Home
Work in Progress
Full Frontal Lion Charge
Video of Lion Call
Book Excerpt
Racing Sunset Video
Exceptional Guide
Reviews
Buy This Book now
About the Author
Bob's Blog-Plog
Other Bob Frump Books
Other Man-Eating Books
Men's Journal Story
Lions and Magic
Other Lion Articles
Book Dedication: T. Masland
Tell a Friend
Contact the Author
Shark Attacks
The Spirit Lions
Custom
e-mail me

The Science of Man-eating
 



The science of ‘Man-eating’* among lions (Panthera leo) with a reconstruction of the natural history of the “Man-eaters of Tsavo”

Julian C. Kerbis Peterhans1 and Thomas Patrick Gnoske2

1 Associate Professor, University College , Roosevelt University , Chicago IL ;

Adjunct Curator, Division of Mammals, Field Museum of Natural History

2 Assistant Collection Manager and Chief Preparator, Division of Birds, Field Museum of Natural History

ABSTRACT

The story of the "Man-eaters of Tsavo" has been retold through script, cinema, and oral tradition in the 100+ years since their infamous 'reign of terror'. Despite their predictably broad popular appeal, the details pertaining to the natural history of these lions have never been reviewed. The skulls and skins of these lions have resided at the Field Museum of Natural History (Chicago) for over 75 years. An analysis of the skull of the primary culprit displays a traumatic injury that may have limited his predatory ability in subduing ‘normal’ prey. A sample of hairs, reflecting the diet of both man-eaters, is preserved in the broken and exposed cavities of their canines. Various additional circumstances likely contributed to their man-eating habit. The Tsavo incident closely followed the debut of rinderpest on the continent, which devastated cattle and buffalo, the primary prey of the Tsavo lion. The Tsavo ‘nyika’ consists of a dense thorn scrub thicket limiting visibility and passage, representing an ideal habitat for an “ambush predator”. Finally, historical review of the literature reveals that 'man-eating' was not an isolated incident at Tsavo. This behaviour was well established in the vicinity of the railway bridge well before these infamous lions appeared, and continued well after their demise, suggesting a recurring opportunity, which may have evolved into a local behavioural tradition. In sum, virtually all of the recognised preconditions for man-eating outbreaks to occur were in effect at Tsavo in the 1890's.

* We use the term 'man-eating' by convention, as attacks are not limited to 'men' and do not always conclude with consumption.

Introduction

The "Man-eaters of Tsavo" first gained infamy in the British press in 1900 (The Spectator, March 3) when they were reported to have caused a temporary halt to the construction of the ‘ Uganda ’ Railway. This railway was to run from the Indian Ocean port of Mombasa, Kenya, to the shores of Lake Victoria, which was then located in Uganda. British engineer J.H. Patterson documented his attempts to dispatch the two adult male lions with detailed journal entries (J.H. Patterson, 1898-1899). These entries became the basis for his first-hand account, first published In the Field (London) at the turn of the century, and later as the basis for his book, The Man-eaters of Tsavo and other East African Adventures (Patterson, 1907). Every Kenyan schoolboy knows the story of the Tsavo lions, first through oral tradition and later through this popular reference. Two Hollywood films have further popularized the story: Bwana Devil (1953) and The Ghost and the Darkness (1997). Despite the popular appeal of this incident, no details concerning the lions themselves, or the circumstances surrounding this notorious man-eating outbreak, were published * between Patterson's 1907 account and the brief review by Kerbis Peterhans, et al. (1998). After reviewing historical accounts, recent literature, the original journals of J.H. Patterson, Game Department records, and the skulls and skins of the ‘Man-eaters’ themselves, we have identified these circumstances. This paper addresses some of the conditions that lead to man-eating in general and how these relate to the incidents at Tsavo over 100 years ago.

Several circumstances have often been associated with Pantherid (big cat) attacks on humans. Sickly, injured, or emaciated pantherids, which are unable to secure their normal prey, can often become man-eaters (Corbett, 1944, 1948, 1954). Predators may also focus on abnormally behaving prey, with certain human behaviors probably qualifying. Among social carnivores (e.g. lions), the favouring of particular prey species (e.g. livestock or humans) can be passed from one generation of predators to the next, with the potential to become a local social tradition (Swayne, 1895; Taylor, 1959; Rushby, 1965). Historical records from Tsavo suggest that predation upon humans was a long-standing phenomenon. Humans were attacked and killed * A probable exception is Preston ’s undated publication, The Genesis of Kenya Colony.

by lions in the Tsavo area well before construction of the railway began (1886, documented in Jackson , 1894) and continue at present, over 100 years after the infamous pair was dispatched (Kenya Wildlife Service, 1994-1998). Another factor, anecdotally mentioned in the literature, is the absence, depletion, or removal of typical prey, causing the predators to seek alternative food sources such as livestock. This brings lions into contact with humans, sometimes resulting in the inclusion of humans in lions’ diets. Lions also develop a taste for humans after being 'provisioned' with dead ones. Environmental variables can be important since heavy cover is essential for “ambush predators” to successfully stalk their prey (Schaller, 1972; Funston, et al. 2001). Seasonal factors are also thought to be important as game disperses during the rains, making them more difficult to locate and secure ( Jackson , 1894). In late 19th century Tsavo, all of these factors were in play concurrently and all may have played a role in the development of the world's most renowned man-eating outbreak. What follows therefore, is a critical review of these different scenarios and the likely role each of them may have played in the events at Tsavo over 100 years ago.

