Category Archives: Ecology

Importance of recent historical records of lions

Lions have enjoyed significant protection in Gujarat’s Gir Forest in India. There are accounts of colonial and local hunters shooting animals outside the state in the late 19th century.

Lions are in severe decline across sub-Saharan Africa, most particularly in West and Central African countries. Even in Eastern or Southeastern Africa many populations are now isolated by geography or fences. The decline of the species across Africa is worryingly similar to the historical disappearances of lions from other regions.

Recent collation and analysis of historical records have given insight into the range of the Barbary lion in North Africa, providing important insights into how lion populations are in decline, the point at which they are most vulnerable to extinction and what the animals tend to do under extreme persecution.

For example, Black et al. (2013) confirm that the range of lions reached far further south into the Saharan Atlas of Algeria than was ever known, even in the mid 1800s when numbers were higher and lion hunting was common in the region. Individual animals appear to have ranged across a wide expanse of arid habitat. This mean that the populations across Algeria could be considered contiguous with populations in the high atlas of southern Morocco.

In India similar reports from 19th century colonial India include hunters taking lions in the Jahore and Marwar districts of Rajasthan in the 1870s, well north of their stronghold in the Gir forest, Gujarat state the last significant home of Asiatic lions over the past 100 years. In 1872 a professional hunter (Bhil Shikari) associated with a Mr. T. W. Miles brought in the skin of a full-grown Asiatic lioness which he had shot on the Anadra side of Mount Abu, the last met with in Sirohi districts. Around the same year a Colonel Hayland bagged four lions near Jaswantpura, in Marwar. These were apparently the last lions seen across the Kutch border into Rajasthan (Adams, 1899).

Lions are a long-lived species, so if isolated individuals range to less-threatened areas they are able to survive for prolonged periods, perhaps giving the impression of a population being maintained. This may not be a reality of persistence, but instead an indication of a single wandering, isolated individual as is suspected today in the recent lion sighting in Gabon.

From a conservation perspective, it is important to separate these exceptional ‘one-off’ sightings, from the true indicators of surviving micro-populations, the latter of which require careful systemic attention. For example, tiny populations of lions were thought to have survived in the remote Saharan Atlas mountains for decades after the species was considered extinct from North Africa (Guggisberg, 1963; Black, 2016). As for the exceptional one-off sightings – well they offer hope*; hope that a suitable habitat can be utilised for future lions population to find genuine refuge.


*Note:  the recent unexpected sighting of a Spix Macaw in the wild, not seen in the area since the last known specimen disappeared in 2000 is a good example of an exception. No one believes the animal has been kept hidden for sixteen years – this individual it is most probably a recent captive release. However the learning point is  – can this animal survive in the remaining habitat, does it display survival behaviours and would in situ conservation of the species be a viable proposition that can be taken forward seriously?



Adams, A. (1899) The Western Rajputana States: A Medico-topographical and General Account of Marwar, Sirohi, Jaisalmir.

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.doi:10.1371/journal.pone.0060174

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. doi:10.1155/2016/6901892

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

Saul H. (2016) Male lion filmed roaming in West African nation of Gabon for the first time in 20 years. The Independent



Dwindling ecology of the Sahara: large animals in decline


A view of the Sahara desert in Algeria – the last Algerian lions survived on the fringes of the Sahara into the 1940s with others surviving in northern coastal forests into the 1950s. (Photo by Florence Devouard)

The Bubal hartebeest is extinct, the dama gazelle and addax survive in only 1% of their former range, and various other hoofed animals from the region are extremely threatened:  the Nubian Ibex clings on in parts of Algeria whilst the Barbary sheep fares a little better.

Of the major predators, the Barbary Lion has certainly been extinct in the wild for 50-60 years (assuming some of its ancestors still survive in a handful  of zoos). The African wild dog no longer inhabits the Sahara desert (and was only present in southern fringes. The Saharan cheetah only inhabits 10 per cent of its former range, surviving largely unnoticed in southern Algeria. Similarly a small leopard population is known to survive in the southern Algerian Sahara, but is lost from 97 percent its historic range.

However there are opportunities for recovery: the scimitar horned oryx is extinct in the wild, but controlled releases in fenced reserves in Tunisia and Chad are looking to re-establish the wild population. Similarly the Addax has been reintroduced in fenced reserves in Morocco and Tunisia. The leopard was thought extinct in the late 1990s, but a small population appears to survive in the Atlas Mountains. Do these examples of experimental reintroductions and relic populations offer hope for wider recovery of North African and Saharan ecosystems?


