Diet Choise of Large Carnivores in Nothern Cameroon

Diet choice of large carnivores in northern Cameroon

Thomas Breuer*Center for Nature Conservation, University of Gottingen, Von-Siebold-Strasse 2, 37075 Gottingen, Germany

Abstract

The diet of African wild dogs, lions and spotted hyenas was

studied in the woodland savannah of Faro National Park

and surrounding hunting zones in northern Cameroon

during the dry season. Faecal analysis revealed that the

diet of the three large carnivores overlaps considerably.

Frequencies of prey remains of large and medium-sized

antelopes were highest, with Buffon’s kob being by far the

most common prey species. Carnivores seem to consume

most abundant prey species. No prey item of domestic

livestock was found in the faeces. Faecal analysis has

proved to be a useful method to reveal diet choice for large

carnivores in northern Cameroon. Detailed information on

the distribution and density of large carnivores and their

prey is needed to give a better picture of the status of

carnivores in northern Cameroon and to help resolve

conflicts with livestock. Further investigations into the

seasonal variation of predation and other factors struc-

turing the large carnivore community in northern Cam-

eroon are recommended.

Key words: Cameroon, carnivores, diet choice, faecal ana-

lysis

Resume

La diete des chiens sauvages africains, des lions, et des

hyenes tachetees fut etudie dans la savane boisee du parc

national de Faro et les zones de chasse environnantes au

nord du Cameroun pendant la saison seche. Des analyses de

la matiere fecale demontrerent que la diete de ces trois

grands carnivores se chevauchent considerablement.

Parmi les restes de proie, les antilopes de taille grande

et moyenne figuraient le plus souvent, avec le kob de Buffon

l’espece de proie la plus repandue de loin. Les carnivores

semblent consommer les especes plus abondantes. Aucune

proie originaire du betail domestique n’a etait trouvee dans

la matiere fecale. L’analyse de la matiere fecale s’est mon-

tree une methode tres utile de decouvrir les choix ali-

mentaires des grands carnivores au nord du Cameroun. Des

donnees exhaustives sur la distribution et la densite des

grands carnivores s’averent necessaires afin de donner une

meilleure image de la situation des carnivores au nord du

Cameroun et de trouver des moyens de recoudre des conflits

avec le betail. Plus de recherche sur les fluctuations

saisonnieres de la predation et d’autres elements qui

impactent sur la communaute de grands carnivores au

nord du Cameroun est preconisee.

Introduction

Information on the feeding ecology of large carnivores

contributes substantially to the understanding of their

behavioural ecology (Mills, 1992). Prey availability and

distribution can influence prey selection and hunting

success (Fuller et al., 1992), as well as activity patterns and

spatial distribution (Henschel & Skinner, 1990).

The feeding ecology of large savannah carnivores has

been well studied in east and southern Africa (Kruuk &

Turner, 1967; Pienaar, 1969; Mills & Biggs, 1993), but

only limited information is available for large carnivores in

west and central Africa (Woodroe, Ginsberg & Macdonald,

1997; Mills & Hofer, 1998; Bauer et al., 2001). Further-

more, studies on carnivore food habits in west and central

Africa have concentrated on tropical forest carnivores,

especially leopards (Panthera pardus) (Hoppe-Dominik,

1984; Hart, Katembo & Punga, 1996; Ososky, 1998;

Henschel, 2001; Ray & Sunquist, 2001). Only a few

studies have been conducted in the savannah (Wanzie,

1986; Ruggiero, 1991; Gross, 1997; Di Silvestre, Novelli &

Bogliani, 2000; Korb, 2000).

The present study was initiated as part of an investiga-

tion into the conservation biology of the endangered*Correspondence: Rubenacherstraße 29, 56220 Kaltenengers,

Germany. E-mail: [email protected]

� 2005 African Journal of Ecology, Afr. J. Ecol., 43, 97–106 97

African wild dog (Lycaon pictus Temminck) in Faro National

Park (FNP), northern Cameroon (Breuer, 2001). The aim of

this paper is to provide information on the frequencies of

prey species found in large carnivore faeces and their diet

overlap in the woodland savannah in and around FNP. The

potential of faecal analysis for determining feeding ecology

of carnivores in Central Africa is discussed and recom-

mendations for future research are made.

