Matt Wedel kindly passed on the photos you see here. They show the Man-eater of Mfuwe, an enormous male lion Panthera leo that terrorised the small town of Mfuwe (and the surrounds) in the Luangwa River Valley of eastern Zambia. The photos were taken in Chicago’s Field Museum where the specimen has been on display since donation in 1998 (sorry about the glare - it's impossible to take good photos without smashing into the case, and the museum generally discourages that sort of thing).

The history and behaviour of man-eating big cats is truly fascinating stuff; the stealth, cunning and occasional reckless confidence they display in getting at their victims, the level and extent of their depredations, and the stealth and cunning that human hunters have to use to put an end to these cats are frequently extraordinary and make fascinating reading. I regularly refer to the 1992 book Man-Eater: Tales of Lion and Tiger Encounters (edited by Edward Hodges-Hill). It includes articles on the Iyenpur tigress, the Jerangau, Champawat, Kempekarai and Mamandur man-eating tigers, the ‘Ogre of Bellundur’ (a tiger), the Tsavo lions, Jim Corbett’s adventures with the Thak man-eater, and many others (Hodges-Hill 1992).

Of all these killer cats, perhaps the most famous were the two male lions who killed a claimed 135 people* in what is now Tsavo National Park during the 1890s. The story of these two lions and their depredations have been the subject of three Hollywood movies, the best known of which is the 1996 The Ghost and The Darkness. An excellent and thorough book, Bruce D. Patterson’s The Lions of Tsavo: Exploring the Legacy of Africa’s Notorious Man-Eaters, discusses this episode in full, placing it within the context of modern research on lion behaviour, conservation and variation (Patterson 2004).

* This number (estimated by Colonel John Henry Patterson, the killer of the two lions) was questioned by Yeakel et al. (2009). Based on stable isotope analysis and estimates of calorific requirements, they reckoned that both lions killed about 34 people in total, a number substantially lower than Patterson's estimate. Yeakel et al. (2009) might be right, but they might not.

It isn’t the only book on the Tsavo lions – there’s also Philip Caputo’s Ghosts of Tsavo: Stalking the Mystery Lions of East Africa (Caputo 2002). Both books discuss the Man-eater of Mfuwe; my text here is based almost entirely on their accounts.

The Mfuwe lion wasn’t killing and eating people back in Victorian times or anything like that. Rather, it was a modern man-eater, its attacks occurring in 1991. The first occurred as two boys were walking along a road at night. One boy fled; by the time game rangers arrived at the attack site, only pieces of clothing and part of the other boy’s skull were found. The second attack occurred at the edge of a village – this time the victim was an adult woman; the lion had broken through the door of her hut to get her. The third attack (occurring at night on a boy who ventured out to meet a friend) was foiled when a game scout fired his gun into the air during the attack, but the boy died of his severe injuries anyway.

Three more kills occurred later in 1991, the last one again involving a woman being dragged from the hut where she lived. Prior to this last kill, it was locally believed that lionesses belonging to the ‘L-pride’ were responsible for the killings, and indeed one of these lionesses was shot dead in August of 1991. The important role of an adult male lion now came to the fore. This lion entered the woman’s hut during daylight and removed a bag containing her laundry. It took the bag to the centre of the village, dropped it, and stood over the bag, roaring. Remarkably, the lion carried the bag about the countryside, leaving it at different locations and sometimes playing with it. Unsurprisingly, perhaps, people now suspected that the lion was no ordinary lion, but a demon or sorcerer in lion form.

Some sources say that people weren’t allowed to hunt and kill this lion because Zambia’s laws about game conservation didn’t allow it. That’s not wholly correct, given the killing of one of the suspected man-eating lionesses by game wardens (they also shot a juvenile male lion at the same time, but didn’t kill it). The other females in the pride were also shot dead later on.