Anon (2013) Sahara Desert’s large mammals Slipping Into Extinction. Environmental News Service. December 5, 2013

Busby et al (2009) Genetic analysis of scat reveals leopard (Panthera pardus) and cheetah (Acinonyx jubatus) in Algeria. Oryx, 43(3), 412–415

Wildlife Conservation Society. “Critically Endangered Cheetahs In Algeria Snapped With Camera Trap.” ScienceDaily. ScienceDaily, 27 February 2009. <>.

Could lions solve a secondary predator issue in North Africa?

Feral dogs photographed by A. Woodhouse

The concept of secondary predator effects on degraded ecosystems is well established. If the large carnivores at the top of the ‘food chain’ are eliminated, then species lower down the pecking order are given a free reign to devastate prey species or the vegetation, if they are grazers (Borrvall and Ebenman, 2006; Duffy, 2002). By inference the presence of primary predators controls the trophic levels below them in the ecosystem.

Whilst large carnivores are necessarily rare, from a trophic, or food energy transfer, perspective (Colinvaux, 1990), they appear to have an amplified impact on the behaviour and presence of other species can be a significant moderating effect in ecosystems. A recent study identified that even perceived presence of large carnivores reduces the feeding routines and impact of secondary predators  in particular in experiments with racoons (Suraci et al 2016).

Do these observations and phenomena relate in any way to the pressures on ecosystems in North Africa? Clearly, land use by mixed flocks of goats and sheep would be changed (to some extent) by the presence of large carnivores, although how this would affect long term landscape recovery is unclear (without some system to manage those flocks). There also is a suspicion that some Barbary macaque populations are be impacted by predation from feral dogs, particularly causing losses in infant monkeys.

Could lions provide an opportunity to deter overgrazing or the presence of feral dogs in those habitats of Barbary macaques?

Further Reading:

Borrvall, C. and Ebenman, B. (2006) Early onset of secondary extinctions in ecological communities following the loss of top predators. Ecology Letters, 9, 435–42

Butynski, T.M., Cortes, J., Waters, S., Fa, J., Hobbelink, M.E., van Lavieren, E., Belbachir, F., Cuzin, F., de Smet, K., Mouna, M., de Iongh, H., Menard, N. & Camperio-Ciani, A. (2008) Macaca sylvanus. The IUCN Red List of Threatened Species 2008: e.T12561A3359140. Downloaded on 26 February 2016.

Colinvaux, P. (1990) Why Big Fierce Animals Are Rare. Penguin Books.

Duffy, J.E. (2003) Biodiversity loss, trophic skew and ecosystem functioning. Ecology Letters,  6: 680–687 doi: 10.1046/j.1461-0248.2003.00494.x

Suraci, J.P., Clinchy M., Dill, L.M., Roberts, D. and Zanette L.Y (2016) Fear of large carnivores causes a trophic cascade. Nature Communications, 7, 10698  doi:10.1038/ncomms10698

Woodhouse, A. (2014) Feral dogs at iMfolozi. Africageographic, Posted: April 30, 2014

A new angle on modern lions’ backstory

Modern lions (Panthera leo) began their exodus out of North Africa towards the end of the Pleistocene, eventually reaching as far as India. Much later, just around 5,000 years ago, another group of lions left the continent, reaching what is today Iran, in the Middle East, forming populations which are now extinct. This understanding of movement in prehistory may have important implications for the conservation of modern lions, since the study by Barnett et al. (2014) identifies that lion populations in West Africa and Central Africa, which have drastically declined over the past few decades, are actually more closely related to the Indian lion than to the more numerous populations of lions in East Africa (for example lions in Somalia or Botswana).

Today in India fewer than 400 Asian lions (P. leo persica) survive in the wild, living on the Kathiawar Peninsula in Gujarat, and this subspecies is listed as ‘Endangered’ by the International Union for the Conservation of Nature. Despite the large geographical distances between them, these lions also seem closely related to the Barbary lions of North Africa. Could a conservation plan involving Indian lions be part of the answer for recovery of North African populations – and could North Africa itself be a potential refuge for the Asiatic lion?


Further Reading:

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

The solitary lion?


photo credit: image/713-Jbel_hebri_rocher

The lion is distinctive in being the only big cat which lives in extended social groups.

The North African ecosystem is
a relatively low energy system (click here) with little food available for carnivores when compared to African savannah or Indian dry forests. Lions
were known to move above the (temporary) snow line in the Atlas Mountains – an even more harsh environment.

This probably explains the reasons for most sightings of Barbary lions in the 18oos and19oos referring to either single animals, pairs, or pairs with cubs; the larger pride structures associated with East Africa are not encountered, and the type of female-dominated family groups which are commonly encountered in India are not described in documented North African sightings.