Material and methods

The study was carried out in the dry season 2000/2001 in

the northern part of FNP and adjoining hunting zones

(between 8�54¢N–8�13¢N and 12�63¢E–13�17¢E) covering

an area over 900 km2 (Breuer, 2001). The dry season was

chosen because of limited visibility because of high grass in

the rainy season. FNP is located in the west of northern

Cameroon and comprises a plateau with mountainous

massifs, limited in the south by the highland of Adamawa

and in the west by the Atlantika-mountains. Several insel-

bergs are situated in the park. The borders of FNP are formed

by naturally occurring rivers, except in the south. Typical

soils in the area are low nutrient and ferruginous tropical

(Brabant & Humbel, 1974).

The study area lies in the Sudano-Guinean zone of

northern Cameroon, dominated by Sudanian savannah

and woodland (White, 1983). The dry season from

November to April is characterized by a hot and dusty

north-east cyclone (Harmattan), and the rainy season

(May to October) is dominated by a south-west monsoon

associated with more humidity (Aubreville, 1950).

Annual rainfall is 1200–1400 mm, with the maximum

falling between August and September (Brabant &

Humbel, 1974). Four different vegetation units are found

in FNP: Isoberlinia spp. woodland, Terminalia spp. and

Burkea africana open savannah and Anogeissus leiocarpus

riparian forest (Donfack et al., 2001). The potential prey

species list in FNP includes Buffon’s kob, Kobus kob

(Erxleben); defassa waterbuck, Kobus ellipsiprymnus

defassa (Ogilby); bushbuck, Tragelaphus scriptus (Pallas);

Giant (Derby’s) Eland, Taurotragus derbianus (Gray); roan

antelope, Hippotragus equinus (Desmarest); hartebeest,

Alcelaphus buselaphus (Pallas), Ourebia ourebi (Zimmer-

mann); red-flanked duiker, Cephalophus rufilatus (Gray);

common duiker, Sylvicapra grimmia (L.); buffalo, Syncerus

caffer (Spartman); warthog, Phacochoerus aethiopicus

(Pallas). Red river hog, Potamochoerus porcus (L.); oribi,

Guinea baboon, Papio papio papio (Desmarest); vervet

monkey, Cercopithecus aethiops (L.); Guereza Colobus,

Colobus guereza (Ruppell); the patas monkey, Erythrocebus

patas (Schreber) and several rodents, hares, small

carnivores and insectivores.

Carnivore faeces were collected in and around FNP,

recording location, date and age, categorized into fresh

(1 day old), recent (around 1–5 days old) and old. Two wild

dog faeces were also collected outside the study area

southeast of Benoue National Park and were included in the

analysis. Only one wild dog faecal sample was analysed

when more than one sample was found at the same spot,

because pack members eat from the same kill (Woodroe

et al., 1997). Spotted hyena (Crocuta crocuta Erxleben) faeces

were collected opportunistically and at latrine sites where

they often defecate to exchange information (Kruuk, 1972;

Mills & Gorman, 1987). Most lion (Panthera leo L.) faeces

were found in dried up riverbeds and in riparian forests and

fewer encountered far from water.

Only those faeces of the same age were collected,

which were at least 5 km apart from each other (Hart

et al., 1996). Carnivore faeces identification was based

on shape, colour, ingested hair (from grooming), diam-

eter, odour, and by the presence of associated field signs,

such as tracks. Faeces were compared with pictures in

field guides. Additionally trackers helped in identification

and ageing of faeces. Faeces of doubtful identity were

excluded from the analysis.

