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
rhino 14
paa (Madoqua) 16
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 | 1 | | | 0/0/1 | |
Cheetah | | 3 (2%) | | | | |
| Male | 2 | | 1 (4) | 0/0/1 | |
| Female | 1 | | 1 (1) | | |
?
Death/Injury/Threat
TABLE 3
Condensed Lion Data,
Incident Log Books 1994-1998
Voi, Tsavo East
National Park
| Total # of
attacks | # times cattle attacked | # cattle
killed | # times
sheep/goat attacked | #
sheep/goat killed | Attacks on
people D/I/T* | Attacks on
others@ |
| 112 | 71 (63%) | 127 | 30 (27%) | 108 | 8 (7%) | |
Males | 61 | 41 (67%) | 66 | 12 (20%) | 39 | 2/3/0 (8%)
| 3 (5%) |
Females | 51 | 30 (59%) | 61 | 18 (35%) | 69 | 0/1/2 (6%)
| |
Females
w/young | 33 | 19 | 47 | 13 | 62 | 0/0/1 | |
Females
w/o young | 18 | 11 | 14 | 5 | 7 | 0/1/1 | |
Percentages calculated by Row rather than Column
* D/I/T Deaths/Injuries/Threats
@ other donkey,
camel, ostrich (Struthio camelus)
an average of 1.45 per
month (TABLE 4). There were a total of
83 incidents with lions over this same 5-year period during the remaining 8
months, actually a higher average of 2.1 per month. However, given the
anomalous nature of the rains in recent years, and the impact of El Nino, a
more thorough investigation would
look at the specific rainfall pattern for each month under consideration, an
angle we are currently pursuing.
However, if the season of vulnerability is
expanded to the continuous 5-month period suggested by Jackson (March-July),
his seasonal hypothesis is confirmed. The March-July data yield 59 lion
incidents over the 25-month ‘wet’ period (2.4/month) compared with 53 incidents
over the 35-month ‘dry’ period (1.5/month). In fact, four of the five busiest
months for ‘incidents’ occurred during these 5 months. This approach to
seasonal data is appropriate because the rains provide temporary pools of water
and new growth: these conditions extend beyond the period of actual rainfall
and encourage ungulates to remain dispersed.
We were hopeful that the field journal of Patterson (1898-1899) would
enable us to test the seasonal hypothesis, but he only documents 14 deaths at
Tsavo in 1898. Remarkably however, all 14 are limited to the typically wettest
months: March (n=4), April (n=4), November (n=3) and December (n=2), with the
slight exception of a single person taken on May 1. Patterson (1907) writes
that some months then transpired (apparently May-October) before lions resumed
their attacks. During this time however, he records 5 deaths from adjacent
areas, including two from the railhead (now advanced), one from Ngomeni (10
miles away), and two others, but none from his camp on the Tsavo River. Nine
deaths were not recorded in his journal and subsequent book. Although this
might appear to confirm the seasonal hypothesis, 1898 was an anomaly in
rainfall patterns. A severe drought began with the failure of the long rainy
season in April/May (Hill, 1976) greatly contributing to the aforementioned
starvation of the Wakamba.
IV. Man-eating: A social tradition
a. Prolonged, localized outbreaks
Although lions have diverse feeding preferences,
specific lion populations are known to specialize on specific prey species.
This indicates a behavioural tradition, which, in a social species, is passed
down from one generation to the next (e.g. a cultural tradition). For the
Busanga Flats, Mloszewski (1983) discusses ‘buffalo-selective predation’
reflected in lion proficiency in tracking buffalo herds, pulling down
individuals, ‘buffalo-killing proficiency’, and even in their ignoring of other
potential prey while seeking out remote buffalo. Other lion populations also
have distinct prey species preferences (e.g. pigs, zebra/wildebeest, reviewed
in Guggisberg, 1975).
In a few instances, lion specialization on
humans has been recorded. Rushby (1965) documented the most prolonged and
devastating outbreak of man-eating by lions in Africa in the Njombe District of
southern Tanzania (1932-1947). This incident may have started with the
establishment of a game free corridor (to prevent the spread of rinderpest,
discussed below) but its longevity is believed to be due to the development of
a social tradition. This was good cattle country, which is often favoured lion
prey. However, these lions became so fixed on humans that they ignored cattle,
even when carrying off herd boys from the kraals.
Between 1932 and 1940 there were no accurate
records of human victims, but detailed records were subsequently maintained. In
the sub-chiefdom of Wangingombe alone, 249 were killed between 1941 and 1946.
These figures account for victims from only one of three sub-chiefdoms where
the killings occurred. In the other two sub-chiefdoms, the killings were at
least as numerous. Additionally, there were killings outside these immediate
boundaries. Rushby concludes that for the entire 15-year period, the death
total was between 1000 and 1500. This is a remarkable figure since the area in
question was only 50 miles by 30 miles. These killings only concluded with the
death of the last of the man-eaters in 1947; in total, 15 suspected man-eating
lions were killed.
These predators conducted their activities with
a strategy that shows similarities with other incidents (Rushby, 1965). At
sundown, lion(s) stalked people in a village, and taking advantage of available
cover, approached as close as possible before a final rush. If two lions
attacked, two victims might be taken. Then, using relays, the lions would carry
their victims up to a mile away. They did not attack the same village on
consecutive occasions and their next attack could be up to 15 miles away. Since
their attacks persisted for 15 years, it is believed that at least 3
generations of man-eating lions were involved. The transmission of a
behavioural specialization (in this case a predatory one focusing on people)
from one generation to the next illustrates a social tradition. According to
Rushby: "There is no doubt that most of these lions were born and brought
up to man-eating" (1965, p.204). Repeated man-eating outbreaks occurred in
the Ankole District of Uganda between 1924 and 1930. The episode did not conclude
until the last of the 17 man-eaters was killed in 1930 (Temple-Perkins, 1955).
The head Game Warden claimed that heredity must play a role (Pitman, 1931).
Identical ‘kill and flee’ strategies were used by lions in Ram Hormuz (Persia)
where, according to Layard (1887), the culprit never appeared in the same place
for two consecutive days. Patterson (1907) repeats this discussion in reference
to the Tsavo pair. Often, man-eating outbreaks only stop with the extermination
of the responsible lions.
b. The Tsavo situation
As discussed earlier, man-eating in and around
Tsavo was not limited to the claim of 135 human deaths during Patterson’s
9-month stay while building the railway bridge over the Tsavo River. Instances
of man-eating occurred before Patterson arrived on the scene and continued
after he departed. To the best of our knowledge, the first recorded case of
man-eating in the Tsavo vicinity was that recorded by Jackson (1894). Jackson’s
camp was attacked by a lion that grabbed the head of a sleeping porter. The lion
succeeded in running of with the cloth head wrap that served to protect the
man’s head when carrying a load. This location is situated adjacent to Tsavo
East and West National Parks and was described as the “gameless wilderness
between Mount Kisigao and Mitati in the Teita country” (p.238). Based on the
records of birds collected by Jackson (Shelley, 1888), we estimate this date to
be between October and December of 1886. In the vicinity of Tsavo, in 1891, a
group of porters returned to camp missing the
trailing member of their party (French-Sheldon, 1892). While a search party was
being organised, they heard the final shrieks of their comrade.
