feeding ecology of the ethiopian wolf in the simien mountains national park, ethiopia

Feeding ecology of the Ethiopian wolf in the SimienMountains National Park, Ethiopia

Mesele Yihune1* and Afework Bekele2

1Department of Biology, Debre Markos University, P.O. Box 269, Debre Markos, Ethiopia, 2Department of Zoological Sciences, Addis Ababa

University, P.O. Box 1176, Addis Ababa, Ethiopia

Abstract

Feeding ecology of the Ethiopian wolf was studied in the

Simien Mountains National Park, Ethiopia, during 2011–

2012 using faecal analysis. A total of 214 faecal samples

were collected from the study area. Data were analysed

using descriptive statistics, and a chi-square test was used

to test seasonal differences in the frequency of prey item

consumed. Percentage frequency of occurrence/scats and

percentage frequency of occurrence/item were calculated.

A Chesson index of prey preference was also calculated.

Eight categories of prey items were identified from the

Ethiopian wolf faecal samples. These comprised five species

of rodents, sheep, birds and grass. Rodents occurred in

most of the samples, and they were the most important

prey items in their frequency of occurrence. Among the

diurnal murid rodents (Arvicanthis abyssinicus, Lophuromys

flavopunctatus and Otomys typus), A. abyssinicus (72%) was

the most common species in the diet followed by L. flavo-

punctatus (24.3%). Arvicanthis abyssinicus was the most

highly preferred prey item, with a Chesson index value of

0.3. There was no significant seasonal difference

(v2 = 2.03, d.f. = 1, P > 0.05) in the frequency of prey

items recorded during the wet and dry seasons. As rodents

are the main prey source of the Ethiopian wolf, they have

to be protected to conserve the Ethiopian wolf in the study

area.

Key words: diet, Ethiopian wolf, prey preference, rodents,

Simien Mountains National Park

R�esum�e

L’�ecologie alimentaire du loup d’�Ethiopie a �et�e �etudi�ee dans

le Parc National des Simien Mountains, en �Ethiopie, en

2011-2012, par l’analyse des excr�ements. Au total, 214

�echantillons de crottes ont �et�e collect�es dans la zone

d’�etude. Les donn�ees ont �et�e analys�ees au moyen de

statistiques descriptives, et un test de v² a servi �a tester les

diff�erences saisonni�eres de la fr�equence des proies con-

somm�ees. Le pourcentage de la fr�equence d’occurrence/

crottes et celui de la fr�equence d’occurrence/�el�ement ont

�et�e calcul�es. Nous avons aussi calcul�e un indice de Chesson

pour la pr�ef�erence d’une proie. Huit cat�egories d’�el�ements

consomm�es ont �et�e identifi�ees �a partir des �echantillons de

crottes de loup d’�Ethiopie. Elles comprennent cinq esp�eces

de rongeurs, des moutons, des oiseaux et de l’herbe. Les

rongeurs �etaient pr�esents dans la plupart des �echantillons

et ils �etaient les �el�ements consomm�es les plus importants

par la fr�equence de leur occurrence. Parmi les rongeurs

murid�es diurnes identifi�es (Arvicanthis abyssinicus, Loph-

uromys flavopunctatus and Otomys typus), A. abyssinicus

�etait l’esp�ece la plus fr�equente dans le r�egime alimentaire

(72%), suivi de L. flavopunctatus (24.3%). A. abyssinicus

�etait la proie favorite, avec un indice de Chesson de 0,3. Il

n’y avait pas de diff�erence saisonni�ere significative (v2 =

2,03, df = 1, P > 0,05) dans la fr�equence des �el�ements

consomm�es en saison s�eche et en saison des pluies. Comme

les rongeurs sont les proies principales du loup d’�Ethiopie,

ils doivent etre prot�eg�es pour pr�eserver ce dernier dans la

zone �etudi�ee.

Introduction

The Ethiopian wolf commonly preys on small mammals

and is specialized in rodent hunting (Sillero-Zubiri, 1994).