Faecal samples were sun dried, ground in a mortar, and

washed in a sieve (1 mm), using hot water to separate

hairs, bones, hoofs, teeth and other prey components from

other organic material. Separated hairs were washed in

acetone, dehydrated in 100% ethanol and dried on filter

paper (Ramakrishnan, Coss & Pelkey, 1999). Hair was

analysed macroscopically (form, length, colour), using a

stereo microscope and microscopically (scale patterns)

under a light microscope following methods described in

Ramakrishnan et al. (1999).

A reference collection of prey hair was made from hairs

of mammals (poached or killed by carnivores) (Breuer,

2001). Scale patterns were compared with this reference

collection and photographs in the literature (Dreyer, 1966;

Feder, 1975; Perrin & Campbell, 1980; Keogh, 1983; Buys

& Keogh, 1984; Keogh, 1985).

Data analysis

Diet choice was calculated using frequency of occurrence

expressed as a percentage. The absolute prey frequency

98 T. Breuer

� 2005 African Journal of Ecology, Afr. J. Ecol., 43, 97–106

(n/N) is the number of prey items of a species (n) in relation

to the faecal samples analysed (N). The relative frequency

(r/R) is the relation of identified prey items (r) to the number

of all prey items found during analysis (R) (Hoppe-Dominik,

1984). No attempt was made to convert the frequency of

occurrence into relative biomass and number of individuals

consumed (Ackermann, Lindsey & Hemker, 1984). No

correction factor is available for lions and spotted hyenas,

although this approach has been applied in a study of wild

dog diet choice in South Africa (Kruger, Lawes & Maddock,

1999), and elsewhere (Karanth & Sunquist, 1995; Hart

et al., 1996; Henschel, 2001).

Prey weight was calculated from 75% of mean female

weight from Kingdon (1997) according to White (1994).

Items not identified to species level and unidentified items

were excluded from calculations of mean prey weight.

Analysis of variance (ANOVA) was performed to identify

differences in mean prey weight. Diet overlap was calcu-

lated using Pianka’s (1973) index

Oab ¼P

n pia pibP

p2ia p2

ib

� �1=2

where pia is the relative frequency of the item i found in the

diet of species a (pib is the relative frequency of i found in

the faeces of species b). The index ranges from 0 (no

overlap) to 1 (complete overlap).

Carnivore niche breadths were calculated using niche

breadth index B ¼P

n 1/p2i (Levins, 1968). Diet diversity

and evenness were calculated using Shannon–Wiener index

(Pielou, 1977). Species accumulation curves were drawn to

calculate the number of samples needed to reveal all prey

species. Relative frequency of different prey species in lion

and spotted hyena faeces were calculated in increments of

10 samples of all faeces analysed to reveal the minimum

number of samples needed to determine diet choice of both

species correctly (Mukherjee, Goyal & Chellam, 1994a).

Results

Diet choice

In total 280 carnivore faecal samples [Lycaon pictus (L.p.):

n ¼ 13; Panthera leo (P.l.): n ¼ 119; Crocuta Crocuta (C.c.):

n ¼ 148] were analysed and 388 mammalian prey items

(L.p.: n ¼ 18; P.l.: n ¼ 173; C.c.: n ¼201) were found.

Only three prey items in spotted hyena faeces and one item

in lion faeces could not been identified. Plant remains were

found in faeces of both lions and spotted hyenas species,

but were not included in the analysis. A mean of

1.38 ± 0.19 prey items per faecal sample for wild dogs,

1.44 ± 0.05 for lions, and 1.36 ± 0.04 for spotted hyenas

was found with a maximum of three items per faecal

sample for all three carnivores.

Analysis revealed a total of nineteen mammalian prey

species ranging from 0.9 to 412.5 kg body weight with

seven, fourteen and sixteen different species for wild dogs,

lions and spotted hyenas respectively. Mean prey weight

was 63.5 ± 12 kg for wild dogs (range 0.9–188.6 kg),

56.7 ± 4.2 kg (range 10.1–412.5 kg) for lions and

54.4 ± 3.9 kg (range 7.5–12.5 kg) for spotted hyenas.