The Mfuwe man-eater’s reign was put to an end by Californian hunter Wayne Hosek. Hosek wasn’t the first to try and kill the Mfuwe lion – a professional hunter, and a Japanese hunter and naturalist, had both tried earlier. Hosek was already in the area with permission to hunt individuals from a list of large mammal species, lion among them. There’s the whole debate here about controlled hunting and its contribution to economy and conservation versus its abuse at the hands of the unscrupulous, the selective impact it has on animal populations, and the ethics of trophy-hunting. I’m somewhat mystified by the idea that anyone can look at a lion, elephant or hippo and feel compelled to shoot it to death, but my personal perspective on this matter is that giving live wildlife an actual dollar value can only be a good thing, at least if you want live wildlife to still be present in future decades.

Anyway, Hosek and professional hunter Charl Beukes (or Buekes? Both spellings are used in the literature) sat for about three weeks in a hide, hoping that a nearby bait of hippo meat would attract the cat and allow them to kill it. Early attempts were unsuccessful – the lion even circled the bait one night, but was so stealthy that it had avoided all detection. Eventually, Hosek succeeded in killing it. On the lion’s death, people swarmed out of the village to spit at the animal and beat it with sticks.

The Mfuwe lion, today on display at the Field Museum in Chicago, is enormous: 3.2 m in total length, 1.2 m tall at the shoulder, and with a mass estimated at 249 kg (Caputo 2002). And, contrary to local descriptions of it being equipped “with a huge mane” (Caputo 2002, p. 3), it’s wholly maneless.

Manelessness in lions – the competing hypotheses

Patterson wrote that Hosek was due to write a book about the Mfuwe man-eater; so far as I can tell this hasn’t happened, but do say if you know otherwise. If you’ve read Patterson’s or Caputo’s books, or if you know a lot about lions in general, you’ll know that the Man-eater of Mfuwe was similar to the man-eating male lions of Tsavo in being maneless. In fact some witnesses did say exactly this, but their descriptions of a maneless lion were assumed to be of a lioness, hence the deliberate targeting of lionesses in efforts to eliminate the man-eater.

Why is manelessness prevalent among lions of this region? Actually, this is a rather controversial subject and numerous hypotheses have been entertained: for a thorough, fully referenced discussion, see Patterson (2004). Perhaps the most intriguing suggestion is that the lions concerned represent a distinct, ancient lineage, phylogenetically separate from other living lions and perhaps being relict, Pleistocene-type lions (Gnoske & Kerbis Peterhans 2000). The authors who promote this hypothesis have used the term ‘Buffalo lions’ for these animals (von Buol 2000) since they describe them as specialised predators of Cape buffalo and other especially big prey. It has also been called the ‘basal lion hypothesis’ (Patterson 2003). Molecular and morphological data does not support this idea: DNA and morphometrics both show that Tsavo-type, maneless lions are deeply nested within Panthera leo, being close to other east and south African lion populations (Patterson 2004, Dubach et al. 2005). [Maneless lion photo below by Mgiganteus.]

The possibility that lions have lost manes because manes are physiologically costly in some environments has also been entertained, as has the idea that manes are reduced and lost in habitats where they provide a disadvantage to moving about among thorn-scrub. Then there’s the idea the maneless lions are old and poor in condition (Patterson (2004) called this the ‘down-and-out hypothesis’), and the alternative idea that their manelessness results from especially high levels of testosterone (that’s right, high levels of testosterone causes hair follicles to shrink: hair reduction and loss is the result). Given that Tsavo-type lions are evolving within a social system where single males, rather than coalitions, associate with female prides, it might be that a particular male physiology is driving manelessness. But, as usual, it seems that several factors are at play: thorn brush and a hot, dry climate might contribute to mane reduction in Tsavo-type lions (Kays & Patterson 2002, Gnoske et al. 2006, Patterson et al. 2006) as well. This issue was previously discussed in the ver 2 article Why the Lion Grew Its Mane, a book review.

Why become a man-eater?