Unlike other medium-sized to large canids, the Ethiopian

wolf has developed an extreme feeding specialization on

high altitude rodents, with a narrow ecological range and

habitat specificity. Their long muzzles, long legs and small

and well-spaced teeth are highly adapted for their feeding*Correspondence: E-mail: [email protected]

484 © 2014 John Wiley & Sons Ltd, Afr. J. Ecol., 52, 484–490

habits and help them to extract their prey from burrows.

Hunting behaviour of the Ethiopian wolf is more similar to

felids than other canids (Sillero-Zubiri & Gottelli, 1995a).

They feed upon rodents such as giant mole rat (Tachyoryctes

macrocephalus), the common mole rat (Tachyoryctes

splendens) and other rodent species such as Arvicanthis

abyssinicus, Arvicanthis blicki, Lophuromys melanonyx, Loph-

uromys flavopunctatus and Otomys typus. Besides rodents,

the Ethiopian wolf rarely feeds on rock hyraxes, lambs and

hares (Sillero-Zubiri, 1994; Sillero-Zubiri & Gottelli,

1995b). Occasionally, it feeds upon sedge (Carex monostac-

hya), which may assist indigestion and control intestinal

parasite (Sillero-Zubiri, 1994). Due to their feeding special-

ization on rodent fauna (highly restricted in space), they are

at present categorized as threatened species in the IUCN

Red List of Threatened Animals (IUCN, 2012).

The Ethiopian wolf utilizes different hunting techniques

in accordance with the size of the prey and forages

throughout the day. The Ethiopian wolf carefully explores

rich food patches and then walks slowly and investigates

holes to localize rodents by hearing. Once the prey is

located, the wolf moves slowly towards it. This may take

up to an hour. Sometimes, the Ethiopian wolf runs in a

zigzag pattern across rodent colonies to pick rodents up

while passing. To dig out rodent holes is also a common

hunting habit in Ethiopian wolf. Activities of the Ethiopian

wolf are usually synchronized with the activities of rodents

in their habitat (Sillero-Zubiri & Gottelli, 1995a; Tefera,

2001). Feeding specialization of the Ethiopian wolf and its

dependence on food resources are highly structured in

space (Marino, 2003). Prey can vary spatially and

seasonally, which can presumably determine their avail-

ability (Huitu, 2000). Ethiopian wolves forage solitarily in

their Afro-alpine habitat (Sillero-Zubiri & Gottelli, 1995a).

This makes them unique in comparison with other similar-

sized canids that are generalist and widely distributed

(Macdonald, 1992). However, rarely, Ethiopian wolves are

facultative cooperative hunters. In areas where resources

are plentiful, Ethiopian wolves appear to tolerate their

relatives that remain in the pack (Kruuk & Macdonald,

1985). This condition helps them to form groups to gain

more prey-rich habitat per individual. In contrast to this,

in the area where resources are scarce, groups are smaller,

typically consisting of pairs of individuals (Marino, 2003).

There are ecological differences in the structure of rodent

communities between different habitats in Ethiopia, nota-

bly the absence of giant mole rat from the central and

north-western blocks (Yalden, Largen & Kock, 1976;

Yalden & Largen, 1992). Hence, the ecology of Ethiopian

wolves in different highlands is also different. Detailed

study on the Ethiopian wolf feeding habit in relation to

prey abundance and preference has not been conducted

before in the Simien Mountains National Park. Therefore,

the present study identifies the diet of the Ethiopian wolf, to

explain seasonal variations in the diet and to determine its

prey abundance and preference in the Simien Mountains

National Park, Ethiopia.

Study area and methods

Study area

The Simien Mountains National Park (SMNP) is part of the

Simien Mountains (between 380000 - 380120E and

130120 - 130190N) in the North Gondar Zone of the

Amhara National Regional State of Ethiopia (Fig. 1). The

total area of SMNP is 412 km2. It consists of chains of

plateaus and grassy plains. The area includes the highest

peak in Ethiopia, Ras Dejen Mountain (4543 m asl). The

massif was formed 25 million years ago (Hurni, 1986).