There was no significant difference in the mean prey

weight of the three large carnivores (ANOVA: F2,384 ¼0.257). All had the same median prey weight of 50.63 kg

(Fig. 1a).

Ungulates made up the biggest portion of the diet of large

carnivores in northern Cameroon (L.p.: 88.9%; P.l. 83.2%;

C.c. 93%) (Fig. 1b). Buffon’s kob was by far the most fre-

quent prey species occurring in carnivore faeces [r/R ¼55.6% (L.p.); 35.3% (P.l.); 59.2% (C.c.)] (Table 1). The

second most important prey for lions and spotted hyenas

was bushbuck (P.l.: 9.3%; C.c.: 9%). Waterbuck accounted

for 16.7% of the diet of wild dogs, 7.5% of lions and 4.5% of

spotted hyenas. Other antelopes consumed were roan

antelope (L.p.: 5.6%; P.l.: 6.9%; C.c.: 2%), oribi (L.p.: 5.6%;

P.l.: 6.4%;C.c.: 5%), Grimm’s duiker (P.l.: 5.2%; C.c.: 4.5%)

and reedbuck (P.l.: 2.9%;C.c.: 3%). Buffalo was only found

in one lion and three spotted hyena samples. Hartebeest

was only found in spotted hyenas samples (C.c.: 1%). Red

river hog are also notably important for lions (P.l.: 6.9%).

Warthogs constituted only 2.3% of the diet of lions and 1%

of the diet of spotted hyenas.

Rodentia, especially crested porcupine, Hystrix cristata

(L.), were found in faeces of all carnivores (L.p.: 7.7%; P.l.:

7.5%; C.c.: 1.5%). A small mammal of the super-family

Muroidea (0.7%) was found in one spotted hyena sample.

The rat-like rodent could not be determined to species or

genus level, because reference material was not available.

The giant pouched rat, Cricetomys gambianus (Waterhouse)

was found in one wild dog sample. Prey remains of pri-

mates were only found in lion (8.1%) and spotted hyena

(3.5%) samples. Baboons accounted for 6.9% of the

occurrences in lion and 2.5% of the items in spotted hyena

faeces. Additionally one carnivore species [Civettictis civetta

(Schreber)] was found in one lion sample. Medium-sized

prey (25.1–100 kg) was found in the majority of carnivore

faeces and makes up the biggest portion in their diet

Diet of large carnivores in Cameroon 99


(L.p.: 55.6%; P.l.: 57%; C.c.: 73.2%) (Fig. 1c). Large prey

was found more often in lion (15.1%) than in spotted

hyena faeces (9.1%), but was most common in wild dog

faeces (22.2%), possibly because of the low sample size.

Dietary diversity and overlap

The diet of the three sympatric carnivores overlaps

extensively. Pianka’s dietary overlap index was extremely

high: 92% between wild dog and lion, 95% between wild

dog and spotted hyena and 93% between lions and hyenas.

Niche breath Index calculated for nineteen different prey

species was greatest for lions (B ¼ 5.99) and similar for

wild dogs (B ¼ 2.84) and spotted hyenas (B ¼ 2.71).

Diversity and evenness index for lions (H¢ ¼ 2.162, J¢ ¼0.819) was higher than for wild dogs (H¢ ¼ 1.432, J¢ ¼0.736) and for spotted hyenas (H¢ ¼ 1.616, J¢ ¼ 0.583)

(Table 1).

Faecal accumulation curves

Faecal accumulation curves showed that 75% of all prey

species were detected after analysing 30 lion (Fig. 2b) and

30 spotted hyena faeces (Fig. 2c). Species accumulation

curves reached asymptote after 60 samples for lions and

110 samples for spotted hyena, but did not asymptote for

wild dogs in this study (Fig. 2a). All prey species in hyena

faeces were found after analysing 100 faeces. Buffalo was

found in one of the last lion samples analysed.

