Category Archives: History

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

Lions in Pakistan or on the shores of the Arabian sea?

Lions survived in the territories of Pakistan up until the mid 1800s, the last of them believed to be killed in 1842 near Kot Diji in Sindh, so its presence there is essentially ancient history.

However there have been recent rumours of lions being spotted in a national park of Azad Jammu and Kashmir, with observers suggesting that these are animals migrating from India. However the lion stronghold of  Gujarat in India (particularly the Gir Forest and the Arabian coastal strip) are hundreds of miles south. These unusual  sightings may relate to mis-identification of other animals (such as leopards) or, more speculatively, released animals from clandestine captive collections (the latter has been suggested for leopard sightings in Mediterranean western Turkey in the late 20th Century).

On the rocks in a Gujarat port – ready to take a swim.

There have been incidents of illegal trade in live African lions into Pakistan in the recent past; how do you get rid of illegal lions? Could that be part of the explanation? In addition of course there are a number of zoos in Pakistan which have captive Asiatic lions (Panthera leo persica) in their animal collections, although these rare specimens would most likely be kept properly secure. Nevertheless the fact that lions are present in the peninsular of north-western India raises some interesting sightings. Animals from the Gir forest (landlocked by extensive tracts of agricultural land) sometimes have to disperse.

By moving across agricultural landscapes and areas of human habitation a small but significant number end up living adjacent to industrial sites, within small coastal forest scrub lands and adjacent to busy ports. The spectacle of a lion walking along a beach or being seen swimming in the sea is, despite its apparently improbability, a matter of fact.


Anon (2016) Lion spotted in Arabian Sea off Gujarat coast, rescued. Pakistan Telegraph (ANI) Sunday 3rd January 2016

Anon (2010) Four lions Imported illegally to Karachi. BBC News South East Asia

Khan, H.N. and Craig, T. (2015) In North-west Pakistan big cats are more feared than global terrorists, The Guardian Weekly/Washington Post 19th September 2015.

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




More on the Northern (near) extinctions

The demise of lions from northern regions (above the Sahara) followed a sequence starting in Europe in ancient historical times, central Asia and Egypt and along the North African coast, then a slow shrinkage from the eastern Mediterranean countries.

Small populations of Asiatic lion (Panthera leo persica) clung on in Syria, Iraq, Iran and Pakistan to the end of the 19th Century and some micro populations continued in the latter three countries into the 20th century, but have only remained in the Gir forest in India since that time.

The Barbary lion (Panthera leo leo) disappeared from coastal Morocco in regions near the larger human population centres in the 19th century, but survived in remote areas of the High Atlas of Morocco and the Saharan Atlas (north central Algeria) and Aures mountains in northeastern Algeria into the second half of the 20th century.

detailed lion extinctions in North Africa and Middle east to India (Black version 2015)


Bartosiewicz, L. (2009) A lion’s share of attention: Archaeoogy and the Historical Record. Acta Archaeologica Academiae Scientiarum Hungaricae. DOI: 10.1556/AArch.59.2008.2.2

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

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?

Insights into late survival & extinction of the Barbary lion

Late sightings of lions in North Africa may seem surprising and certainly some of the stories have oddities about them (Black et al. 2013). What should not be forgotten, however, is how distinguishable a lion is from other similar species in the region – the chances of people mis-identifying a lion are relatively slim.

Even among the most recent sightings of Barbary lions in Morocco and Algeria the variation of circumstances when people see the animal is huge – for example one sighting is a photograph taken while flying over the Atlas mountains, another is lion observed by locals on a bus, whilst several others occur when the animal was shot (Lee et al. 2015).

Assessment of historical sightings by expert panel reveled the reliability of these sightings is generally high, so varied sources can be sensibly used to map out the story of the species’ decline.

Supp1 Figure

Observations of lions in North Africa 1895 – 1960 Grey shading indicates Mediterranean scrub land. Triangular markers indicate lion sightings; (sightings 7–21) in Morocco (western Maghreb) and sightings 133-149 in Algeria (eastern Maghreb). Black circles denote human population centers. Dashed lines indicate national boundaries.

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

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



To worship the lion

Sketch of a pride of Cave lions. Note the lack of manes on the big standing male. Art by Tabitha Paterson (@TabithaPaterson)

Sketch of a pride of Cave lions. Note the lack of manes on the big standing male. Art by Tabitha Paterson (@TabithaPaterson)

We think of lions, today, as African animals. This is mostly true. However, there is still a tiny refugium of non-African lions, isolated in the Kathiawar peninsula of India, and centred on the Gir forest reserve. Here, 400 or so Asian lions eke out an existence, beset on all sides by people and farmland, the last remnants of an empire that once spread from Tunisia via Turkey to the Tigris and beyond.  But, even this is only a fraction of the range that lions once held.

During the Pleistocene, highly differentiated lion subspecies (or perhaps separate species, opinion is divided) roamed from Spain to Siberia, through the steppes of Beringia, and into the Americas as far south as Mexico. Their fossils are surprisingly common in Britain too. In fact, excavation of the site of Trafalgar square uncovered a number of lion fossils where now their equally impressive bronze cousins lie today. The cave lion (Panthera spelaea) occupied all of Eurasia and Beringia. The closely related American lion (Panthera atrox) was found over the contiguous lower 48 states.