Notes on the Natural History of the 'Man-eaters of Tsavo'

Unlike the Tsavo of today with large tracts of open expanse, the Tsavo of the 1890's was composed of a nearly impenetrable, thorn thicket known as nyika. The quest for ivory during the 19th century had eliminated elephants (Loxodonta africana) from much of eastern Kenya , including most of Tsavo (Thorbahn, 1979). This is why many of the porters transporting ivory through Tsavo at the time, originated far up-country, sometimes even from Uganda (Patterson, 1907). Elephants are a keystone species and have a major impact on the vegetation and the large mammal community. The elimination of elephants from Tsavo caused a proliferation of dense woody and thorny undergrowth and eliminated herds of grazing ungulates from the vicinity. Browsers, including dik-dik (Madoqua sp.) and black rhino (Diceros bicornis) increased. A review of Patterson's field journals (1898-1899) quantifies the animal species he encountered (TABLE 1). There were no wildebeest (Connochaetes taurinus). Hartebeest (hartebeest/hirola, Alcelaphus buselaphus/hunteri) and zebra (Equus burchelli/grevyi) were around but not common. Patterson never once refers to buffalo (Syncerus caffer) or cattle (Bos taurus ssp.) in his journal. This can be traced to the arrival of rinderpest on the continent in 1891, which had a devastating impact on the bovine populations (Lugard, 1893; Mettam, 1937). By the 1960’s, the Tsavo elephant population had recovered and Tsavo became known as one of the largest elephant sanctuaries in East Africa (Leuthold and Leuthold, 1976). Therefore, the two large mammal species (elephant and buffalo) comprising most of the mammalian biomass in Tsavo today (Ibid.) were virtually absent in the 1890’s. In sum, by 1898, Tsavo had already been severely impacted by humans, resulting in vast anthropogenic differences between the Tsavo environment of the 1890's and that of today.

The 'Man-eaters of Tsavo' themselves consist of two adult male lions that are virtually devoid of manes in the conventional sense. They display a slight sagittal crest of hair that can be typical of some maneless lions. The second lion killed (FMNH 23969, FIGURE 1) had slight cheek tufts as well as darkened chest patches while the first lion was without these adornments. They were photographed by Colonel J.H. Patterson (1907) after they were dispatched in 1898. Except for the severely broken canine and remodeled mandible and cranium of the primary culprit (FMNH 23970; FIGURE 2), the man-eaters of Tsavo were adult male lions in their prime

TABLE 1

FAUNA OF TSAVO AS REPRESENTED IN THE FIELD JOURNAL OF JH PATTERSON, 1898-1899

Number of                                                       Number of Times                    Times

Group                                             Mentioned                                                       Mentioned

Common name

Invertebrates                                                              Mammals

Tsetse fly                                 1                                  black panther               1

Mosquito                                 1                                  baboon                                    1

2                                  rabbit                           1

Reptiles                                                                           rat                                1

Crocodile                                 1                                  giraffe                          1

Birds                                                                                   donkey                                    2

long-billed                               1                                  monkey                       2

water fowl                               1                                  hippo                           2

owl                                          1                                  waterbuck                    2

spur fowl                                 2                                  bushbuck                     3

sand grouse                             3                                  leopard                        3

ostrich                                     4                                  zebra                            4

birds                                        5                                  wild cat                        4

partridge                                  6                                  hartebeest                    4

guinea fowl                             12                                small deer                    5

35                                goat                             11

rhino                            14

paa (Madoqua)            16

lion                              70

147

 

 

 


6 ? - 8 ? years of age). Their squamoso-parietal sutures are obliterated making them at least 6 1/2 years old (Smuts, et al. 1978), but their maxillo-premaxillary sutures are not even partially closed, making them under 9 years old. Their teeth also betray this age range as they are yellowing and there is visible wear on the canine, incisor, P3, and P4 (>5-6 years of age, Ibid.). Although these individuals may have been siblings due to their association and similar size and morphology, FMNH 23969 displays more apical wear than FMNH 23970. We doubt that their difference in age extends beyond one or two years.

Both animals were huge individuals as Patterson’s published photos (1907) demonstrate. Patterson uses the following in describing the first lion: “enormous brute”, “powerful beast in every way", (Patterson, 1898-1899). FMNH 23970 was 9’8” long (nose to tail) and 44” in height at the shoulder while FMNH 23969 was 9’6” long and 48” in height at the shoulder (Ibid.). Shoulder heights for East African male lions average 37.8" in height and 8'9" in length according to Meinertzhagen (1938) who collected in the Athi-Kapiti Plains and in the Serengeti. Guggisberg (1975) provides an additional measurement: 3’ in height at the shoulder and 9’ long.