Five years later (October, 1896), a party led
by Ansorge was travelling through Tsavo (Ansorge, 1899). The group was warned
by missionaries at Kibwezi not to stay in the vicinity of Ngomeni due to the
presence of ‘man-eaters’. In order to avoid Ngomeni, the group crossed the
Tsavo River and camped on the opposite side (right bank). Nevertheless, lions
attacked the group at night and one of the porters was carried off. After a
short chase, the man was retrieved 200 yards from the camp. The lion did not
return to Ansorge’s camp that night but evidently visited a Wakamba caravan
camp 30 minutes away. A Wakamba native was carried off and killed at this camp
and the entire party fled abandoning their provisions and some trade goods.
This could have been the same lion culprit(s)
responsible for the human deaths during the construction of the railway bridge
two years later. Two months before Patterson’s arrival in Tsavo, railway
engineer R.O. Preston recorded that lions killed two railhead workers. When a
search was conducted, they found only their heads, hands and feet and bits of
other skeletons (Preston, n.d.). Preston later writes that a total of 17
Punjabi workers were killed during his tenure in the vicinity. Less than one
year later, after both man-eaters were slain, J. H. Patterson remarked that a
total of 28 Indian workers were killed during the Tsavo phase of construction
(Patterson, 1907). It is not clear if these totals overlap or if they are
completely independent. There were no further human victims after the death of
the first man-eater in December of 1898. If we add the 17 Punjabi victims
mentioned by Preston and the single Akamba taken from Ansorge’s caravan in
1896, the total number of documented victims could rise to 46. However, given
his propensity to highlight the nature of the
carnage, we would find it difficult to believe
that Patterson did not include the 17 victims discussed by Preston in his
totals.
Despite the deaths of the two primary culprits near the bridge over the
Tsavo River, killings along the railway continued. In March of 1899, about 3
months after the second man-eater was shot, Engineer O’Hara, the head of a
road-building crew working between Voi and Taveta, was dragged out of his tent
and killed by a lion about 12 miles from Voi (Patterson, 1907; Preston, n.d.).
In June of 1900, the ‘Kima Killer’ pulled Superintendent C.H. Ryall out of a
railway carriage at Kima Station, 120 miles up the railway from Tsavo station
(Patterson, 1907; Anonymous, 1961). Man-eating lions had already taken a number
of victims in the vicinity of Kima, including one Asian and a number of
Africans, as well as from stations at Makindu and Simba (respectively, 65 and
80 miles from Tsavo station).
Man-eating instances were
again recorded in Tsavo during World War I. General Phillips, based in Voi for
3 months in 1915, lost three Indian soldiers to lions in the vicinity of the
Tsavo River (Trzebinski, 1986). Percival (1928) lost great numbers of sentries
guarding the railway lines from German saboteurs. Subsequently, man-eating
subsided in Kenya, and indeed was eliminated for a time, after the near
extermination of lions in the late 1910's and 1920's. Instances of man-eating
eventually returned on a sporadic basis beginning with a killing along the Tana
River in 1928 (Kenya Game Department, 1928): “For the first time in years an authentic case of man-eating was
reported on the Tana. A Somali was taken near Sankuri and eaten.”
During the 1950's, in the vicinity of Ngulia Camp in Tsavo West NP, a
man-eating lion remained for several years, and tried to carry off a worker
from a road crew during a camping safari led by lion expert Guggisberg (1961).
In 1965, J. Perrott (Kingsley-Heath, 1965) killed a man-eater on the Darajani
Track in Tsavo East. This lion had a porcupine quill embedded in its nostril.
J. Stanley (pers. comm.) describes an incident in 1972 when a herdsman was
killed by lions in Tsavo East N. P. Two recent deaths due to lion are recorded
in Kenya Wildlife Service archives (1994-1998). In sum, man-eating lions have
been active in Tsavo for over 100 years.
In Tsavo, other predators attacked people as
well. Both Lugard (1893) and Ansorge (1899) mention that hyaenas carried off
sleeping porters at night. The victims were often the infirm, emaciated and
abandoned (Ibid). In the Ngulia Hills of Tsavo, a leopard that had been
provisioned as a tourist attraction became a man-eater in the 1980’s (Isiche,
personal communication).
V. Selection of Abnormally Behaving Prey
Lions often select ill or abnormally
behaving prey (Schaller, 1972). Several examples suggest that lions also select
‘abnormally' behaving humans. For a lion, abnormal human behavior might include
the deranged or inebriated. Capstick (1981) describes a man-eating outbreak in
Zambia that began with the killing of the “village idiot” from the town of
Kalundi. Late night drunkenness coincided with a man-eating outbreak in and
around Queen Elizabeth National Park (Uganda) in 1993. Thirteen adult male
humans were taken by this adult female, with excellent teeth, known as the
Kazinga Man-eater along the Kazinga channel in western Uganda (Kurtis
Productions, 1999). Taylor (1959) also discusses the killing of a drunk by a
leopard in Mozambique. In a similar vein, George Rushby, quoted in the
Tanganyika Game Department, Annual Report, (1951-1952) describes the following:
‘In Ubena, a kind of bamboo is cultivated for
the wine it produces during the rainy season. This wine becomes very potent
when kept for a few days and during the tapping season, which has just
commenced, there is much drunkenness in this district. When I was hunting the
Ubena man eater, I frequently travelled along the main and side roads in my
truck at night hoping to meet the lions. During the wine season, I came upon
men late at night staggering or lying alongside the road in various stages of
intoxication and these were aware that man-eaters were operating in that
particular area at that time. It is possible that man-eating started in that
area through some hungry lion coming across a dead drunk man at night time and
could not resist an easy meal’.
Perhaps lions interpret the laboured movements
of cargo-laden bicyclists as ‘abnormal behavior’. In a number of recent
man-eating incidents in the vicinity of Queen Elizabeth National Park
(1991-1992), several victims were taken from bicycles by an aged male lion with
worn teeth and blunt claws (L. Sieffert, pers. comm.). The recent video,
Maneaters (Kurtis
Productions, 1999), depicts these incidents and shows lions lurking in tall
grass on opposite sides of a road frequented by bicyclists in the area. Taylor
(1959) describes an identical situation in Mozambique in which a pair of lions
on opposite sides of the path killed a bicyclist.
Regarding the Tsavo situation however, there is
no evidence of ‘abnormal’ behaviour on the part of the workers at Tsavo and no
documented selection of Tsavo victims according to their activities. Aberrant
behaviour cannot be documented as a contributing factor to the events at Tsavo
in 1898-1899.
VI. Vegetation and Habitat factors
Dense cover facilitates attacks by ambush
predators (Mitchel, et al. 1965; Schaller, 1972; Prinns, 1996; Funston, et al.
2001). Even in the open Serengeti ecosystem, Schaller documents that 75% of
lion kills occur near cover. Favoured areas include thickets, tall grasslands,
and woodlands. Despite relatively small areas encompassing these habitats,
40-41% of all kills occurred near rivers where dense vegetation and broken
terrain facilitate ambush. Lion predation on buffalo was especially focused on
river edges.