As much as I’d like to continue discussing variation within lions, my time is up, and this is still a topic I plan to return to at length. One more thing is worth saying: why do some big cats become bold man-eaters in the first place?

It’s a well known bit of lore that man-eating big cats are sometimes injured or sickly individuals that take to killing humans because it’s easy, and because the animals are desperate enough to disregard normal caution or fear. The Mfuwe lion was described as looking “a little green around the gills” when seen in the field, and had sustained a major injury to the lower jaw that had resulted in pustulous lesions (Patterson 2004, pp. 76-77). The two Tsavo lions both had craniodental injuries that might have prevented normal predatory behaviour (Yeakel et al. 2009). However, this 'infirmity hypothesis' doesn’t explain all cases, since many problem big cats have been perfectly healthy.

Other individuals become bold and experienced following a chance encounter with an unfortunate human, and then deliberately target people in future. Others switch to humans as prey because funeral rites, atrocities resulting from slavery and so on mean that human remains left out in the open are opportunistically scavenged and hence ‘turn on’ a culture of human-eating in some individuals or groups. Finally, shortages of natural prey caused by drought, disease epidemics and extirpation by people also seem to cause lions, tigers and other big cats to switch to humans through desperation.

Human encroachment on big cat habitat, the invasion of habitat by domestic livestock and by people hunting for bushmeat – in short, human disruption of ecosystems – means that big cats both become more familiar with people as potential prey items, and are increasingly pushed into a corner where humans become ever more desirable and available prey items. And some or all of these phenomena might combine, meaning that the cause of man-eating behaviour is not exactly simple. Of incidental interest is that human predators have sometimes used outbreaks of man-eating in lions and other cats to conceal their own murderous habits. Of the several creepy stories on record, perhaps the weirdest is that concerning the ‘lion-men of Singida’. These were actually drugged young women “sewn into animal hides, and fitted with deadly prosthetic claws” (Patterson 2004).

For previous articles on lions and other cats, see...

Refs - -

Caputo, P. 2002. Ghosts of Tsavo: Stalking the Mystery Lions of East Africa. National Geographic Society, Washington, D. C.

Dubach, J., Patterson, B., Briggs, M., Venzke, K., Flamand, J., Stander, P., Scheepers, L., & Kays, R. (2005). Molecular genetic variation across the southern and eastern geographic ranges of the African lion, Panthera leo Conservation Genetics, 6 (1), 15-24 DOI: 10.1007/s10592-004-7729-6

Gnoske, T. P., Celesia, G. G. & Kerbis Peterhans, J. C. 2006. Dissociation between mane development and sexual maturity in lions (Panthera leo): solution to the Tsavo riddle?Journal of Zoology 270, 551-560.

- . & Kerbis Peterhans, J. 2000. Cave lions: the truth behind biblical myths. In The Field 71, 2-6.

Hodges-Hill, E. (ed) 1992. Man-Eater. Tales of Lion and Tiger Encounters. Cockbird Press Ltd, Heathfield.

Kays, R. W. & Patterson, B. D. 2002. Mane variation in African lions and its social correlates. Canadian Journal of Zoology 80, 471-478.

Patterson, B. D. 2004. The Lions of Tsavo: Exploring the Legacy of Africa's Notorious Man-Eaters. McGraw-Hill, New York.

- ., Kays, R. W., Kasiki, S. M. & Sebestyen, V. M. 2006. Developmental effects of climate on the lion's mane (Panthera leo). Journal of Mammalogy 87, 193-200.

von Buol, P. 2000. 'Buffalo' lions. A feline missing link? Swara: the Magazine of the East African Wildlife Society 23 (2), 20-25.

Yeakel, J. D., Patterson, B. D., Fox-Dobbs, K., Okumura, M. M., Cerling, T. E., Moore, J. W., Koch, P. L. & Dominy, N. J. 2009. Cooperation and individuality among man-eating lions. Proceedings of the National Academy of Sciences 106, 19040-19043.