The Simien Mountains National Park represents one of

the most marvellous natural areas in the world. The

presence of a high number of endemic species, unique

biophysical features and its international significance led

SMNP to become a World Heritage Site in 1978. In 1996,

it was listed as World Heritage Site in Danger due to the

declining number of Walia ibex (Capra walie), agricultural

encroachment, loss of biodiversity and impacts of road

construction (Falch & Keiner, 2000). However, at present,

the number of Walia ibex is increasing from time to time

due to different conservation practices (Ejigu, 2012).

There are different types of soil in the SMNP. The

difference in soil types is mainly caused by differences in

land-use practice, geological events, topography and

climate (Falch & Keiner, 2000). The climate of the SMNP

varies from area to area. Generally, highlands have a

relatively low temperature. They are cold in the early

mornings of the dry season (Hurni, 1986). The SMNP is

part of the Afro-alpine centre of plant diversity with high

level of endemism (Debonnet, Melamari & Bomhard,

2006). The Park consists of a representative area of the

Ethiopian tropical highland biome including the vegeta-

tion characteristics (Debonnet, Melamari & Bomhard,

2006). It harbours various endemic animals due to its

unique topography and ecosystem characteristics (Hurni,

1986).

© 2014 John Wiley & Sons Ltd, Afr. J. Ecol., 52, 484–490

Feeding ecology of the Ethiopian wolf 485

Methods

Feeding ecology of the Ethiopian wolf was studied during

2011–2012 using faecal dropping analysis. A total of

214 faecal samples of the Ethiopian wolf were collected

from different sites in the study area during wet and dry

seasons of the study period. Information such as sample

sites, date of collection, approximate age of faeces, time of

collection, altitude of the site of collection, habitat type

and global positioning system (GPS) coordinates were

recorded for each of the samples and was labelled. Age of

faecal samples was grouped into fresh (a day old), recent

(2–5 days old) and old (>5 days) (Breuer, 2005). Very

old faecal droppings were not collected, rather fresh and

recent faecal samples were considered in the analysis.

Ethiopian wolf droppings were distinguished from other

sympatric carnivores in the study area (common jackal

and domestic dogs) based on shape, colour, ingested hair,

diameter and odour following Breuer (2005). In addition

to this, faecal samples were not collected from areas of

human activity to reduce the risk of collecting domestic

dog faeces. Faecal samples were not collected from nearby

locations (every 400 m) to avoid samples from the same

individual.

The collected samples were sun-dried, ground in a

mortar, and thoroughly washed using hot water and

sieved in a sieve (0.5 mm) to separate macrocomponents

such as hairs, bones, claws and teeth and nonfood items

from organic materials. All separated macroscopic mate-

rials from scats were sorted out and categorized as hair,

bone, tooth, flesh (skin, meat and tendon), grass and

nonfood items such as plastic materials, following

Reynolds & Aebischer (1991). Separated hairs were

washed in acetone, dehydrated in 100% ethanol and dried

on filter paper. Hair was analysed macroscopically using

hand lens and type, length and colour (Breuer, 2005).

Sample hairs were compared with rodent specimen in

Zoological and Natural History Museum, Addis Ababa

University. Presence of any of the body parts of rodents

was recorded for each of the samples analysed.

To estimate the biomass of rodents and prepare reference

collection, a rodent trapping study using capture–mark–

recapture method was conducted. A permanent

70 9 70 m live trapping grid was established in five

different habitat types for three consecutive days and nights

repeatedly during the wet and dry seasons. Grids were

established in five different habitat types, namely Festuca-

Lobelia, Lobelia stand, Festuca grassland, Helichrysum-

Festuca and Festuca-Carex grassland. In each trapping site,

a standard square (seven rows by seven columns) trapping

grid was established during wet and dry seasons (Linzey &

Kesner, 1997). A total of 49 Sherman live traps

(5.5 9 6.5 9 16 cm) were set per grid at every 10 m

interval between points. Traps were baited with peanut

butter and barley flour. Traps were covered with leaves and

grasses to prevent death of rodents due to extremely cold

weather and to avoid damage of traps by otherwild animals.