Some commentators suggest that the behaviour of the Barbary lion was more similar to the Siberian tiger – relatively solitary. There are certainly many examples of single animals being cornered and shot in isolation.

Reading links:

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. doi:10.1371/journal.pone.0060174

Black, S. (2015) Room to move in?

The sixth vision: could the former northern range become a refuge for Panthera leo?

One perspective on Barbary lion and North Africa revolves around the ‘romantic’ notion of reintroducing this charismatic species back into its former range. A second perspective is more pragmatic and equally visionary; using the species as a focus for driving the restoration of habitats in the region. A third vision, more pragmatic perhaps, would be to drive a tourist industry for economic benefit. A fourth vision is to preserve genetic diversity in Panthera leo or even selectively ‘breeding-back’ the Barbary lion, by retaining the genes held by lions from the Moroccan Royal collection (although this could still be achieved in captivity). A fifth vision would see North Africa developed as a new enclave for Panthera leo persica (currently only extant in India) – from the wild population most closely related to the Barbary lion.

Could there be a sixth vision – to provide an enclave for Panthera leo as climate change disrupts the suitability of existing habitats south of the Sahara? A recent paper shows the risk of decline in current habitats suitable for lions (Peterson et al 2014). Up in the northern strip of Africa, along the Mediterranean coast there are potentially some small enclaves of habitat. Would it be prudent to make these a refuge for lions?

If so, which lions would we put there? What sort of ecosystem should develop as a result (prey, landscape, human use)? Which might be the best locations? What controls might be needed to protect humans, livestock and lions?

Peterson and Radocy Climnate change predictions


Peterson A.T., Radocy, T., Hall, E., Peterhans, J.C.K., and Celesia, G.G. (2014) The potential distribution of the Vulnerable African lion Pathera leo in the face of changing global climate. Oryx 06/2014; 48(04):1-10. DOI:10.1017/S0030605312000919

The Northern Lions

The last micro populations of lions north of the Sahara held out in the North West of Africa (Algeria and Morocco) and in Iraq, Iran and India in the East. Of these, just 400 remain in the Gir Forest with a small number now appear to be established in the Kodinar Coastal strip in southern Gujarat, India.

Lion distribution map inc 20th century in northThe last record of lions in Iraq was possibly the two shot by a Turkish governor in 1914 near Mosul and later in 1918 in the lower Tigris . The last Iranian lions had largely dissappeared in the 1940s with sporadic sightings by railway engineers in the years during the second world war. A lion was also thought to have been seen near Quetta in Pakistan in 1935.

In 1963, the last pride of five Persian lions was hunted in the Dasht-i Arzhan districy of Fars Province in Iran. According to Guggisberg, national newspapers and media “celebrated” the killing of these lions with pictures and fanfare. The pride consisted of a female with four cubs in a cave, the male had been shot already. Just as in earlier accounts from Algeria in the 1880s, the female was shot on the spot, and the cubs were taken as trophies. No subsequent sightings have been reported from Iran, although an attempt was made to reintroduce lions into the region in the 1970s, but the animals dissappeared, presumably shot.


Divyabhanusinh A. (2008) The Story of Asia’s Lions. The Marg Foundation.

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

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

Room to move in?

Picture a valley in North Africa, described by Yamaguchi & Haddane (2002):
Forests 02 00777f3 1024“…Between the Middle and High Atlas lies a rocky mountainous area where green oaks dominate the landscape…where the endangered Barbary leopard may still survive… Barbary macaques (Macaca sylvanus), Barbary sheep (Ammotragus lervia) and wild boars live there, and Cuvier’s gazelles (Gazella cuvieri) and Barbary red deer (Cervus elaphus barbarus) may also be reintroduced…lions may be released into a securely-fenced semi-natural enclosure…to live with minimum human intervention…, releasing them into an open area is out of question… “

In the North African biosphere, the prey biomass densities are much lower than in the savannahs of Sub-Saharan  Africa or the dry forests of western India. The Moroccan Atlas is likely, on average, to be able to support probably less than four lions per 100km2. In the Gir Forest in Gujarat, India, where domestic livestock as a supplementary prey base, the carrying capacity of lions is estimated  at around 15 lions per 100 km2 (Banerjee et al., 2013).

Other comparisons with the higher energy, prey dense  and open landscape of the Gujarat (where lions can leave the forest for surrounding human-dominated areas in times of food shortfalls) show how constrained the landscape in North Africa would be:

  • Gir Forest, Gujarat (1500 km2) – 400 lions
  • Kodinar coastal forest, Gujarat (60 km2) 12+ lions

Translocation of lions in India has been planned for some time. The planned release site in Kuno Wildlife Sanctuary (an area of 900 km2) would involve reintroduction of only 5-8 animals intially (with plans to move 1-2 adult males every 5 years out of the sanctuary). So, even in a relatively large and currently established wildlife habitat with a reasonably dense prey base, only a very small number of lions would be released.