Tigers are also ambush predators and show
parallels with lions. Attacks on people by tigers in the Dudwa National Park,
India, were facilitated by an invasion of villagers into the reserve in order
to cut grass, collect firewood, and especially, by the tall sugar cane around
the periphery (Singh, 1984). The sugar cane fields, analogous to normal tiger
habitat, provide ideal cover for females, and an ideal setting to ambush
people. Such attacks might also reflect vigorous female defense of their young,
which may be nurtured and hidden in the grass thickets. In recent years, more
than 100 people were killed by tigers in Dudwa (Wolfe and Sleeper, 1995).
Cover can also be an important factor when
lions stalk people. Taylor asserts (1959) that a lion will never attack a
person across a wide opening. In the Lindi District of Tanzania, large game is
not plentiful and wild pig and warthog are the favoured prey of lions for much
of the year. However, during the wet season, ‘matette’ grasses grow high and
the rustling of these grasses alert warthog to the approach of lions. With
increasing matette grass height, the ability of lion to capture pigs declines.
During this time of year, man-eating incidents multiply (Harvey, 1932; Ionides,
1938; Taylor, 1959). Taylor (1959) discusses the tendency of man-eaters (both
lions and leopards) to seek cover in this 'matette' grass. Tanzanian Game
ranger Mahenge reports:
‘It is thought that the arrival of the
man-eater around Ifakara during the rains may be due to the fact that the lion
cannot move about in the long grass without their natural prey seeing the tips
of the grass moving and consequently have to resort to feeding on humans who
use the same paths day after day. It may also be that some of the lions get cut
off by the floods which surround the Ifakara area and do not have game to feed
on’ (Mahenge, 1951-52).
Hunting guide and safari operator Richard
Bonham independently reports (personal communication) that these same phenomena
are operational today in south easterrn Tanzania. Alternatively, the withering
away and/or burning of grasses in areas with a different prey base, may lead to
an increase in the frequency of lion attacks on livestock and people. Neumann
(1898) writes that in South Africa, after the grass has been burned during the
dry winter season, lions are unable to approach their normal open country prey.
Instead, they turn to livestock, particularly native cattle. For Kafue National
Park (Zambia), Mitchel et al. (1965) claim that warthog are more easily killed
in the wet season when the grass is tall. Lion preference shifts to buffalo
during the dry season when they are concentrated along rivers. At this time,
the tall grass has been burned by brush fires and grazing species have been
forced onto the plains. It would be interesting to explore these contrasting
seasonal predatory activities and trace the grass and pig species involved.
A belt of dry thorn bush thicket persists along
the east African coast from Lamu to the Zambezi. This habitat never contains a
high density of prey and is prone to man-eating outbreaks (Taylor, 1959). Lions
in this thorn bush belt typically occur in small groups of two and three (not
including cubs) and the mature males are often maneless. In the late 1890's,
due to the demise of elephants from the ivory trade, the predominant vegetation
in the region of Tsavo East National Park was this so-called ‘nyika'. This
thicket facilitates the ambush of Tsavo lions. Patterson and his crew were
often able to hear the screams of lion victims and even able to hear the
crunching of human bones, but could not see the victims. Patterson describes
these thickets as too dense to pursue the man-eaters. His photograph of the
railway line in 1898 (FIGURE 8) illustrates the dense nature of the thickets. Such an environment
encourages man-eating behaviour. Twenty years later, Percival (1928) still
referred to this thick cover and encroaching bush as facilitating fatal attacks
by lions on railroad sentries during World War I. The modern Tsavo environment
has since changed remarkably. Increases in elephant numbers through the 1960’s
resulted in the removal of much of the brushy vegetation, opening up large
tracts of open plains and changing the components of the mammalian biomass.
Tsavo has became one of the largest elephant and buffalo sanctuaries in East
Africa (Leuthold and Leuthold, 1976).
VII. Depletion of 'Typical' Prey and Prey
Depauperate Regions
A frequently cited cause of man-eating
behaviour has been thought to be the absence of, or the depletion of, 'typical'
prey. In particular, disease (e.g. rinderpest), game control efforts, habitat
dynamics, and destruction of natural habitats by humans, can have major
negative impacts on game populations and their dependent predators. The prime
cattle district of Uganda is named Ankole, a name now applied to the breed of
long-horned cattle that are raised there. Beginning in 1924, the most severe
outbreak of man-eating ever recorded in the country occurred in the vicinity of
Sanga. Comprehensive records were not kept but the administrator in charge
states that in less than 4 months, no less than 161 people were killed in 1924
(Temple-Perkins, 1955). Pitman claims (1931) that this incident of man-eating
was due to an outbreak of rinderpest a few years earlier. In an effort to wipe
out this disease, it was decided to destroy the wild ungulates in the region.
Between the removal of their natural prey and the effects of the disease, lions
were left without any prey base. The effects were disastrous. Despite bearing
no apparent injuries, the Ankole District lions turned their attention to
cattle, and thence to man. These circumstances were compounded by the fact that
a major thoroughfare from Rwanda to Uganda ran through the district. Rwandese
immigrants and labourers, traversing this route on foot, and camping in the
open at night without fires, were taken without a formal record of their
disappearance (Temple-Perkins, 1955). Attacks were so bold that entire villages
were abandoned. One of the more famous Sanga man-eaters killed 84 people while
another took over 40 (Temple-Perkins, 1955).
Rushby (1965) attributes the origins of the
Njombe incident (discussed earlier) to the establishment of a game-free area in
southern Tanzania designed to prevent the spread of rinderpest to the south. A
150-mile long pole fence was erected along the southern border with Zambia.
Three teams of European game observers and African scouts had orders to shoot
out all game within 5 miles on both sides of the game fence. Concerns for
protecting livestock from the spread of this disease outweighed those for the
protection of local peoples. The numbers of wild, hoofed animals killed,
steadily increased from 245 in 1943 to 1059 in 1946. The area near Njombe is on
a plateau of 4-6000' in elevation and was already known to be poor in game;
most of the potential prey consisted of small antelope and pigs. Results were
devastating; an estimated 1500 people may have been killed by lions over a
15-year period. Similar records come from Ndola, Zambia, where man-eaters would
appear following great epidemics of rinderpest (Radcliffe-Holmes, 1929).
Perfectly healthy and prime aged lions were among the culprits, suggesting that
injuries were not to blame.
In various localities throughout Africa,
buffalo are the favoured prey of lions (Chobe National Park, Botswana; McBride,
1984; Central African Republic, Stockenstroom, 1987; Lake Manyara, Makacha and
Schaller, 1969, Prins and Iason, 1989, Prins, 1996; Kafue National Park,
Zambia, Mitchel, et al. 1965; Kruger National Park, South Africa, Funston et
al., 1998, 2001; Busanga Flats, Zambia, Mloszewski, 1983). Where they are
buffalo specialists, lions associate themselves with particular buffalo herds
(Mloszewski, 1983). In Tsavo East National Park, buffalo are the favoured prey
of male lions (Lansing & Kerbis Peterhans 2000). Here too, lions associate
with specific buffalo herds (personal communication: Russell, 1998; Packer,
2001). Buffalo are especially sensitive to the disease known as rinderpest.