Patterson claimed (1907) that the man-eaters were prime-aged, healthy males and was so cited by subsequent authors (e.g. Selous, 1908; Akeley, 1923; Bradley, 1926; Guggisberg, 1961; Cloudsley-Thompson, 1967). We can only assume that this referred to the condition of their limbs and torso and that he did not inspect their dentition. Residing in the collections of Field Museum for 75 years, the two Tsavo lion skulls had not been differentiated from one another in any way. One of us (TPG) rediscovered these skulls in the collections of the Field Museum in the 1980's and subsequently deduced which was the first man-eater shot by referring to Patterson’s discussion of the deaths of each (Patterson, 1907). The first lion (FMNH 23970) was killed by shots to the body, while the second lion was shot with several bullets, including one to the head that shattered its zygomatic arch (FMNH 23969). Although Patterson states that he may

have shot off the tip of the canine of the first man-eater slain (Patterson 1907, 1925), both lions are missing the apical end of their lower left canines due to pre-existing traumas

One of us (TPG) first noticed that the skull and mandible of FMNH 23970 were malformed due to a severely broken canine with exposed pulp vacuity. This observation was first noted in Kerbis Peterhans et al. (1997) with further details provided by Neiburger and Patterson (2000). This injury probably occurred fairly early in life as it led to obvious remodeling of the jaws (FIGURES 3,4). In particular, the root of the broken lower canine had been re-oriented towards the horizontal plane, and the mandible and associated dentition were completely asymmetric due to mal-occlusion. This injury (perhaps caused by a kick or a blow from the head/horn of a buffalo or zebra) could have prevented the lion from efficiently killing its normal prey.

The second man-eater (FMNH 23969) had slightly damaged teeth, specifically a broken upper left carnassial and broken lower right canine (photo in Patterson, 1907). The break to the upper left carnassial (P4) is fairly fresh (several months in age) as reflected in its still sharp edges. The broken canine tip had been worn smooth from months, perhaps years, of wear. Despite these breaks, we do not believe these injuries were serious enough to affect this animal’s predatory behavior.

Their tenure as man-eaters may have been first discussed by Ansorge (1899) who discussed lion attacks on a caravan in 1896 at the very crossing point of the Tsavo River over which the railway bridge was subsequently constructed. At this time, the Tsavo man-eaters would have been approximately 4 1/2 to 6 1/2 years old, full-grown ‘sub-adults’. R.O. Preston followed Ansorge at the river crossing but preceded Patterson. Preston arrived in Tsavo in January of 1897 (Miller, 1971) and promptly recorded the deaths of two railhead workers due to lions (Preston, n.d.). J. H. Patterson arrived in March of 1898, and in late April 1898, attacks were again reported. By the time the ‘reign of terror’ was over, Patterson stated that the lions “had devoured between them no less than twenty-eight Indian coolies” (1907, p.107). The figure of 28 was cited by subsequent authors (Selous, 1908; Guggisberg, 1961; Miller, 1971). The first man-eater was killed on December 9, 1898 and the second on December 29, 1898 (Patterson, 1898-1899). Despite a gap of 20 days between the deaths of the two lions, there were no humans from the railway crew taken by the remaining man-eater. This leads us to believe that the first individual slain (FMNH 23970), with the long-term dental trauma, was the primary culprit. If we add the two deaths recorded by Preston prior to Patterson’s arrival and the death discussed by Ansorge, the documented total could increase to 31.

Between March and December of 1898, a minimum of 28 humans was taken by these notorious lions. Over the years however, these figures have changed and the legend has grown. In his journal entries (1898-1899) Patterson only discusses the deaths of 14 individuals. In his 1907 book, he mentions 28 deaths of the imported and well-paid Indian laborers. We assume this figure is accurate since the British maintained precise accounting details (many of which are summarized in Hill, 1976). His only reference to victimized African laborers is summarized as “scores of unfortunate African natives of whom no official record was kept” (1907, p.107). In 1925 however, Patterson published a follow-up account where he states that 107 Africans were among the victims, thereby increasing the total killed on his watch to “135 Indian and African artisans and laborers” (p.1). Neiburger and Patterson (2000) expand the description to “135 armed men”. Later, these lions are described as: “evading exposure to modern hunters with high-power firearms” and taking “135 well-armed railroad workers” from “well-fortified camps” (Neiburger and Patterson, 2002). Caputo (2000) expands the carnage to 140 people. The fact is, that there is no published record indicating that any more than 28 were officially recognized as being victims, although our review of the historical literature can possibly add three more.