Range of lions since the Pleistocene. Image by Ross Barnett

Range of lions since the Pleistocene. Image by Ross Barnett

The cave lion is, and was, a pretty special felid. Considerably larger than modern lions, it was the apex predator of the Pleistocene food web (with perhaps some competition from Homotherium).  As it lived in Europe at the same time as anatomically modern humans, it has been depicted in numerous pieces of parietal and portable art. The cave walls of Chauvet and Lascaux contain brilliantly realistic images of this extinct animal, showing that it lived in prides, and that males were maneless. We know this because in a few images, the adult male scrotum is obvious, and the mane is absent.

Pride of cave lions from Chauvet cave. Public domain image.

Pride of cave lions from Chauvet cave. Public domain image.

It also seems that early Europeans had some kind of cultural affinity for the cave lion. One of the most amazing pieces of art to come from this period, exquisitely crafted from mammoth ivory, shows a half-lion, half-human chimera. This löwenmensch, as it is known in german, testifies to some kind of ritual or mythic importance for the cave lion in the culture of the time. Like the venus figurines, löwenmensch, have been found at multiple sites, showing that the idea was not just an isolated one but shared amongst communities.

Löwenmensch from Hohlenstein-Stadel. Image by Dagmar Hollmann via Wikimedia Commons. License: CC BY-SA 3.0

Löwenmensch from Hohlenstein-Stadel. Image by Dagmar Hollmann via Wikimedia Commons. License: CC BY-SA 3.0

Further reading:

Nice piece by the Telegraph, featuring our very own Ross Barnett: ‘Super-sized lions’ roamed UK in Ice Age.

Barnett, R., et al. (2014), ‘Revealing the maternal deomgraphic history of Panthera leo using ancient DNA and a spatially explicit genealogical analysis’, BMC Evolutionary Biology, 14, 70. [Full Article]

Barnett, R., et al. (2009), ‘Phylogeography of lions (Panthera leo ssp.) reveals three distinct taxa and late Pleistocene reduction in genetic diversity’, Molecular Ecology, 18 (8), 1668-77. [Abstract]

Conard, N. J. (2003), ‘Palaeolithic ivory sculptures from southwestern Germany and the origins of figurative art’, Nature, 426 (6968), 830-32. [Abstract]

Franks, J. W. (1960), ‘Interglacial deposits at Trafalgar Square’, The New Phytologist, 59, 145-150Montellano-Ballesteros, M. and Carbot-Chanona, G. (2009), ‘Panthera leo atrox (Mammalia: Carnivora: Felidae) in Chiapas, Mexico’, The Southwestern Naturalist, 54 (2), 217-22. [Abstract]

Montellano-Ballesteros, M., and G. Carbot-Chanona (2009). ‘Panthera Leo Atrox (Mammalia: Carnivora: Felidae) in Chiapas, Mexico.’ The Southwestern Naturalist 54, no. 2 , 217-22. [Abstract]

Packer, C. and Clotte, J. (2000), ‘When Lions Ruled France’, Natural History, 109, 52-57. [Full Article]


Posted on BarbaryLion with thanks to Ross Barnett and colleagues at TwilightBeasts: Jan Freedman (@janfreedman), Tabitha Paterson (@TabithaPaterson), and Rena Maguire (@justrena).
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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.

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.

Old-fashioned human-wildife conflict

There are very few taxidermy specimens of Barbary Lions. Over the past 15 years my colleague, Dr Nobuyuki Yamaguchi has spent months attempting to locate these treasures, following the footsteps of Vratislav Mazak, the famous biologist who had previously tracked down many specimens in the 1960s. Sadly some of these items have gone missing over the past 40 years.

Velizar Simeonovski

“The thief of Beja” by V. Simeonovski (click image to see full size version)

Only one specimen, in Leiden’s Naturalis Museum in the Netherlands, includes clear information on its provenance. Leading widlife artist Velizar Simeonovski has recreated the scene, showing the male lion being shot at close range (thumbnail link, right) by a local Tunisian defending his livestock. Velizar also offers an interesting commentary on the story.

Th male lion at Beja was shot in 1823, over a hundred years before the last lions dissappeared from the region. Whilst many of the subsequent encounters between people and lions in North Africa include livestock predation by lions it is also true that lions became more adept at withdrawing into remote areas away from human contact. In the 20th century only one third of the 30 encounters with lions resulted in the animal being shot and only two incidents involved livestock attacks.


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

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

Simeonovski, V. (2014) “The thief of Beja” 13 February 1823, the vicinity of Beja , Tunisia.

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

Here, in times past, lions roamed…

Ain Talawan1ss

Lion country: Ain Talawan (Photo: N Yamaguchi)

This landscape is more reminiscent of Scotland than Africa, but is a valley in north-eastern Algeria. Accounts of lions in this region continued up to the early 20th century (Black et al 2013). One of the more memorable regular encounters by local people with lions used to occur in this region.

The stream above the valley runs near a track which was frequently used by local people taking goods to market.

However the stream, although fairly unassuming (see the photo below) was an importnat water point for lions. This meant that people regularly had to drive lions away from the area with sticks when travelling through the area. A colleague in Algeria has collected verbal accounts from an old man who used to travel this route in the 1920s.

An unassuming stream in hills above Ain Talawan. (Photo: N Yamaguchi)

The scene is reminiscent of the encounters in the Gir Forest in Gujarat, India which the local Maldhari people  experience with the Asiatic lion (Panthera leo persica).

A Maldhari and his livestock in the Gir forest. He has a stick to steer his animals and to protect them and himself against lions.









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

Moroccan Royal Lions