Though it was not diagnosed at the time, rinderpest first arrived on the
continent (Somalia) in 1889. Dr. Brock, a veterinarian and a colleague of
Patterson’s in Tsavo, was studying the disease in the coastal region of Kenya
(Patterson, 1907). The disease ran through Kenya by mid-1890 (Lugard, 1893;
Mettam, 1937) with devastating results:
“The enormous extent of the devastation it has
caused in Africa can hardly be exaggerated. Most of the tribes possessed vast
herds of thousands on thousands of cattle, and of these, in some localities,
hardly one is left; in others, the deaths have been limited to perhaps 90%’
….Never before....have the cattle died in such vast numbers; never before has
the wild game suffered. Nearly all the buffalo and eland are gone. The giraffe
have suffered, and many of the small antelope...The pig (wart-hog) seem to have
nearly all died....It is noticeable that the animals nearest akin to the cattle
have died-viz., the buffalo and the most bovine of the antelopes, the eland’
(Lugard 1893, p.525, 527-8).
According to Hill (1976), a second outbreak
occurred in 1897:
“Rinderpest had broken out in the Ulu District
and in Kikuyu land towards the end of 1897, and thousands of cattle died. Many
wildebeest, buffalo, hartebeests, buffalo and eland also died, apparently from
the disease, but the loss of game was not so severe as during the rinderpest
epidemic of 1890-1" (p.176).
Portal (1894) states that buffalo declined from
hundreds of thousands throughout German and British East Africa to virtual
extinction between 1890 and 1891. While reflecting on his travelling and
hunting in the Tsavo region between 1893 and 1897, Neumann (1898) stated that
game of any type was exceedingly scarce and that buffalo and cattle, in
particular, were almost extinct from the great cattle plague of some years
earlier. By 1903, cattle had still not recovered to the point where export
could begin (Anonymous, 1947). There can be little doubt that favoured lion
prey, particularly the bovines (cattle, buffalo) were still at low densities in
Tsavo in the late 1890's (see also Wilson, 1913). As a first time visitor to
East Africa in 1897, Patterson had no baseline for comparison. In his journal
of 1898-1899, Patterson does not once refer to cattle or buffalo, while he
refers to other domestic and game animals (not including lions) on 115
occasions (TABLE 1).
Even if water-dependent buffalo were to have been available to lions in Tsavo
in 1897-98, they would have been found at Kanderi Swamp, as well as other
riverine and lacustrine settings. However, Patterson (1907) also describes the
draining of the swamps around Voi in order to control the spread of malaria.
When Patterson returned to Tsavo in 1907 he was surprised that the cattle
population had almost completely recovered (Patterson, 1909). This confirms
that cattle must have been almost locally exterminated by disease during his
earlier work in Tsavo.
During World War I, Percival describes (1928)
the dependence of his troops on wild game, estimating that over 40,000 wild
ungulates were necessary to feed the troops based in Tsavo during the 2 years
of warfare. This dependency on wild game must have further stressed the Tsavo
carnivores. Shortly after the troops departed from Tsavo, they moved into
Masailand where they continued their depredation of wild game. The Masai
promptly complained of the highest level ever of cattle-killing by lions
(Ibid.).
VIII. Maimed Pantherids
The classically
recognized circumstances for man-eating revolve around malnourished, aged, or
wounded animals. The earliest discussant of aged lions attacking people appears
to be Polybius (cited by Pliny; Bostock and Riley, 1855), who states that in
their old age, lions become man-eaters,
attacking people in their
towns because they no longer have the strength to pursue wild prey. Pliny
further adds that such animals are aged as their teeth can be completely worn
down.
Corbett provided detailed
observations of such ailments during his years as the primary dispatcher of
man-eating tigers and leopards on the Indian sub-continent during the first
half of the 20th century (1944, 1948, 1954). Such animals are unable
to pursue, or hold and suffocate, their normal prey. Damaged limbs, teeth, and
embedded porcupine quills appear to be the most common afflictions of the
tigers, lions and leopards that become man-eaters. Of eight man-eating tigers
and two leopards extensively discussed by Corbett (1944, 1948, 1954), severe
physical anomalies were common to all: five of the individuals had at least one
broken canine, three displayed severe porcupine quill damage, and one had an
infected shotgun wound. One of the individuals with a broken canine was
extremely old with worn-out and frayed claws. Three of the five female tigers
had accompanying cubs.
In Africa,
Stevenson-Hamilton (1954) states that embedded porcupine quills may be the
biggest threat to lions after humans. He describes one feeble attack on a pack
horse by an emaciated lion in his prime years that had its foot pads infested
with porcupine quills. Kingsley-Heath (1965) and Piggott shot a large, mature,
emaciated lion in Darajani (adjacent to Tsavo) that had just killed a human. It
was photographed (FIGURE 9) with a
porcupine quill embedded in its nostril. Its perfect canines however, suggested
that he was of prime age. No less than three of the eight man-eating tigers
discussed in detail by Corbet (1944, 1948, 1954) were victims of porcupine
quills.
Taylor (1959) describes
an incident when a local commissioner in Mozambique decided to trap a lion for
close-up photographs. The lion, trapped by its front paw, ultimately tore off
its own foot in order to escape. Now debilitated, it became a notorious
man-eater, even climbing stairways in attempts to get at its victims. One of
the recent man-eaters in the vicinity of Queen Elizabeth National Park (Uganda)
had been caught in a poacher’s snare during the mid-1990s. The damaged
condition of its forelimb is recorded in its skeleton that is still preserved
near the Mweya Lodge (Gnoske, personal observation). In July 1991, one of us
(JCKP) was a first hand observer of an analogous situation in the Ituri Forest
(Democratic Republic of Congo). There was panic in the town of Epulu as a
prime-age leopard suddenly began killing domestic dogs in town. It was
imperative to capture this leopard before people became victims. The leopard
was captured and destroyed. The classic symbol of this doomed predator was the
loss of its left-front paw to a human
snare. The animal had
little recourse but to then turn to the town that had unsuspectingly maimed him
in the first place.
However, man-eating is not
a guaranteed outcome of the serious trauma affecting the predatory behavior of
large Pantherids, especially the lion. The Pipal Pani Tiger (Corbett, 1944) was
gored and seriously wounded by a buffalo. It was later shot in the shoulder by
a local hunter leaving him mildly lame. At this time it became a regular cattle
killer. When it was finally killed, it was never known to have attacked humans.
The Hardwar Tiger, whose body was said to be in ‘perfect’ condition" and
measured 9'10" between the pegs, was a notorious cattle killer but never
attacked humans. After being killed, the skull was prepared and the right
maxillary canine was found growing in an oblique direction through the palate
into the middle of the mouth (Colyer, 1951). The author concludes that the
tiger must have suffered a serious injury to the deciduous canine in its youth.
Both the Field Museum and
the United States National Museum have lion specimens with dental problems
comparable to, or exceeding those of the primary Tsavo culprit. None were
documented as having been man-eaters. Fifty percent of the length of the right
upper canine of FMNH 75608 (FIGURE 10)
has been broken away, while the left upper canine has been broken to the gum
line. Two additional lion specimens from areas adjacent to Tsavo housed at the
United States National Museum (Moshi, Tanzania; Taveta, Kenya) suffered
traumatic dental injuries leaving the pulp cavities exposed, but were not
documented to have been man-eaters. Finally, the 12 injured lions we have
observed in Tsavo over the past few years were not known to have attacked
people or their livestock.