If we are to assume however, that the figure of 135 is accurate, and further assume that 135 people would provide ca. 50 pounds of edible meat per individual, the lions would have been left with 6750 pounds of meat over the 10-month period. Schaller (1972) has estimated an adult male lion’s daily intake to be ca. 15.4 pounds of meat per day, resulting in 5605 pounds per year, per lion, or 9,342 pounds for both adult male Tsavo lions for the 10-month period. Although these figures are well within an order of magnitude, there is no convincing evidence in Patterson’s journals that this many people were killed by lions during his tenure in Tsavo. Furthermore, it appears that ‘normal’ prey were also consumed over this time period.

Although depicted as dedicated man-eaters, both lions came to bait and attacked livestock. The first lion slain approached a dead donkey used as bait, while the second carried off 6 tethered goats over a two-day period (17-18 December, 1898; Patterson 1898-1899). The severe injury to FMNH 23970 did not prevent it from feeding on ‘normal’ prey afterwards. Studies of their dietary preferences are currently being conducted through the analysis of several thousand animal hairs, removed from the broken canines of the man-eaters. Preliminary results provided by Ogeto Mwebi, indicate that these hairs are mostly from the lions themselves, lodged in the canine vacuities during bouts of grooming. Mwebi (personal communication) has tentatively identified the following prey species: zebra, porcupine (Hystrix cristata), warthog (Phacochoerus aethiopicus), impala (Aepycerus melampus), eland (Taurotragus oryx), and oryx (Oryx gazella). To date, there is no evidence of human hair, which suggests that immediately following the break to the canine of the first man-eater, humans did not suddenly appear on the menu. If we find do evidence of humans, we will be able to distinguish African from Indian victims. However, since lions prefer human viscera and large fleshy parts, including the buttocks, thighs and arms (see figures of human victims in Kingsley-Heath, 1965; Beard, 1988), hair bearing areas such as the cranium and perhaps the pubis may not be consumed. This may explain the absence of human hair in our samples. What is clear is that these lions were not obligatory man-eaters and that they continued to pursue prey other than humans.

I. Fossil Record and Modern History of Man-eating

For as long as they have co-existed, primates, along with ungulates, have been the primary base of prey for African Pantherids (Brain, 1981). For most of their history, extinct and living Hominids have represented little more than a vulnerable, slow moving, bipedal source of protein for the big cats. In the Plio/Pleistocene of South Africa, Australopithecus robustus was the single most common prey item for leopards (Panthera pardus) with a minimum of 88 individuals represented at the site of Swartkrans (Brain, 1981). The newly discovered South African site of Drimolen promises similar results (Keyser, et al. 2000). The earliest reputed hominid fossil was also said to have been killed and dismembered by a carnivore (Fox, 2000). While Homo sp. seems to have been more effective in avoiding predation than Australopithecus, based on fewer remains in fossil lair sites (Brain, 1981), early Homo, nevertheless, remained on the menu. In fact, paleoanthropological material from South Africa, which has provided most of the fossil evidence for the early stages of

human evolution, appears to have been primarily accumulated by large predators, particularly Pantherids and Hyaenids (Brain, 1981; Kerbis Peterhans, 1990).

European Paleolithic rock art depicts cave lions embedded with arrows or spears (Figure 1 in Frobenius,1933; Begouen and Breuil, 1958; Ruspoli, 1987). One of the earliest historical depictions of man-eating that we have found is the nearly 5,000 year old "Battlefield Pallette" depicting a lion eating, and/or killing, dead or wounded Libyans during a clash with the Egyptians (Aldred, 1980). Bushmen rock artists illustrate numerous scenes of lions dismembering humans as well as human retaliation (Tongue, 1909). Stow (1905) writes that lion fed upon the flesh of Bushmen even more than their sheep. With the advent of the colonial era in Africa and Asia, documentation of man-eating became more regular. This coincided with the exploration of continental interiors in the 19th century, the debut of “big game trophy hunting”, and the construction of inland railways in the late 19th and early 20th centuries (Sowerby, 1923; Hill, 1976).

Man-eating is not unusual, nor necessarily 'aberrant'. From the late 19th century up until the present, man-eating incidents by Pantherids in Africa and Asia have continued. During a 5-year period in the 1920's, 7,000 human deaths due to tigers (Panthera tigris) were reported in India (McDougal, 1987). The Sanga man-eaters killed over 161people in SW Uganda in the mind to late 1920’s (Temple-Perkins, 1955). In the 1930's, lions killed around 1500 people in a 150 square mile area of southern Tanzania (Rushby, 1965). Between 1978 and 1984, tigers killed a minimum of 128 people in Kheri, a small Indian District in Uttar Pradesh. Three hundred eighteen people were killed by tigers in the mangrove swamp forests of the Sundarbans Tiger Reserve between 1975 and 1981 (Sanyal, 1987). A minimum of 193 lion attacks resulting in 28 human deaths, were recorded from areas adjacent to the Gir Forest (India) between 1977 and 1991(Saberwal, et al. 1994). In sum, human and Pantherid conflicts have existed throughout their coexistence with evidence for sustained and localized outbreaks of attacks on humans.