Table 1 Frequency of occurrence of different prey species of large

carnivores in northern Cameroon, revealed by faecal analysis

Species

Wild dog

(n ¼ 13)

(%)

Lion

(n ¼ 119)

(%)

Spotted hyena

(n ¼ 148) (%)

Artiodactyla

Potamochoerus porcus – 6.9 –

Phacochoerus africanus – 2.3 1.0

Cephalophus rufilatus – – 2.5

Ourebia ourebi 5.6 6.4 5.0

Sylvicapra grimmia 5.6 5.2 4.5

Tragelaphus scriptus – 9.2 9.0

Redunca redunca – 2.9 3.0

Kobus ellipsipymnus 16.7 7.5 4.5

Kobus kob 55.6 35.3 59.2

Alcelaphus buselaphus – – 1.0

Hippotragus equinus 5.6 6.9 2.0

Syncerus caffer – 6.0 1.5

Primates

Papio anubis – 6.9 2.5

Colobus guereza – 1.2 0.5

Cercopithecus tantalus – – 0.5

Carnivora

Civettictis civetta – 0.6 –

Rodentia

Hystrix cristata 5.6 7.5 1.5

Cricetomys gambianus 5.6 – –

Muridae

uk – – 0.5

Not identified – 0.6 1.5

0

20

40

60

80

Wild dog Lion Spotted hyena

Pre

y w

eig

ht

(kg

)

Mean prey weight (kg)Median prey weight (kg)

0

20

40

60

80

100

Artiodactyla

Rodentia

Primates

Muridae

Carnivora

Not identifie

d

Fre

qu

ency

of

occ

ure

nce

(%

) Wild dog (n = 13)Lion (n = 119)Spotted hyena (n = 148)

0

20

40

60

80

Wild dog (n = 13) Lion (n = 119) Spotted hyena(n = 148)

Fre

qu

ency

of

occ

ure

nce

(%

)

0–55.1–2525.1–100> 100

(a)

(b)

(c)

Fig 1 Mean prey weight (±SE) (a), frequency of occurrence of

different mammal orders (b), and relative proportion of different

prey weights (kg) (c) consumed by large carnivores in Faro

National Park, northern Cameroon

100 T. Breuer


There was little variation in the frequency of occurrence

of the five most important prey species (comprising 66.5

and 82.1% of the prey items of lions and spotted hyenas)

after 70 lion samples (Fig. 3a) and 70 spotted hyenas

samples (Fig. 3b), although variation of Buffon’s kob in the

diet of spotted hyena samples was more than 8% even after

70 samples.

Discussion

Sample size

In contrast to many lion and spotted hyena samples, only

thirteen wild dog faeces could be analysed during the

whole study period for several reasons. Wild dogs live at

lower densities than the other carnivores (Creel & Creel,

1998), especially in northern Cameroon (Breuer, 2003).

Their faeces are rare and difficult to find, because wild dogs

of one pack often defecate at their daily resting-place in

thickets with limited access (Woodroe et al., 1997). In

contrast the faeces of spotted hyenas are visible from a

distance by their white colour, and many faeces can be

collected at latrine sites (Kruuk, 1972; Sillero-Zubiri &

Gotelli, 1992). Like leopards (Ramakrishnan et al., 1999;

Parnell, 2000), lions defecate on prominent places and in

the sand of seasonal riverbeds (Bosch, 1976).

Most of the prey species were detected after analysing 30

samples. After analysing 70 lion samples there was no

great variation in the frequency of occurrence of different

prey species. We recommend at least 70 samples to

determine lion and spotted hyena diet choice, but even

after analysis of more than 100 faeces, one might find

more prey species. Variation in the diet choice of spotted

hyenas might be caused by local variation in prey num-

bers, because the last samples were collected in the eastern

tip of FNP, which harbours the highest numbers of Buf-

fon’s kob (Breuer, 2002). The sample size for wild dogs is

too low to indicate the number of samples needed to reveal

all prey species. The number of wild dog samples in this

study might not be large enough for a reliable estimate and

it might give a false impression of diet composition and

mean prey weight. A study of hyenas showed that 25

faeces per month are a representative number to give ro-

bust results on diet choice (Henschel & Skinner, 1990).