Additional complications of home range size and number of groups would be a major constraint for any lion reintroduction in North Africa. Would North African animals live in large prides as cmomonly encountered on the African savannah, in family groups, or would a single lioness holds resource territory while male coalitions attempt to maximize female groups within their range, as in India? (Black et al., 2013; Yadvendradev et al, 2009)?

Whilst having wild lions back in North Africa is a big dream for some, it has practical limitations. And then there are the significant needs of local people…


Banerjee K, Jhala YV, Chauhan KS, Dave CV (2013) Living with Lions: The Economics of Coexistence in the Gir Forests, India. PLoS ONE 8(1): e49457. doi:10.1371/journal.pone.0049457

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. doi:10.1371/journal.pone.0060174

Linares, J.C.,  Taïqui, L. and  Camarero, J.J. (2011) Increasing Drought Sensitivity and Decline of Atlas Cedar (Cedrus atlantica) in the Moroccan Middle Atlas Forests Forests, 2(3), 777-796; doi:10.3390/f2030777

Yamaguchi N, Haddane B. (2002) The North African Barbary lion and the Atlas Lion Project. International Zoo News 49 (321): 465-481.

Yadvendradev, V.J. et al. (2009) Home range and habitat preference of female lions (Panthera leo persica) in Gir forests, India. Biodiversity and Conservation.DOI: 10.1007/s10531-009-9648-9

Lion food – try it Tunisian style

lion and boar mosaic

Mosaic of lions eating wild boar, 2nd century AD, Museum of El-Jem, El-Jem Thydrus, Tunisia

This 2nd century Roman mosaic from Tunisia is a fine example of its genre and interesting for its depiction of lions, a not unfamiliar scene in artwork of the era.

The picture manages to look both familiar and unfamiliar at the same time. Two lions devouring prey, yet it is two male lions. The prey is relatively large compared to the lions but it is only a pig. The scene is a  dry and sparse climate and there are depictions of human presence a temple or palace perhaps… all somewhat fanciful, surely?


Wild boar (Sus scrofa) in India.    Photo S Black

There are many pre-Roman artistic depictions of lions devouring prey in Mediterranean art (bulls, deer etc.). Perhaps fewer of lions eating wild boar. We do know that wild boar are the prey of lions in India today and boar would have been prey elsewhere in the former middle eastern range of the species. They would also have been a key prey for lions in North Africa as depicted in the mosaic. If lions were still present in the wilds of Algeria or Morocco today, then wild boar would probably be the best food source (other than domestic flocks).

We know that Barbary lions tended to operate in small family groups rather than the prides familiar in savannah landscapes, so it is interesting to see two males in this depiction – does this offer a clue to how Barbary lions behaved, perhaps?

In reality almost all lion depictions of the classical era show male lions and almost never lionesses, so little can be gained from this depiction. The Tunisian mosaic is most probably a purely artistic interpretation of events. It is satisifying, however, to see the lions depicted with a known wild prey species form the region. In classical European artwork lion prey is more often depicted as domestic cattle, donkeys, horses or wild deer. Interestingly 18th and 19th century art often depicts lions attacking horses and domestic camels.

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. doi:10.1371/journal.pone.0060174

Yamaguchi N, Haddane B. 2002. The North African Barbary lion and the Atlas Lion Project. International Zoo News 49 (321): 465-481.

Lion decline indicates mass extinction

Estimates suggest that today there are around 32,000 to 35,000 lions left. Sixty years ago this  figure was somewere between 400,000 and 500,000, so in a lifetime we have seen a decline of 90 percent or more. Over that period the lion population in India (Panthera leo persica) has clung on and grown from just under 200 animals to about 350 – 400 animals today (from a low in the 1890s of about 20 individuals). Even in the 1940s lions were observed as far west as Iran but the entire Middle East no longer is home to these animals. Over this period the North African population has also dissappeared (with the last sightings in the late 1950s).

Worse still, the remaining bulk of population in sub-saharan Africa (see figure below) is patchy and sometimes involves tiny micropopulations, particularly in Central and West Africa, which could easily follow the fate of Middle Eastern and North African populations.

Riggio et al (2012) Densities of lions in Africa


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. doi:10.1371/journal.pone.0060174

Riggio, J., Jacobson, A., Dollar, L. et al. (2012) The size of savannah Africa: a lion’s (Panthera leo) view.Biodiversity Conservation. DOI 10.1007/s10531-012-0381-4