II. Access to injured, sick or dead humans

a. Slave and Trade Caravan Routes


Most carnivores, especially lions and hyaenas (Crocuta crocuta), are facultative predators and scavengers, actively hunting or scavenging depending on the circumstances (Schaller, 1972; Kruuk, 1972). The slave trade in East Africa promoted the consumption of humans, as dead or dying slaves were abandoned along caravan routes. A slave trader told A.J. Swann (cited in New, 1874) that any slave too weak to carry his load was abandoned or killed at once in order to discourage others from refusing to carry their burdens. During Livingstone’s travels along caravan routes in central Africa, he often encountered human remains. He estimated that only one in five slaves reached the coast alive (cited in Miller, 1971). With 20,000 slaves imported each year to Zanzibar alone, there was probably a minimum of 80,000 humans lost annually along the northern

caravan routes. This huge number of abandoned, sickly, dying, and dead, represented a regular bonanza to any carnivore prepared to take advantage of them.

The ‘Uganda railway’ was built along a Swahili caravan route that had been used for decades. The caravan route maps of Wakefield (1870) are quite similar to those of Johnston (1899) and the railway line is shown to closely parallel the route used by ‘explorers and caravans’ (Hill, 1976). At the Tsavo River, Patterson (1907) describes his tent as being pitched close to the former caravan

route to ‘Uganda’. He provides a photograph of the crossing point of the caravan trail over the Tsavo River (FIGURE 5). Preston (n.d.) discusses this same caravan trail, stretching along the right (south) bank of the Sabaki River.

Trade routes also generate extensive “ debris fields” and condition the local carnivores to a highly predictable food source. People and goods moving along the caravan route included human slaves, porters, ivory, luggage, and foodstuffs. Coast-bound Wakamba traders transported livestock including sheep (Ovis aries), goats (Capra hircus) and cattle. In the late 19th century, Eurpoean expeditions joined the mix, travelling along the same routes as those plied by Arab and African traders. First hand accounts of these caravans include those by French-Sheldon (1892), Lugard (1893), Neumann (1898), Ansorge (1899), Patterson (1907), and Preston (n.d.).

Despite proclamations banning the international trade in slaves (1873, 1876), and the similar edict in 1890, an illicit slave trade continued through the Tsavo region during the late 19th century; slave ownership was not banned until 1907 (Miller, 1971). One of the reasons the British claimed to have constructed the railway was that its development would help to put an end to the illicit slave trade, which still flourished in the late 19th century:

‘After a railway has existed for some time there cannot be....any other kind of locomotion to compete with it....If a railway could exist from this lake to the coast, caravans could no more be employed as they are employed now to carry ivory’ (Lord Salisbury, 1891 quoted in Hill, 1976, p.54).

It was reasoned that an efficient and rapid alternative in the transport of goods would eliminate the profitability of all caravans.

However, even the end of the formalised slave trade did not end the regular loss of human life along this route. Selous (1908) depicts lions posted around campfires waiting for opportunities (FIGURE 6). Porters hired to carry ivory and trade goods were abandoned in the bush after dying from exposure (Portal, 1894) or suffering injuries or disease (French-Sheldon, 1892; Lugard, 1893). French-Sheldon’s caravan came across an abandoned ‘ill wretch’ at Buru in 1891, ‘dying from hunger and neglect who was unable to proceed with a caravan....and turned adrift, without adequate means, to reach the coast as best he could or drop dead in the bush’ (p.203). At the end of a long days march, porters often limped into their camps with broken bones and severe injuries (French-Sheldon, 1892). Lugard (1893) describes liberating slaves in a caravan he encountered along the Voi River (Tsavo) in 1890, and speaks of a paralysed ‘porter’ being abandoned to hyenas at the Voi River. He concludes, ‘I have never seen anything approaching the carelessness of human life and the callousness to human suffering which seem to characterise some methods of African travel’ (Vol II, p. 544).

French-Sheldon (1892), Ansorge (1899) and Patterson (1907), passed westward along the Tsavo caravan route and criss-crossed with caravans moving east. In Patterson’s chapter, entitled ‘The Stricken Caravan’ (1907), he describes a caravan of 4,000 porters (Basoga and Baganda) moving from Uganda to Mombassa in 1898. On their return to Uganda, the entourage again passed Patterson’s camp on the Athi-Kapiti Plains. An epidemic of dysentery had swept through the contingent, perhaps linked to their drastic change in diet along the way. After each day’s march, dozens were left along the route; the group could not stop as food and water were always at a premium. Thirteen dropped out in the vicinity of Patterson’s tent and despite his best efforts, only seven were saved. Along the track, Patterson estimated finding a swollen corpse every 100 yards. When passing one of their abandoned camps, Patterson encountered about 12 fresh graves that had already been disinterred by scavenging hyenas.