Other authors recommend at last 30 faecal samples for

lions in Asia (Mukherjee, Goyal & Chellam, 1994b). Kru-

ger et al. (1999) have analysed 78 wild dog faeces for

determining diet choice.

Diet choice

Wild dogs mainly prey on medium-sized antelopes (Fuller

& Kat, 1990; Mills & Biggs, 1993; Creel & Creel, 1995;

Kruger et al., 1999). Although only thirteen wild dog

faecal samples were analysed, Buffon’s kob clearly made up

the largest portion of wild dogs’ diet. Frequency of occur-

rence of Buffon’s kob in lion faeces was proportionally

lower but still comprised more than a third of all prey

items. The higher B-value for lions shows a more diversi-

fied diet than that of spotted hyenas and wild dogs. Similar

feeding behaviour has been found in another study area

in Central Africa (Ruggiero, 1991) and elsewhere (Van

Orsdol, 1984), where lions hunted most of the time near

water, killing porcupines and all kinds of antelopes and

with a preference for kob and hartebeest. Lions in FNP are

associated with riparian forest and they depend on prey

012345678

0 1 2 3

(a)

(b)

(c)

4 5 6 7 8 9 10 11 12 13No. of faeces analysed

No. of faeces analysed

Cu

mu

lati

ve n

o.

of

spec

ies

Cu

mu

lati

ve n

o.

of

spec

ies

Cu

mu

lati

ve n

o.

of

spec

ies

No. of faeces analysed

02468

10121416

0 10 20 30 40 50 60 70 80 90 100

110

119

0

5

10

15

20

0 20 40 60 80 100

120

140

Fig 2 Wild dog (a), lion (b), and spotted hyena (c) species/sample

size curves revealed by faecal analysis



near water. Lions also eat small antelopes, but depend on

large and medium-sized ungulates (Mills & Biggs, 1993).

Diet choice of baboons and other carnivores might reflect

lions’ ability to kill potential competitors (see below).

Spotted hyenas’ main prey was also Buffon’s kob, poss-

ibly because it was the most abundant prey species in and

around FNP (Breuer, 2002). Hyenas are opportunistic

predators, feeding on the most abundant prey (Holecamp

et al., 1997). Common ungulate species are eaten more

often than rare species and diet choice of spotted hyenas is

related to ungulate size, abundance, and availability

(Henschel & Skinner, 1990; Di Silvestre et al., 2000).

Additionally Buffon’s kob is often found in cable snares in

and around FNP (Breuer, 2002.). Game which is captured

in snares, is often taken by scavenging carnivores (Noss,

1998).

Although hartebeest is the second most numerous prey

species in FNP (Gomse & Mahop, 2000), no sign of its

presence in lion and wild dog faeces was found. The low

sample size for wild dogs might be a reason for this, but

cannot explain the absence in lions’ diet. One reason is their

spatial distribution in FNP. Most of the hartebeests are found

in the centre and south of FNP (Gomse & Mahop, 2000), and

they are rarely seen in the northern part (Breuer, 2002).

In contrast to findings in West Africa (Di Silvestre et al.,

2000), mainly medium-sized antelopes, especially Buffon’s

kob are consumed by large carnivores in FNP, which

confirms findings in other areas in Central Africa (Wanzie,

1986; Ruggiero, 1991; Baggett, 1998). It is not clear if

Buffon’s kob is a favoured prey species or is just chosen

because it is abundant and other antelopes are rare. Their

spatial distribution may have an influence on carnivore

distribution, because Buffon’s kob is always found in close

proximity to water (Breuer, 2002). Only a few monkey

prey items were found because they are difficult to capture

(Di Silvestre et al., 2000). In contrast to a study in Senegal

(Di Silvestre et al., 2000), few buffalo and other large prey

species were found in the faeces of carnivores in FNP.

Large mammals are rare in FNP and seem to be subject to

heavy poaching (Breuer, 2002).

