Do these observations have any basis in fact?

Certainly there were lions in Morocco up to and including the second world war, although they were seen further south. One was shot in the High Atlas Mountains as late as 1942 in the Tizi‐n‐Tichka pass, and a few years before a pair were seen south of the Atlas ranges on the Saharan fringes, with a further group seen in the same area in the mid 1930s. All of the known 20th century sightings were south of Fez, often in the areas around Ifrane, Azrou, Kenifra and further south around Toubkal or further south again beyond Assa.

The last known sighting in the north (the Rif Mountains and up towards Tetouan) was of a lion killed in 1895. However this does not rule out lions holding on in that region much later in small groups, especially if areas were not visited by people. For comparison, in Algeria several small lion populations were known up to the 1930s and up to the late 1940s, even though many sources suggest the disappeared by the 1890s. The last known sighting in Algeria was in 1956.

Extinction models show that, accounting for the frequency and spacing of sightings, lions could have persisted in both Morocco and Algeria up to the early 1960s (Black et al 2013; Lee et al, 2015). Only the destruction of habitat along the Mediterranean coast during the French-Algerian War suggests that lions might have disappeared earlier, perhaps by 1958.

Of course fear of lions (real or imagined) only tells part of the story of concerns by local people in the Rif Mountains in the 1940s. The other factor which may have concerned people in the area would be leopards. They still persist in Morocco today and would have been an important threat to livestock and, as we know from other regions, also a threat to people.

Further Reading:

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

Guggisberg C.A.W. (1963) Simba: the life of the lion. London: Bailey Bros. and Swinfen

Lee TE, Black SA, Fellous A, Yamaguchi N, Angelici FM, Al Hikmani H, Reed JM, Elphick CS, Roberts DL. (2015) Assessing uncertainty in sighting records: an example of the Barbary lion.PeerJ 3:e1224

Schnitzler, A.E. (2011) Past and Present Distribution of the North African-Asian lion subgroup: a review. mammal Review, 41, 3.

Rural, sub-urban & urban cats: understanding unintended impacts in the dynamics of big cat conservation

Alley cat by Nayan Khanolkar

A leopard ghosts through an alley in a suburb of Mumbai bordering Sanjay Gandhi National Park. Leopards often enter the streets at night, which can lead to conflict with humans. This outstanding photo taken by India’s Nayan Khanolkar, was Urban category winner in the Natural History Museum’s 2016 Wildlife Photographer of the Year.

Leopards have been shown to shift territories in areas where tiger conservation in protected areas has allowed the larger species to dominate a habitat. This means that leopards, a highly adaptable species, are pushed into rural and urban areas. Inevitably, this brings them into conflict with people.

Although leopards may adapt and co-habit with tigers and people, depending on the prey-base available to all three species, leopards themselves are the underdogs in these interactions. There is some evidence that leopards adapt with nocturnal dispersal into human landscapes (to avoid contact with the humans already living there). The photograph above is an illustration of this sort of behaviour (other insights in this short clip )

It appears that leopards avoid tigers in space, but humans in time and these differences in behaviour have implications for managing conservation in areas where human-dominated landscapes border leopard and tiger territories.


Further reading:

Carter, N., Jasny, M., Gurung, B. and Liu J. (2015) Impacts of people and tigers on leopard spatiotemporal activity patterns in a global biodiversity hotspot. Global Ecology and Conservation 3(1): 149:162.

Nichols, S. (2014) Conservation targeting tigers pushes leopards to change. Michigan State University