The inverse situation is also true. Healthy
prime age and sub-adult lions often engage in attacks on domestic stock and
humans. The Field Museum recently acquired (1998) the skull of the
"Man-eater of Mfuwe" (FMNH 163109), one of the largest man-eaters on
record and with near-perfect teeth in the prime of his life (ca.5-6 years old
per criteria in Smuts, et al. 1978). Upon the arrival of this specimen at Field
Museum, we immediately noted a bony swelling with abscess on its mandible.
Although one might be tempted to attribute its man-eating behavior to this
injury, the occlusion of the teeth was not effected and the canines are in
perfect condition. It is doubtful that this trauma contributed to its man-eating
habits. A number of prime-condition Kenyan lions had serious reputations as
cattle-killers but never became man-eaters. These include 6 individuals with
near-perfect teeth and skeletons housed at the National Museums of Kenya
(Department of Osteology): 2 individuals from Laikipia, 2 from the Tana
watershed near Kangudo, and two from the vicinity of the Aberdares and Mt.
Kenya. Two individuals were sub-adult, 2 were prime and one was post-prime.
Interestingly, the post-prime lion is the heaviest free ranging lion ever
recorded at 272 kilograms. Despite their apparent healthy skulls and skeletons,
these individuals did become stock raiders, perhaps due to ever-impinging
development impacting their prey base.
Lion preference for
buffalo, an often dangerous prey species, can lead to debilitating injuries,
even death (Roosevelt and Heller, 1914; Mangani, 1962; Schaller, 1972;
Sinclair, 1977; Mloszewski, 1983). Today, in Tsavo East National Park, buffalo
are the primary prey for lions (Lansing and Kerbis Peterhans, 2000; Gnoske,
personal observation, 2001), especially along the Voi River drainage and other
permanent waterways where buffalo congregate. In April of 1997, one maneless
male lion was dispatched by KWS after suffering a broken foreleg during an encounter
with a buffalo (Muhanga, personal communication). Our own observations in Tsavo
document possible scenarios of impairment. Between 1998 and 2001, both of us
observed mature lions with potentially debilitating injuries. These animals
were located in fully protected areas adjacent to Voi, just within the southern
boundary of Tsavo East N.P. In 1998, after killing a female buffalo, one lion
was limping to the extent that KWS rangers considered its removal. A second
lion suffered an injury to its right eye as well as to its forelimb. A third
individual was filmed with blood coming from both ears. Two photographs by H.
Schuetz taken in 2001 depict adult male lions, suffering from severe dental
trauma (FIGURES 11,12), on fresh kills
of elephant and buffalo respectively. In total, we documented 12 injured lions,
10 of which were males. Four males and one female had moderately to severely
broken canine teeth. Six males exhibited head trauma, one male suffered from
back trauma, and three males and one female exhibited injured forelimbs.
Several of these individuals displayed a combination of injuries. Despite these
injuries, there is no evidence that these animals left the park to become
man-eaters or cattle rustlers; they continued to pursue wild prey.
In order to investigate
possible correlations between injuries, sex, and age with the appearance of
problem lions in Tsavo today, we reviewed 29 'problem lion’ skulls from the
‘trophy room’ of Tsavo East National Park Headquarters at Voi. We follow Smuts
et al. (1978) in recognizing 3-4 year olds (e.g. ‘sub-adult’; Schaller, 1972)
as those lions with a completely erupted and virtually unworn dentition,
whereas 5-7 year olds (prime-age) display dental yellowing and visible canine
wear as well as wear on the tips of the highest cusps of P3 and P4.
Our analysis (TABLE 5) shows that 10
were from healthy prime-age individuals and 14 from sub-adults with only two of
these 24 individuals exhibiting slightly damaged carnassial teeth. Of the
remaining 5 aged or past-prime individuals, one displayed an upper left
carnassial broken in life, two had moderately worn and broken canines, one was
past prime with good canines, and one could be considered senile with severely
worn canines. In sum, these skulls were derived from generally healthy,
sub-adult and prime aged individuals with intact dentitions. As detailed
earlier, one of the Tsavo man-eaters had a severely broken and mal-occluding
canine that may have contributed to its man-eating habits, while the other
individual had a broken carnassial that would not have affected its predatory
habits.
Table 5
Analysis of 29 Skulls of Problem Lions
Housed at Tsavo East National Park- Problem Animal Control Offices, Voi
Kenya Wildlife Service
Sub-adult (n=14) Prime
Adult (n=10) Post-Prime
(n=5)
11: Male, perfect teeth 5: Male, perfect teeth 2: Male, 2 worn/damaged canines ea.
2: Female, perfect teeth 1:
Male, broken P4 2:
Female (prob), mod. worn teeth
1: Female, damaged P4’s 4:
Female, perfect teeth 1:
Female, blunt canines, broken P4
We also sexed these
animals by measuring their canine teeth. Smuts et al. (1978) documented no
overlap in unworn canine heights of adult male and female lions from Kruger
National Park, South Africa (n=122). Unless labels dictated otherwise, we
assumed a minimum unworn canine height of 4.7mm would represent male lions
(females ranging from 3.7mm to 4.6 mm). Sex determination from canine height
corresponds with our independent sex determinations of lion skulls: males
usually have an upper P4 over 3.6mm (TPG observations). Nineteen of
29 sexable lion skulls (66%) from Tsavo East culling operations are from males.
Of the 19 Tsavo males, 11 had unworn canines (3-4 year olds) while only 6
displayed apical canine wear, defined by Smuts et al. (1967) as a characteristic
of lions in the 5-6 year age class. In sum, problem lions adjacent to Tsavo
East NP, Kenya, are typically male (66%) in the 3-4 year old age category (58%
of the males). Although the data is minimal, it appears that as lions age,
females become potentially as problematic as older males: 7/14 (50%) of our
Tsavo sample in the prime and aged categories are female. However, this may
simply be due to their greater frequency in the population. Based on tooth wear
(Van Valkenberg, 1988), female lions are longer-lived, which may account for
their greater frequency in the ‘older’ category.
It is likely that
debilitating injuries to the limbs are more likely to lead to man-eating than a
severely broken canine. However, we are unable to confirm this due to the lack
of post-cranial skeletons from animal control offices. D. King (unpublished
report) observed a female radio-collared lion in Tsavo, with canines worn to
nubs, killing an adult female eland. Lions do not necessarily kill large
bovines with their canines, but more often by breaking the necks or suffocating
their victims (Selous, 1907, 1908; Gnoske, personal observation). Similar
observations have been made on tigers for several locations in India (reviewed
in Fletcher, 1911). These activities require a vice-like grip rather than an
impaling stab wound. The forelimbs
however, must function
together. One forelimb is used to secure the victim’s head and to control the
horns while the other assists in pushing the animal off its feet, often
breaking its neck.
Discussion
This review of select
man-eating incidents confirms that certain circumstances can lead to the
development of man-eating behavior among large Pantherids, possibly including
the Man-eaters of Tsavo. These circumstances include: inadvertent or purposeful
provisioning, abandoned human corpses, livestock availability, lion
demographics and social traditions, peculiar behavior of the prey, seasonal
factors, dense cover, prey-depleted landscapes, and impairment or injury.