Traps were checked twice a day early in the morning

(between 06:00 and 07:00 hours) and late in the afternoon

(between 17:00 and 18:00 hours). Animals caught from

Fig 1 Map of the Simien Mountains

National Park and the study sites


486 Mesele Yihune and Afework Bekele

06:00 to 18:00 hours were recorded as diurnal, and those

trapped from 18:00 to 06:00 hours were nocturnal. Each

trapped animal was identified, marked by toe clipping and

released in the site from where it was trapped (Linzey &

Kesner, 1997; Clausnitzer, 2003). Information such as

weight, sex, approximate age (juvenile, subadult and adult)

based on their weight and pelage colour and reproductive

condition was recorded (Bekele, 1996).

Data analysis

Data were analysed using frequency of food item and

volumetric methods following Zabala & Zuberogoitia

(2003). Percentage frequency of occurrence/scats (%FO/

S) was calculated. Statistical analyses were performed with

percentage frequency of occurrence/item (%FO/I). %FO/S

is the frequency by which a food item occurs in the scat

sample, whereas %FO/I is the number of each prey species

divided by the total number of items identified where item

is defined as an occurrence of a prey species in an

individual faecal sample (Angerbjorn, Tannerfeldt &

Erlinge, 1999).

To determine preference for each rodent prey category,

their volume in the diet and their biomass were used to

calculate a Chesson index (Chesson, 1983; Vos, 2000) as

follows:

l ¼ rn�1ðXm

j¼1

rjn�1j Þ�1

Where r = the volume of each prey category in the diet;

n = relative biomass of the sameprey category in the area;

m = number of prey categories;

l = Chesson index of preference.

The sum of l across all possible species or prey

categories equals 1, and the larger the value for individual

species or prey category, the more preferred is the prey.

Biomass of each rodent prey species was calculated

using density multiplied by average weight of each species

in the study area.

Results

Eight categories of prey items were identified from the

Ethiopian wolf droppings analysed (n = 214). These com-

prised six species of mammals including five species of

rodents and sheep, birds and grass. Rodents occurred in

most samples, and they were the most predominant prey in

terms of frequency of occurrence. Nonfood items such as

plastics and cigarette paper were also identified in the

scats.

Among the diurnal murid rodent species (A. abyssinicus,

L. flavopunctatus and Otomys typus), A. abyssinicus (72%)

was the most common species in the diet followed by

L. flavopunctatus (24.3%), and O. typus (14%) was the least

common murid rodent species in the diet. The nocturnal

Stenocephalemys griseicauda (18.7%) was the second least

common murid rodent in the diet of the Ethiopian wolf

(Table 1).

In considering the frequency of occurrence per item

(%FO/I), A. abyssinicus (37.1%) had the highest proportion

among murids followed by L. flavopunctatus (12.5%),

whereas O. typus (7.2%) accounted the least percentage

by volume. Common mole rat, T. splendens, accounted for

13% by volume. On the other hand, sheep contributed

14.5% of the diet by volume (Table 2).