b. Burial practises, epidemics, warfare


Local burial practises can also encourage the development of man-eating behaviour by providing easy access to human corpses. The Masai simply abandon their dead, and sometimes the 'near dead', in the bush to hyenas and lions (also Kikuyu, Percival, 1925; Saitoti, 1980; Read, 1984). French-Sheldon (1892) describes coming across the corpse of a deceased Masai, weighted in armlets and leglets. Selous (1908) describes an incident near Bulawayo, Zimbabwe where an elderly woman was dragged alive and tied to a tree for the hyaenas. In the same volume Selous depicts hyenas hovering around an abandoned corpse (FIGURE 7). Melliss (1895) describes picking up an elderly woman that could not keep up with her clan, and who was left to fend for herself along a Somali caravan route. Gouldsbury (1916) writes of the Somali:

"Women generally, and more especially when old and decrepit, are of very little account among the Somali. If unable to keep up with the Kafila <caravan> on the march, they are often abandoned and left exhausted on the side of the road, either to follow as best they can or to be devoured by a hungry lion, should one happen to pass that way" (p.167).

In the Taita Hills, above the Tsavo caravan route, the Wataita dig up their dead and then place skulls in rock shelters, while the people of Taveta disperse the bones of the dead in the bush and place the skulls in sacred tree hollows, rock shelters, or earthen pots (French-Sheldon, 1892; Adamson, 1967). The Wakamba peoples inhabiting the Tsavo region were said to leave the bodies of ‘peasants and women’ and even the mortally infirm in the bush ‘for the hyaenas to devour’ (Hobley, 1910; Lindblom, 1920).

In India, Corbett documents several instances of man-eating outbreaks by leopards and tigers. These include the Rudrapryag Leopard, which killed 150 humans (1918-1926) following an influenza outbreak that killed over one million people (Corbett, 1948), and the Panar Leopard which killed 400 humans (1905-1907) following a severe cholera outbreak (Corbett, 1954). Many Hindu families cannot afford the expensive funeral pyre necessary for a proper cremation. Instead, the dead are abandoned with a red-hot coal in their mouth. It is possible, but doubtful, that Hindu railway workers dying in Tsavo were disposed of in this fashion. There was no shortage of firewood; lions were kept at bay during the night with non-stop fires. In any case, not all of the imports were Hindi, as Patterson himself (1907) speaks of the rivalries between Muslim and Hindi laborers.

Between 1898 and 1899, an outbreak of small pox and famine in Kenya killed thousands of Kikuyu and Ukumbani. The dead and near dead were abandoned in the bush to be devoured by carnivores (Percvial, 1925). Once the diseases abated and the supplies of corpses ran out, the large numbers of hyenas turned their attentions towards living humans. Hyenas started taking children at dusk and grabbing adults as they slept (Percival, 1925). Miller (1971) estimates that 25,000 Africans were felled by disease and starvation during this time (1898-1900). Such inadvertent provisioning of carnivores has often been correlated with attacks on humans. Even today, provisioning of hyenas by tourist outfitters can lead to human fatalities (Singer, 2000).

Patterson (1907) and Hill (1976) again provide analogies for the situation in Tsavo in 1897-1898. A severe famine, caused in part by a severe drought caused thousands of local Wakamba to die of starvation during the time the railway was being built through Tsavo (Miller, 1971). In their desperation, Wakamba would attack and slaughter isolated railway maintenance gangs in order to steal their food (Patterson, 1907). Death among the Wakamba due to a smallpox outbreak was also devastating at this time. Despite massive relief efforts, there was not enough treatment (lymph) to go around (Miller, 1971). The human toll among the Indian coolies imported to work on the railroad was also substantial: 340 of 7,131 died between 1897 and March of 1898. Death was mostly due to malaria, diarrhea, and dysentery; an additional 705 were debilitated (Hill, 1976), with many being returned to India.

During civil unrest and conflict, large Pantherids have been known to kill humans after having regular access to human corpses. As discussed earlier, the 5,000-year-old ‘Battlefield Palette’ depicts lions consuming humans following a battle in Egypt (Aldred, 1980). In the Arakan district of Burma, tigers had not previously claimed many human victims but during World War II, they suddenly turned their attentions to live and dead soldiers (McDougal, 1987, 1993). Caputo (personal communication) describes at least three instances of tiger attacks on humans during his time in Viet Nam (1966-1969), an observation confirmed by Jackson (1985).

A similar scenario was been documented in Tsavo during World War I (Lettow-Vorbeck, 1920). The Germans, colonial powers in Tanganyika, and the British, who colonised Kenya, had military conflicts in the vicinity of Tsavo National Park with lions also taking their share. General Phillips, based in Voi for three months in 1915, lost three Indian soldiers to lion in the vicinity of the Tsavo River (Trzenbinski, 1986). When the war began, noted game ranger A. B. Percival was charged with defending the railway against sabotage from the Germans. Sentries were posted and according to Percival (1928):

‘Lions were a veritable curse; man after man on sentry duty was taken, till it seemed sheer cruelty to put a man on such duty at all. Sentries were doubled and still men were taken. It needs little imagination to realise the case: two men at their post, bush all round and close up-it was inviting attack by lions’ (p. 286).

During WW I, the wounded and deceased not recovered by nightfall, were assumed to be taken by this carnivorous entourage (Percival, 1928).