However, we are able to eliminate several scenarios in regard to the Tsavo
situation of 1898. For example, no available data suggest that laborers in
Tsavo behaved in a peculiar manner, thereby subjecting themselves to lion
attack. Although there appears to be a seasonal component (rainy season vs. dry
season) in the timing of attacks on bomas in Tsavo today, we discount any
association with the onset of the rains in the Tsavo of 1898, due to the noted
delay in the onset of the rains that year (Hill, 1976).
Both pastoral peoples and
hunter-gatherers are vulnerable to Pantherid confrontation and attack. Treves
and Naughton-Treves (1999) have shown that individuals employed in
meat-gathering activities can be victims of large carnivore attack while
poaching in parks or while attempting to drive large Pantherids off of their
kills (6 reports resulting in 9 casualties). The Darajani Man-eater killed a
poacher on the outskirts of Tsavo (Kingsley-Heath, 1965). Marks (1976)
describes conflicts endured by the Bisa, of Zambia, who secure a major
component of their meat by scavenging buffalo directly from lions. Similar
cases are documented today in Queen Elizabeth National Park (Sieffert, personal
communication). Pastoral communities, adjacent to protected areas, are also
especially vulnerable. The only documented precise location of an attack on
people in Tsavo in recent times states that the attack occurred in a 'cattle
boma' or corral, to someone defending or providing for their livestock. Corbett
also discusses several instances of tigers attacking pastoralists looking for
their livestock. As discussed earlier, numerous authors have anecdotally
claimed that the transition from domestic stock depredations, to conflict with
humans, to predation on humans, is an oft-repeated scenario. In parts of India
where tigers are protected, programs are underway which provide a source of
wild, hoofed stock in order to satisfy predatory demand.
Our review of problem lion
skulls derived from animals culled by the Kenya Wildlife Service shows an
overall predominance (83%) of subadult (3-4 year olds) and prime-aged (5-7 year
olds) individuals rather than aged individuals (over 7 years old). Among the
subadult and prime-aged lions, 70% are males. Sex ratios, determined from the
skulls of 'problem' animals, are complemented by documented attacks of male
lions on livestock and humans in Tsavo East National Park (54% and 83%
respectively, Table 2). This disproportionate impact by male lions may in fact
be much higher if one were to assume heavily female-skewed sex ratios among
adult lions as have been described for most lion populations. This male cohort,
despite absolutely smaller numbers, appears to be the predominant ‘problem’
demographic component of the Tsavo lion population. The bias in male
‘trouble-makers’ among the Pantherids is most pronounced among leopards where
143 of 152 (94%) known man-eaters are males (Turnbull-Kemp, 1967).
This demographic cohort
mirrors the age at which ‘subadult’ males leave or are forced out of their
natal groups (prides), described by Schaller (1972) as being between 2 1/2 and
3 1/2 years of age. Saberwal (1994) comes to similar conclusions for Indian
lions: " subadult lions were represented disproportionately in conflicts
with humans" (p.504) and "subadult lions involved in attacks are
likely those displaced from or dispersing from their natal territories"
(p. 506). The second largest sex and age cohort, is represented by prime-aged
male lions (Table 3). This is the category into which the Tsavo man-eaters
belong.
Corbett's observations
(1944, 1948, 1954) document the frequently damaged condition of man-eating
tigers and leopards in India. Under certain circumstances, these observations
have been applied to African lions, perhaps including the man-eaters of Tsavo.
However, the theory championed by Corbett (1944, 1948, 1954) that aged or
impaired Pantherids are the primary culprits in human/carnivore conflict needs
to be modified. Our data suggest that only 15% of disruptive lions (including
maneaters) are so impaired. No data are available to indicate whether 15% is a
high incidence of impairment among lion populations. Schaller (1972) suggests
that only 10% of lions in the Serengeti reach old age. We believe that
impairments to the appendages (broken or snared limbs, porcupine quills) are
much more likely to lead to attacks on livestock or people than broken teeth.
Severely broken canines and extensive canine wear may have an effect on a
lion’s ability to grip ungulate prey, but trauma to incisors, premolars and
molars have little effect in subduing prey. It is clear that severe canine
impairment needs to be distinguished from other dental traumas when discussing
impacts on predatory behavior.
Even so, lion with
severely broken canines have not been shown to be particularly troublesome. The
cohort of sub-adult and prime aged males dispatched near Tsavo East N.P., are
generally healthy individuals, at least from the cranio-dental perspective
(Table 3). These individuals, collected by Problem Animal Control (PAC), were
responsible for having attacked livestock and threatening people. They are not
aged or infirm. In fact, the frequency of broken canines (4/29=14%, Table 3)
and broken carnassials (3/29=10%, Table 3) in the PAC skulls are equal to or
less than their frequency in lion skulls housed in museum collections. For
example, Van Valkenburgh (1988) documented 26 broken canines (21.5%) and 13
broken carnassials (10.7%) in a review of 121 lion skulls in museums.
Lions dispatched by the
PAC unit contrast the group of living lions, documented by us within Tsavo East
and outside of the areas patrolled by PAC, which do display significant
cranio-dental trauma. It would appear that lions with serious cranio-dental
trauma are not typically becoming cattle-killers and threatening people. It is
also possible however, that this group knows that if they remain within the
official boundaries of the park, they will not be eliminated. Although there
are a number of authentic and well-documented cases of impaired lions becoming
man-eaters (Pliny, cited in Rawlinson, 1992; Selous, 1908; Taylor, 1959;
Kingsley-Heath, 1965), they account for a low proportion of the cattle-killing
and man-eating incidents recorded. Further, many of these examples illustrate
problems unrelated to cranio-dental trauma (e.g. porcupine quills, starvation,
old age, impaired limbs, etc.).
The most comprehensive
review of the physical condition of man-eating Pantherids (lions, tigers, and
leopards) has been produced by Turnbull-Kemp (1967, Table 2), who states that
“a high percentage of the man-eaters among the three species considered here
are mature individuals bearing no obvious injury and certainly unhampered by
any wound or the like” (p.133). We have partially reproduced his Table here (TABLE
6). Only 22% of the lions and 11% of the
leopards displayed an injury in Turnbull-Kemp’s data set (Ibid.).
Unfortunately, there is no breakdown on the nature of the injuries. If aged
individuals are added, the figures rise only to 24% and 12% respectively. The
data from Treves and Naughton-Treves (1999) provides further support for these
low numbers: only 14% of the lion attacks (n=275) and 15% of the leopard
attacks (n=114) were attributed to injured animals. Approximately 13% (3/29) of
the skulls from the KWS animal control office at Voi, showed evidence of
serious dental trauma or advanced age. Finally, Thessiger (personal
communication) states that the 30 problem lions he shot in Northern Darfur
(Sudan, 1935-1937) were generally healthy and of prime-age. Although certain
injuries are overlooked (e.g. the Tsavo maneaters), the vast majority of
attacks appear to be conducted by healthy animals. These unique data sets
(TABLES 5-6; Turnbull-Kemp, 1967; Naughton-Treves, 1999) stand in contrast to
the often-quoted statement that man-eating lions are generally impaired
animals. In fact, the greatest proponent of this belief was Corbet who
popularized this notion based on a spectacular group of (but limited number of)
man-eating tigers that may be more prone to develop this habit due to their
solitary habits. Post-cranial injury may play an occasional role and animals
with impaired limbs need to be rehabilitated or eliminated, a policy currently
practiced by the Kenya Wildlife Service. The great majority of man-eating and
cattle killing lions, are sub-adult or prime-aged animals in good health.