There was no significant seasonal difference (v2 = 2.03,

d.f. = 1, P > 0.05) in the frequency of prey items between

Table 1 Frequency occurrence of prey items of the Ethiopian wolf

in the SMNP

Prey categories

Frequency of occurrence/scats (%FO/S)

n (%)

A. abyssinicus 154 (72.0)

L. flavopunctatus 52 (24.3)

S. griseicauda 40 (18.7)

O. typus 30 (14.0)

T. splendens 54 (25.2)

Sheep 60 (28.0)

Bird 10 (4.8)

Grass 15 (7.0)

Table 2 Percentage volume of prey items of the Ethiopian wolf in

the SMNP using faecal dropping samples (%FO/I)

Prey categories

Frequency of occurrence/ /item (%FO/I)

n (%)

A. abyssinicus 154 (37.1)

L. flavopunctatus 52 (12.5)

S. griseicauda 40 (9.7)

O. typus 30 (7.2)

T. splendens 54 (13.0)

Sheep 60 (14.5)

Bird 10 (2.4)

Grass 15 (3.6)

Total 415 (100.0)



the wet and dry seasons, and so, all the data are merged

together irrespective of seasons.

Prey abundance and preference

The four common species of rodents (A. abyssinicus,

L. flavopunctatus, O. typus and S. griseicauda) accounted

for 66.5% of the prey by volume. The biomass of these

rodents in five different habitats was estimated from the

live trap data during rodent surveys. The biomass of

T. splendens was also estimated (Table 3).

Arvicanthis abyssinicus was the most highly preferred

prey item, with a Chesson index value of 0.3. O. typus, and

L. flavopunctatus was the second highly preferred prey

items, with a Chesson index value of 0.24 each. T. splen-

dens was the third preferred prey item, with a Chesson

index value of 0.2. The nocturnal S. griseicauda was the

least preferred species among the rodents, with a Chesson

index value of 0.05.

Discussion

Knowledge of diet composition of predators is fundamental

for understanding their ecology (Scheiffarth, 2001). Canids

are typically generalist feeders (Macdonald, 1992). How-

ever, rodents are the primary diet of the Ethiopian wolf

which accounted for a total of 79.5% by volume of all prey

consumed in the SMNP. Similarly, research conducted in

Guassa area of Menz, Ethiopia, indicated that the primary

diet of Ethiopian wolves constituted rodents which

accounted for 88.1% by volume (Tefera, 2001). Marino

(2003) also noted that murid rodents dominated the diet of

the Ethiopian wolf in five study areas (Simien Mountains,

north Wollo, south Wollo, southern highlands (Arsi) and

Bale Mountains) and they accounted for 96% of all prey

occurrences in the scats together. In addition to this,

previous study in the SMNP also showed that rodents

constituted 97.8% of the Ethiopianwolf diet (Yihune, Bekele

& Tefera, 2008). Ethiopian wolves are believed to be rodent

hunters across their ranges due to their habitat and food

specificity (Sillero-Zubiri, 1994).

Three species of diurnal murids (A. abyssinicus, L. flavo-

punctatus and O. typus) were the main prey type that

accounted for 56.5% by volume, while the common mole

rat, T. splendens, accounted for 13% by volume. According

to Tefera (2001), diurnal murids including A. abyssinicus,

L. flavopunctatus and O. typus were the main prey of the

Ethiopian wolves. T. splendens accounted for 16.6% by

volume in the diet. A study in the Bale Mountains National

Park (Malcolm, 1997) and Guassa area of Menz (Tefera,

2001) revealed the absence of the giant mole rat

(T. macrocephalus) in these areas, and hence, the common

mole rats (T. splendens) formed the diet of the Ethiopian

wolf. Sheep constituted 28% by volume in the diet. This is

an indication that there might be relatively severe sheep

loss to the Ethiopian wolf and an indication of diet shift of

the Ethiopian wolf and increasing conflict with the local

community. The nocturnal murid, S. griseicauda,

accounted for 18.7% by volume in the diet. In the

northern Highlands, where the climate is drier and human

population is denser, Ethiopian wolves frequently feed on

rat-sized prey, including nocturnal species (Marino,

Mitchell & Johnson, 2009). Nonfood items such as plastics

and cigarette paper were also found in the faecal samples

of wolves. This might have resulted from unmanaged

waste disposal system in the Park and wolf habitats by

tourists and Park communities. Such nondigestible mate-

rials may cause death to the Ethiopian wolf, and it can be

considered as a kind of threat unless it is regulated. In the

present study, there were no shrew species encountered in

the faecal samples. Shrews were rejected by the Ethiopian

wolf during food preference trials (Sillero-Zubiri & Gottelli,

1995a). This indicated that foraging activity of the

Ethiopian wolf was related to the activity of rodents

(Sillero-Zubiri & Gottelli, 1995a). Furthermore, it might be

related to the imbalance of cost and gain of energy because

shrew species are too small to replace the energy that the

Ethiopian wolf expends to capture them.