0

0.1

0.2

0.3

0.4

0.5

0 20 40 60 80 100

119

Kobus kob

(a)

(b)

Tragelaphusscriptus

Kobusellipsipymnus

Hystrixcristata

Hippotragusequinus

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0 20 40 60 80 100

120

140

Kobus kob

Kobusellipsipymnus

Ourebia ourebi

Sylvicapragrimmia

Tragelaphusscriptus

Fig 3 Cumulative frequency of occurrence

of the five most common prey species of

lions (a) and spotted hyena (b) as related to

the number of faecal samples analysed in

and around Faro National Park, northern

Cameroon

102 T. Breuer


As in other parts of northern Cameroon (Bauer, 2003)

large carnivores near FNP were reported having killed

livestock (Breuer, 2002). No domestic animal items were,

however, found in the carnivore faeces. Generally sheep

and goat are subject to predation by spotted hyenas, and

lions prey on larger livestock (Kruuk, 1972). Lions usually

kill livestock at night in northern Cameroon (Breuer,

2002), although people report that lions around Waza

National Park mostly kill cattle during daytime when

livestock is moving to and from drinking points and only

occasionally at night in the villages (Bauer & Kari, 2001).

Hyenas kill livestock at night only (Breuer, 2002) and

more often in the rainy season (Bauer & Kari, 2001). Wild

dogs in northern Cameroon are often blamed and killed for

livestock losses, especially by traditional herdsmen. As a

result, they have declined dramatically in northern

Cameroon (Breuer, 2003). Even after a reported case of

livestock predation north of FNP, no signs of a livestock kill

could be attributed to wild dogs (Breuer, 2001).

Diet overlap and feeding competition

Indices show that the diets of the three carnivores overlap

substantially, which would suggest high feeding competi-

tion. Major prey species are concentrated along water-

courses (Breuer, 2002) and exploitation competition

(sensu Mills, 1991) near water is expected. Several studies

have shown that the diets of large carnivores overlaps

strikingly (Kruuk & Turner, 1967; Mills & Biggs, 1993)

and other factors, such as preferred habitat and hunting

period promote ecological separation (Creel, Spong & Creel,

2001). Baboons also hunt medium-sized antelopes in FNP

and hair of Buffon’s kob and bushbuck was found in their

faeces (Breuer, 2001). As they are very abundant in the

park and adjoining hunting zones, their role as an indirect

competitor is not negligible.

The potential of faecal analysis for carnivore research

in northern Cameroon

Several methods have been used to study feeding ecology,

and each of these methods has advantages and disadvan-

tages (Mills, 1992). In this study diet choice was deter-

mined by faecal analysis. Faecal analysis allows a

continuous determination of diet choice (Putman, 1984),

and has been used in many different habitats to determine

feeding ecology of several carnivore species (Hoppe-Domi-

nik, 1984; Henschel & Skinner, 1990; Mukherjee et al.,

1994a; Karanth & Sunquist, 1995; Hart et al., 1996;

Kruger et al., 1999).

As in other studies, a maximum of three prey species

was found in a single faecal sample, demonstrating that

items from the same prey species are found in the faeces for

up to 5 days (Reynolds & Aebischer, 1991; Bothma & Le

Riche, 1994). A mean of 1.4 prey species was found in the

thirteen faecal samples of wild dogs, which is similar to the

results of Kruger et al. (1999), who found a mean 1.5

species in the 78 samples analysed. Mukherjee et al.

(1994b)) found that 72% of faecal samples contained only

one prey species, which is larger than my findings (56%)

but may reflect the low number of large prey in lion faeces

in northern Cameroon. The findings of prey items in

spotted hyena faeces confirm the results of Bearder (1977),

who found 1.51 prey items in spotted hyena faeces in a

woodland savanna in southern Africa.