 

III. Predation on Livestock: Human/Carnivore conflict

a. Historical sources

The initial attraction of large carnivores to humans can also center on a supply of livestock and pack animals. Herodotus may have been the first to discuss lions killing domestic stock (baggage camels, Camelus dromedarius), when they were described attacking Xerxes' caravans during his trip through Paeonia (Rawlinson, 1992). In Africa, the first long-horned, Nile valley cattle (Bos primigenius) were domesticated by 4,000 BC (Rouse, 1972). The first long-horned, humped cattle (Bos taurus indicus) arrived in Africa about 2,000 B.C., when they crossed from southern Arabia into the horn of Africa. The presence of domestic stock, particularly cattle, contributed to future associations of livestock, humans and lions.

After coming into contact and conflict with humans while killing domestic stock (Swayne, 1895; Neumann, 1898; Borradaile, 1928), predators educate themselves on the sights, sounds, and activity patterns of humans; they may switch to herdsman and humans in general (Layard, 1887; Caldwell, 1925; Percival 1925, 1928; Pitman, 1931; Hunter, 1952; Guggisberg, 1975; McDougal, 1987). Four of the eight man-eating tigers discussed by Corbett (1944, 1948, 1954) were also notorious cattle killers, confrontations that led to direct conflict with humans. For example, the Chuka man-eater (Corbett, 1954) attacked and killed cattle. On one occasion, it was disturbed by, and subsequently killed, a young herdsman looking for his cattle, thereby beginning his man-eating career. The Chowgarh man-eater alternated between preying upon cattle and humans and ultimately became more focused on humans after its large cub was killed. This cub had assisted her in bringing down cattle.


Swayne (1895) discussed a parasitic relationship between humans and lions in Somalia. This revolved around a detailed calendar of caravan movements which was influenced by shipping schedules and seasonal availability of caravan-transported goods:

"The movements of the native encampments seem chiefly to influence the changes in the quarters of the lions, the latter following the karias <herds> as they move to fresh pasture. When a family with its flocks and herds and its karias, moves, its attendant lions, if there should be any, accompany it, being sometimes man-eaters and more often cattle-eaters. Last June my own caravan, while returning to the coast from Ogaden, was followed by a pair of hungry lions. We discovered this by chance, when some scouts of mine, happened to go back along the road" (293-294).

Pitman (1931) describes an incident in SW Uganda in 1928 when four lions claimed 15 cattle in a few weeks. Little attention was paid to this incident and due to a communication gap, nothing more was heard for several months. Within 9 months, humans were included among the victims. Before they were destroyed, they had claimed more than 250 head of cattle and no less than 22 humans.

In historic Tsavo, at least three circumstances brought about massive movements of people and their accompanying livestock: caravans, the building of the railway, and troop deployments during World War I. When mass movements occur, infirm and injured livestock and food debris are left to the carnivores. The situation for Tsavo was particularly acute, as the region is known as the ‘nyika’, a dry, thorny shrubland with particularly low prey density (Leuthold and Leuthold, 1976).

French-Sheldon (1892), Lugard (1893), Ansorge (1899), and Patterson (1907) all state that caravans, including those passing through Tsavo, were routinely stalked by hyenas and/or lions, attracted by the food refuse as well as the livestock (sheep, goats, cattle, and donkeys (Equus asinus) that often accompanied these expeditions. French-Sheldon (1892) writes:

”When we camped for the night we were obliged to form a hedge of thorn-bushes and circle the encampment with huge bonfires to keep the wild beasts from attacking us. It was terrifying to hear the continuous roar of lions resounding on all sides.....and to see the glare of hyena eyes in the darkness of the umbrageous surroundings. A sense of abject helplessness momentarily possessed me….”(p.162).

Bringing large numbers of domestic ungulates into a naturally prey-depauperate region as Tsavo, is an obvious attractions to the local carnivore community.

During the mid-1890's, advance railway crews in the Tsavo area were charged with clearing bush and the construction of embankments ahead of the Uganda Railway. Hill (1976) writes that in 1897-1898 over 90% of the 1502 camels, donkeys and cattle used for transport had died. These animals were victims of tse tse flies, lack of water in the Taru desert, (Patterson, 1907; Hill, 1976) and other ailments. During World War I, there was substantial mortality among the thousands of transport animals fuelling the war effort against the Germans in the Tsavo region. The British were actually grateful that carcasses of dead and dying transport animals (up to 20 per day) could simply be abandoned in the bush, 100 yards from camp. Lions and hyenas followed in the wake of troop movements and ‘were assured of a feast every night’ (Percival 1928, p. 295). Percival states that, with such inadvertent provisioning, the boldness of the lions and hyenas around army camps grew. He continues:

‘When on safari in the reserve in 1919-1920, I was struck by the boldness of the hyenas; they hung round the camp at night, coming to such close quarters that they kept the dogs barking incessantly. I felt quite sure that this unusual temerity, greater than I have known the brutes to display anywhere else, may be explained by their experience during the war' (p.295).