Chesson index indicated that the Ethiopian wolf primar-

ily prefers A. abyssinicus followed by L. flavopunctatus and

O. typus. Similarly, A. abyssinicus was the first preferred

prey item by the Ethiopian wolf followed by L. flavopunct-

atus and T. splendens in Guassa area of Menz (Tefera,

2001). However, T. splendens was the third preferred prey

item by the Ethiopian wolf in SMNP, and S. griseicauda was

Table 3 Biomass (kg/km2) of rodent prey species of the Ethiopian

wolf in the study area

Species Biomass (kg/km2)

A. abyssinicus 705.0

L. flavopunctatus 263.5

O. typus 151.9

S. griseicauda 991.4

T. splendens 321.0

Total 2432.8



the least preferred prey item. Similar results were obtained

in Guassa area where S. griseicauda was poorly represented

in the diet of the Ethiopian wolf (Tefera, 2001). According

to Marino (2003), Ethiopian wolves rarely feed upon

nocturnal rats in the Bale Mountains National Park. There

was no significant seasonal difference (v2 = 2.03, d.f. = 1,

P > 0.05) in the diet of the Ethiopian wolf between wet

and dry seasons as observed in Guassa area (Tefera, 2001).

Although the Ethiopian wolf primarily feeds upon rodents

in the SMNP, the rodent biomass is lower than in the Bale

Mountains National Park and Guassa area of Menz. This

might be related to livestock impact in the area and

topographic, climatic and vegetation variations. According

to the present study, the Ethiopian wolf largely depends on

the rodent community in the SMNP, and hence, rodents

have major ecological importance in the study area.

Hence, any negative impact on the rodent community can

influence the fate of the Ethiopian wolf population in

SMNP and will lead to diet shift to sheep. Therefore,

rodents should be conserved by making their habitat free

from human and livestock interventions to support the

threatened Ethiopian wolf population in the study area.

However, this needs awareness creation and proper

compensation to humans.

Acknowledgements

We would like to thank Addis Ababa University for

providing fund to conduct this study. We are also grateful

to the Ethiopian Wildlife Conservation Authority for

permission to conduct the study in the SMNP.

References

Angerbjorn, A., Tannerfeldt, M. & Erlinge, S. (1999) Predator -

prey relationships: arctic foxes and lemmings. J. Anim. Ecol. 68,

34–49.

Bekele, A. (1996) Population dynamics of the Ethiopian endemic

rodent, Praomys albipes in the Menagesha State Forest. J. Zool.

Lond. 238, 1–12.

Breuer, T. (2005) Diet choice of large carnivores in northern

Cameroon. Afr. J. Ecol. 43, 97–106.

Chesson, J. (1983) The estimation and analysis of preference

and its relationship to foraging models. Ecology 64,

1297–1304.

Clausnitzer, V. (2003) Rodents of Mt. Elgon, Uganda: ecology,

biogeography, and the significance of fire. Ecotrop. Mono. 3,

1–176.

Debonnet, G., Melamari, L. & Bomhard, B. (2006) Reactive

Monitoring Mission to Simien Mountains National Park,

Ethiopia. WHC / IUCN Mission Report, UNESCO, Paris and

IUCN, Gland.

Ejigu, D. (2012) Population status and ecology of walia ibex (Capra

walie): a study to address its conservation in Simien Mountains

National Park, Ethiopia, PhD thesis. Addis Ababa University,

Addis Ababa.