Direct observation is mostly used to document lion

feeding behaviour (Mills, 1992). Additionally there exists

no correction factor to estimate the relative proportion of

biomass of different prey species and individuals consumed

by lions. Faecal analysis has been widely used to determine

diet choice of spotted hyenas (Henschel & Skinner, 1990;

Sillero-Zubiri & Gotelli, 1992; Di Silvestre et al., 2000;

Korb, 2000). Quantitative data from hyena faeces is open

to a number of pitfalls, because of their scavenging beha-

viour and regurgitation (Kruuk, 1972). Hyenas generally

hunt large animals in groups but forage individually for

small prey (Holecamp et al., 1997). Individuals travel

separately or in cohesive groups (Bearder, 1977; Sillero-

Zubiri & Gotelli, 1993; Holecamp et al., 1997). Korb

(2000) assumed that spotted hyenas in west Africa do not

live in groups. Faecal analysis does determine if prey was

killed or scavenged. Buffalo, mostly scavenged, proved to

be overestimated using faecal analysis, while medium-sized

ungulates were underestimated (Henschel & Skinner,

1990). Items from a small mammal (<2 kg) were found in

one sample. Kruger et al. (1999) also found small antelope

items (C. rufilatus) in the faeces, although direct observa-

tions could not confirm this. This is typical of faecal ana-

lysis, which often identifies small prey species, which are

underestimated in studies using kill samples or are over-

looked by direct observations (Mills, 1992; Karanth &

Sunquist, 1995). The presence of a roan antelope in the

small sample size of wild dogs contributes to the high mean

prey weight. Faecal analysis neither gives information on

the age and the sex of the prey nor does it decide if the

animal was hunted or scavenged.



Faecal analysis seems to be a good method to reveal diet

choice of large carnivores in northern Cameroon and in

similar habitats (Korb, 2000). Many protected areas in

northern Cameroon, especially FNP have no or limited road

access and direct observations of the carnivores are very

rare. Faeces are much more often encountered and can

give further information on genetic structure, reproduc-

tion, sex-ratio and parasitology (Kohn & Wayne, 1997).

Conclusion

Competition between carnivores is an important factor

affecting structure of carnivore communities (Woodroe,

2001). Knowing carnivores’ feeding ecology helps docu-

ment the impact on prey species, interspecific competition,

and livestock kills. Further investigations are needed to

determine which differences between the large carnivores

permit their coexistence. Additionally, information is

required on seasonal variation in feeding ecology and diet

choice. Antelopes have a different spatial distribution and

group size in the rainy season (Stark, 1986; Tsague, 1996;

Fischer & Linsenmaier, 2001). It would be interesting to

determine if this has an influence on the diet choice, dis-

tribution and social behaviour of large carnivores as

shown in other parts of Africa (Van Orsdol, 1984; Dun-

ham, 1992; Hanby, Bygott & Packer, 1995; Mills, 1995).

Livestock predation must be analysed in detail, including

kills on cattle migration routes.

A detailed evaluation of the effect of large carnivores on

the abundance and distribution of prey species can only be

answered if the densities and distribution of all large

carnivore species are known. Poaching may have an

enormous effect on the diet choice of large carnivores in

FNP (Breuer, 2002). Detailed information on distribution

and numbers of large carnivores will contribute substan-

tially to the understanding their behavioural ecology in

northern Cameroon (Woodroe et al., 1997; Mills & Hofer,

1998; Bauer et al., 2003). This is valuable data to facilitate

appropriate management for carnivores (Ormerod, 2002),

especially for the conservation of the highly endangered

African wild dog (Breuer, 2003).

Acknowledgements

We thank the delegation of the Ministry of Environment

and Forests in the North Province for the permit to conduct

this study in and around FNP. We would like to thank

WWF-NSSP (Northern Sudanian Savannah Project) for

invaluable logistical support. Fieldwork and laboratory

work at the Garoua Wildlife College was possible by

financial support from WWF and North Carolina Zoologi-

cal Park. We are grateful to all the trackers and game

guards, especially Mr Wassou and other rural people for

providing valuable information and their hospitality. Spe-

cial thanks to Ndjobdi Seini, the conservator of FNP.

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106 T. Breuer


Diet Choise of Large Carnivores in Nothern Cameroon

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