Percival portrayed lions of the coastal region as particularly large and aggressive. Soldiers mounted on horseback came across lions that:

’learned that food in the shape of horses abandoned on account of injuries or sickness were to be had without exertion on their part, and they dogged the mounted men day and night. As this was near the German lines, firing was strictly forbidden; hence the lions grew bold and exceedingly troublesome. They would come fearlessly up to the very outskirts of a camp’ (p.287).

b. Human/carnivore conflict in Tsavo: species profiles, sex, age, and seasonal data

In Kenya today, pastoral peoples can be compensated for the loss of domestic stock. Kenya Wildlife Service has established ‘incident report’ log-books that document the circumstances surrounding conflicts between carnivores and humans. Between 1994 and 1998, a total of 121 incidents between large carnivores and humans and their livestock were registered by KWS officials based in Voi (Tsavo East National Park, TABLE 2). Lions were responsible for 93% of these incidents. Collectively, cheetah, leopard and hyenas accounted for only 7% of the incidents; they exclusively attacked sheep and goats. In areas where they co-occur with lions, leopards typically select for smaller prey (Kerbis Peterhans, 1990; Fay, et al. 1995; Treves and Naughton-Treves, 1999). Conversely, lion predominately attacked cattle (63%), with sheep and goats secondary (27%).

Specific data for lions is detailed in TABLE 3. Male lions attacked cattle more often (41/71=58%) than female lions (30/71=42%), while females preferentially attacked sheep and goats (18/30=60% vs. 12/30=40%). Multiple killings were relatively common. Females with cubs killed more individuals per attack than either male lions or females without cubs, averaging 4.8 sheep/goats and 2.5 cows per attack. Male lions averaged 3.25 goats and 1.6 cows per attack, while solitary females killed an average of 1.4 sheep/goats and 1.3 cows per attack. Female lions with young were more prone to multiple kills, taking at least four goats on six occasions (on one occasion 20 were killed); 3-6 cows were killed on eight occasions. One male lion killed 18 goats, while individual leopards killed eight and 10 goats on separate occasions, obvious examples of 'surplus killing' (sensu Kruuk, 1972). According to Kenya Wildlife Service archives, the most recent incidents of ‘man-eating’ at Tsavo East N.P. were recorded in October 1994 and July 1998.

Assuming an equal sex ratio, male lions were more likely to attack domestic stock than females (61 vs. 51 attacks). Data from most African localities indicate that adult female lions typically outnumber adult males by ratios ranging from 1.4:1 to 4:1: Lake Manyara (Makacha and Schaller 1969: 3 to 1), Kafue (Mitchel et al. 1965: 1.7:1), Serengeti plains (Adamson 1964: 3:1, Schaller 1972: 2:1), and Kruger National Parks (Anonymous 1960: 1.4:1). Schaller (1972) describes more equal, but still male-heavy sex ratios in the Serengeti woodlands while Rogers (1974) describes the same for lions in Selous National Park, Tanzania. Preliminary data from Tsavo (Russell, personal communication) indicate a predominance of females. Among 64 lion distributed between 5 prides, Russell only documented 10 adult males. This suggests that the impact of male lions in Tsavo is especially disproportionate to their numbers. However, according to KWS archives, female lion with dependent cubs can be especially destructive; they accounted for the highest number of kills per attack. Between 1994 and 1998, adult male lions were mostly responsible for attacks on humans in Tsavo East National Park, accounting for 5 of 6 attacks (83%) and both deaths.

Several authors have speculated on possible seasonal correlations of human/lion conflict. For eastern Africa, Jackson (1894) claims that attacks should occur after the rains, between March and the end of July, when prey are dispersed. Guggisberg (1961) restates this hypothesis, but provides no data. He asserts that during the wet season in Tsavo, lions become more troublesome for pastoralists. He attributes this to the dispersal of game making them harder to secure. He further adds that the rustling of the taller grass during this time of year alerts potential prey to the approach of predators. However, modern data from the Tsavo East incident report logs do not support Guggisberg’s assertion. The wettest months in Voi, Kenya (Tsavo East NP) are typically March, April, November, and December (Kenya Government, 1970). During the 5-year period for which we have data, there were a total of 29 incidents with lions reported during these 20 months (4 months x 5 years),

TABLE 2

Kenya Wildlife Service, Incident Log Books

1994-1998 representing 121 Incidents

Voi, Tsavo East National Park

 

# of attacks by:

Total # of attacks by Carnivora

(%)

Total # of cattle attacks (# killed)

# times sheep & goat attacked (# killed)

Attacks on people

D/I/T*

Other attacks

Lion

 

112 (93%)

71 (127)

30 (108)

2/4/2

3

Leopard

 

4 (3%)

 

 

 

 

 

Males

3

 

3 (21)

 

 

 

Females

1

 

1 (5)

 

 

Hyaena

 

2 (2%)

 

 

 

 

 

Male

1

 

1 (1)

 

 

 

Female