It is also important to
point out that this cohort of sub-adult and prime aged males
(TABLE 5) are healthy
individuals, at least from the cranio-dental perspective. These individuals
were responsible for having attacked livestock and threatening people. They are
not aged or infirmed. They contrast the group of male lions, documented by us
within Tsavo East
TABLE
6
Age and Condition Data on 241 known Man-eaters at
Death
(from Turnbull-Kemp, 1967)
Lion Tiger Leopard
# in sample 89 74 78
AGED (POST-PRIME) (18%) (35%)* (12%)
uninjured 12
(14%) 9
(12%) 7
(9%)
injured 2
(2%) 15
(20%) 1
(1+%)
aged dentition 2
(2%) 2
( 3%) 1
(1+%)
MATURE (PRIME) (33%) (47%) (85%)
uninjured 24
(27%) 28
(38%) 62
(79.5%)
injured 5
( 6%) 7
( 9.5%) 4
( 5+%)
IMMATURE (SUBADULT) (50%) (18%) (4%)
uinjured 32
(36%) 12
(16+%) 0
injured 12
(14%) 1
( 1+%) 3
(4%)
*figure corrected from Turnbull-Kemp’s
figure of 55.1%
figures rounded to nearest
%
and adjacent to the areas
patrolled by KWS Problem Animal Control Units, that do display significant
cranio-dental trauma. It would appear that lions with serious cranio-dental
trauma are not often becoming cattle-killers and threatening people. It is also
possible that this group knows that if they remain within the official
boundaries of the park, they will not be destroyed. Although there have been a
number of authentic and well-documented published cases of impaired lions
becoming man-eaters, they appear to account for a low proportion of the
cattle-killing and man-eating incidents recorded.
With regard to age, most (82%) of the
man-eating lions discussed by Turnbull-Kemp were either ‘mature’ (33%) or
‘immature’ (50%), whereas only 18% were classified as ‘aged’. Tigers and
leopards on the other hand, display unique and distinct patterns. The great
majority of the man-eating leopards (85%) were ‘mature’ animals. Compared with
lions, twice as many of the man-eating tigers were listed as ‘aged’ (35% vs.
18%). As suggested by Turnbull-Kemp (1967), the solitary tiger might be more
prone to man-eating when its physical condition (old age or injury) prevents it
from subduing ungulate prey. When its condition deteriorates, the more social
lion might be more likely to take advantage of prey brought down by affiliates,
allies or ‘pride-mates’. Conversely, lions are 2 1/2 to 3 times more likely to
become man-eaters when ‘immature’ (50% in lions vs. 18% in tigers), an age
category he defines as being approximately 3 years of age (equivalent to
Schaller’s <1972> ‘subadult’ category). This age group is the same
problem cohort reflected in our data set from Tsavo East National Park.
Leopards differ from both lion and tiger as they are basically mature and
uninjured when they take to man-eating (85% mature, 80% mature and uninjured). Leopards are the smallest of these
three large cat species and might find adult male humans a more formidable
adversary. This may be why leopards are twice as likely to attack human women
and children than lions (67% vs. 33%, n=138;Treves and Naughton-Treves, 1999)
and also why larger male leopards are more likely to become man-eaters than the
smaller females.
The Tsavo man-eaters can be classified in
Turnbull-Kemp’s (1967) ‘mature’ category. Although this is not consistent with
Turnbull Kemp’s highest risk group (‘immature’ at 50%), fully 1/3 of the
man-eaters Turnbull-Kemp documents do fall into this age cohort. It is also
possible however, that the Tsavo lions began their man-eating career two years
earlier as documented by Ansorge (1899). The fact that they were male, further
places them into the ‘trouble-maker’ category. However, given that the two
acted together, there was no reason why the severely damaged canine and
remodeled skull of one of them, turned both of them into man-eaters. Their
social bonds, immense size, apparently healthy limbs, and mature age would
allow them to tackle ‘normal’ prey. Indeed, this has been confirmed with the
species profile represented by hairs extracted from their broken canines.
Adult male lions in Tsavo (Lansing and
Kerbis Peterhans, 2000) and elsewhere preferentially target buffalo. When
buffalo populations crash, buffalo-dependent lions can switch to alternative
food supplies, including humans. Africa's most notorious episodes of man-eating
were associated with the depletion of buffalo, or other prey animals. These
include the Tsavo incident, the Ankole outbreak in Uganda (rinderpest outbreak
killing buffalo and resultant game eradication) and the Njombe incident in southern
Tanzania (associated with the development of a game free corridor). Domestic
cattle on the outskirts of protected areas can act as buffalo surrogates for
these lions; lions apparently see little difference between the two. Injuries
from encounters with buffalo can also impair lion ability to secure normal wild
prey and push them into attacks on domestic stock and perhaps humans.
'Provisioning' of carnivores can lead to
instances of man-eating. Provisioning of lions in the Gir Forest (India) was
abolished following a correlation between lion provisioning and man-eating
outbreaks (Saberwal et al., 1994). Recent attacks on tourists adjacent to, and
within national parks, by leopards and hyenas have led to a policy of
non-provisioning for these large carnivores in parts of Africa, including parts
of Kenya. Similarly, traditional practices of abandoning the dead in the bush
can potentially stimulate predators to seize live individuals. It is especially
ironic that cattle-loving Masai might attract lions to their bomas by
abandoning their dead in the bush. Deceased humans were available in Tsavo
(1898/99) in the form of local ‘burial’ practices, hundreds of Wakamba dead
from starvation, hundreds of railway workers dead from disease, and a recurring
opportunity of abandoned or injured porters and slaves. Non-indigenous porters
from outside localities often died from dysentery and a suite of pathogens to
which they were not accustomed.
In regards to ' provisioning', it is
important to note that many of the man-eaters we discuss here came to animal
baits. These were not 'dedicated' or 'obligatory' man-eaters. For instance, the
Mfuwe man-eater was attracted with hippopotamus parts (Hippopotamus
amphibious), Corbet's man-eaters were
regularly taken with cattle and goats, and the Tsavo man-eaters were lured to
their deaths with a donkey and three tethered goats respectively. The Tsavo
pair continued to pursue wild prey immediately following the severe trauma
experienced by FMNH 23970.
Civil strife also plays a
role in providing Pantherids with human corpses as we have documented for Viet
Nam and World War II. In order to avoid detection, rebels will travel off-road,
through uninhabited, predator-infested landscapes. Areas prone to man-eating
today include Queen Elizabeth National Park (Uganda) and Kruger National Park
(South Africa). Queen Elizabeth borders the Democratic Republic of Congo while
Kruger borders Mozambique; both of these protected areas have functioned as
highways for 'rebel' infiltrators who come into contact with large predators,
sometimes with fatal results. Four rebels were recently killed by lions in
western Uganda (New Vision, 13 May 1997) while infiltrating from the Democratic
Republic of Congo. In order to keep a low profile, rebels also avoid using
fires at night, further encouraging predators.
We have documented over 30 years of nearly
continuous recorded predation of humans along the caravan and railway route in
the Tsavo region, suggesting that man-eating was a regular part of lion life in
this area. An entourage of predators was attracted to caravan routes
through which people, porters, livestock, and their transport animals passed.
Caravan thoroughfares were oases in the thorny desert and were likely adopted
as human hunting grounds by Tsavo lions for generations. The camps of the
railway crew were extensive as they supported 2000-3000 individuals (Patterson,
1907) and provided a plentiful source of food debris and dead transport
animals.
Several circumstances contributed to highlight
the area of the Tsavo bridge as an attractive feeding ground. This is the point
where the Tsavo River intercepts the traditional caravan route. Patterson notes
that this was the unofficial point to ford the river for generations of
caravans. This point is less than four kilometres from the confluence of the
Tsavo and Athi Rivers. The Tsavo and Athi Rivers are the only year-round source
of water in the vicinity. The Tsavo drains the glaciers of nearby Mt.
Kilimanjaro. To anyone who has ambled through the equatorial heat of Tsavo, the
Tsavo River is a welcome oasis in the heart of a sub-desert, thorn-scrub
thicket. Even today, green-topped trees are restricted to the river’s edge.
Patterson himself joyfully described a walking excursion from his camp near the
bridge: "walking stealthily along in the delightful shade of the
overhanging palms" (1907, p.). The area of the bridge was a rest and
refuelling point for land caravans, all to eager too set up camp in the shade,
adjacent to fresh water…..just in time for marauding nocturnal carnivores. This
is why we believe that this area was prone to man-eating throughout the days of
the caravan.
For a long-lived species with a long period of
infant/ juvenile dependency, any regularly practised predatory behaviour can
become a cultural tradition, passed down from one generation to the next
(Taylor, 1959; Rushby, 1965). To call such behaviour 'aberrant' may be
acceptable from an anthropogenic perspective but is 'normal' behaviour for the
relevant predator. Over thirty years of documented man-eating at the turn of
the 20th century in Tsavo may represent six generations of lions,
all of whom may have been part of the same social tradition. The 15-year reign
of the infamous "man-eaters of Njombe", the Sanga man-eaters, and the
modern behavior of the swamp tigers of the Sundarbarns also fall into this
category. At the turn of the 20th century, the well-travelled
caravan route was replaced with a parallel rail line and ultimately an
automobile route. These actions denied area carnivores access to vulnerable
campers. In their frustration with the 'iron horse' in the first years of the
railway, the big cats were even documented removing humans from railway
carriages (Patterson, 1907; Anonymous, 1961). Their human depredations focused
on the railway stations of Makindu, Kima, Voi and Simba (Miller, 1971).
The mega-fauna of Tsavo
has been historically dynamic. In three major ways, the faunal composition
during Patterson’s time contributed to the propensity of Tsavo lions to select
people as prey. The Tsavo nyika was quite limited in its capacity to maintain a
high prey biomass and was dominated by large rhino and small antelope (e.g.
dik-dik) well above and below the size preferred by lion. Taylor (1959) notes
the propensity of lions to become man-eaters in such habitats. According to
Patterson’s journal entries (1898-1899) medium-sized, gregarious ungulates were
either absent (wildebeest) or sparse (zebra). Humans became even more
vulnerable to Tsavo lions with the dramatic reduction of buffalo, their favored
prey under normal conditions. The biomass of buffalo in Tsavo is normally the
highest among potential lion prey species (Leuthold and Leuthold, 1976).
However, two waves of rinderpest had decimated the buffalo herds and the
draining of the Kanderi Swamp prevented a prompt comeback. Ivory hunters had
already eliminated the elephant population. In his journal (1898-1899), despite
185 references to the fauna of Tsavo, Patterson never once mentions buffalo,
cattle or elephants. The absence of elephants had yet another effect on Tsavo
lions: the thorny shrub vegetation became overgrown providing easy ambush and
exit for predatory lions.
We cannot claim that any
single cause will guarantee that a lion will turn into a ‘man-eater’, but it is
clear that a variety of causes will increase the likelihood. In regard to the
Tsavo situation, we have discounted several factors often proposed to account
for man-eating outbreaks. These include peculiar behgavior of the prey,
seasonal factors and broken teeth. However, several important pre-conditions
for the development of man-eating were in place in Tsavo during the 1890’s.
These include a prey-depleted landscape, long standing behavioral traditions,
inadvertent provisioning of humans and their livestock, and habitat factors. Although the human toll at Tsavo was
claimed to exceed 100 individuals, it seems the total could have been higher.
Given the circumstances at Tsavo in the 1890's, instead of asking how so many
humans could have been dispatched, we wonder why there weren't more.
Acknowledgements
An interdisciplinary study such as this owes
thanks to a variety of individuals from diverse backgrounds. We are especially
indebted to the following individuals and institutions for their support: The
Office of the President, Kenya (Permit No. MOES&T 13/001/30C 57), Eli Lilly
Foundation; Barbara Brown Fund (Field Museum of Natural History); Bill Kurtis
and Seamus Gallagher (Kurtis Productions); Stephanie Stephens and Nina Cummings
(Library and Photo Archives, Field Museum); Samuel Andanje, Naftali Kio, James
Isiche, John Muhanga, Samuel Kasiki, Richard Bagine (Kenya Wildlife Service);
Ogeto Mwebi (National Museums of Kenya); Robert Kityo and Ludwig Sieffert
(Makerere University), Craig Packer (University of Minnesota), Troy Nowak
(Illinois State Police), Richard Bonham, Gordon Boy, Peter Buol, Phil Caputo,
Dennis King, Alexander Maitland, Lisa Ostovits, Alan Patterson, John Perrott,
Anthony Russell, Wilfred Thessiger, and Vincent Wan. For the original illustrations
(Figures 3,4 & 10) and photographs (Figures 11,12), we acknowledge Kathryn
Keith and Harald Schuetz respectively. For comments on our manuscript we
ackowledge Tom Butynski, Luke Hunter, Dave Willard and Jose Tello.
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Headings to Figures
Figure 1 The second Tsavo man-eater
slain (FMNH 23969).
Figure 2 The first Tsavo man-eater
slain (FMNH 23970).
Figure 3 Lateral
view of FMNH 23970 depicting forward projection of right upper canine. Compare
with vertical left upper canine visible in rear (courtesy of K. Keith).
Figure 4 Anterior
view of FMNH 23970 depicting broken cavity of lower right canine. Note
asymmetry of upper incisors and canines due to broken and missing mandibular
teeth (courtesy of K. Keith).
Figure 5 Caravan
crossing the Tsavo River (Patterson, 1907).
Figure 6 Lion hovering around camp
(Selous, 1908).
Figure 7 Hyenas circling corpse
(Selous, 1908).
Figure 8 The
dense Tsavo bush (nyika) along the railway (Patterson, 1907).
Figure 9 The
Darjani Man-eater with embedded porcupine quill in nostril (courtesy of J.
Perrot).
Figure
10 Pearson lion, FMNH 75608 (courtesy
of K. Keith).
Figure 11 Tsavo lion with
dental trauma, on elephant carcass (courtesy of H. Schuetz, 2001).
Figure 12 Tsavo lion with
dental trauma, on buffalo carcass (courtesy of H. Schuetz, 2001).
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