Falch, F. & Keiner, M. (2000) Simien Mountains National Park

Management Plan, Final Draft. Amhara National Regional State,

Bahir Dar.

Huitu, O. (2000)Wolf (Canis lupus) diet and prey species selectivity in

Kainuu, MSc thesis. University of Jyvaskyla, Finland.

Hurni, S.J. (1986) Management Plan: Simien Mountains National

Park and Surrounding Rural Area. UNESCO World Heritage

Committee, Gland.

IUCN (2012) 2012 IUCN Red List of Threatened Species. Available

at: http://www.iucnredlist.org (Accessed on 23 October 2012).

Kruuk, H. & Macdonald, D.W. (1985) Group territories of

carnivores: empires and enclaves. In: Behavioural Ecology.

Ecological Consequences of Adaptive Behaviour (Eds R. M. SIBLY and

R. H. SMITH). Blackwell, Oxford.

Linzey, A.V. & Kesner, M.H. (1997) Small mammals of a woodland

savannah ecosystem in Zimbabwe: density and habitat

occupancy patterns. J. Zool. Lond. 243, 137–152.

Macdonald, D.W. (1992) The Velvet Claw. BBC Books, London.

Malcolm, J. (1997) The diet of the Ethiopian wolf (Canis simensis,

R€uppell) from a grassland area of the Bale Mountains, Ethiopia.

Afr. J. Ecol. 35, 162–164.

Marino, J. (2003) Spatial ecology of the Ethiopian wolf, Canis

simensis, PhD thesis, University of Oxford, UK.

Marino, J., Mitchell, R. & Johnson, P.J. (2009) Dietary

specialization and climatic-linked variations in extant

populations of Ethiopian wolves. Afr. J. Ecol. 48, 517–525.

Reynolds, J.C. & Aebischer, N.J. (1991) Comparison and

quantification of carnivore diet by faecal analysis: a critique,

with recommendations, based on a study of the Fox Vulpes

vulpes. Mamm. Rev. 21, 97–122.

Scheiffarth, G. (2001) The diet of Bar-tailed Godwits Limala

lapponica in the Wadden sea: combining visual observation and

faecal analysis. Ardea 89, 481–494.

Sillero-Zubiri, C. (1994) Behavioural ecology of the Ethiopian wolf

(Canis simensis), PhD thesis, University of Oxford, UK.

Sillero-Zubiri, C. & Gottelli, D. (1995a) Diet and feeding

behaviour of Ethiopian wolves (Canis simensis). J. Mammal. 76,

531–541.

Sillero-Zubiri, C. & Gottelli, D. (1995b) Spatial organization in the

Ethiopian wolf Canis simensis: Large packs and small stable

home ranges. J. Zool. Lond. 237, 65–81.

Tefera, Z. (2001) Common property resource management of an Afro

alpine habitat: supporting a population of a critically endangered

Ethiopian wolf (Canis simensis), PhD thesis, Durrel Institute of

Conservation and Ecology, University of Kent, UK.

Vos, J. (2000) Food habits and livestock depredation to two

Iberian wolf packs (Canis lupus signatus) in the north of Portugal.

J. Zool. Lond. 251, 457–462.



Yalden, D.W. & Largen, M.J. (1992) The endemic mammals of

Ethiopia. Mamm. Rev. 22, 115–150.

Yalden, D.W., Largen, M.J. & Kock, D. (1976) Catalogue of the

mammals of Ethiopia. Insectivora and Rodentia. Mon. Zool. Ital.

NS Suppl. 8, 1–118.

Yihune, M., Bekele, A. & Tefera, Z. (2008) Human-Ethiopian wolf

conflict in and around the Simien Mountains National Park,

Ethiopia. Int. J. Ecol. Env. Sci. 34, 149–155.

Zabala, J. & Zuberogoitia, I. (2003) Badger,Meles meles (Mustelidae,

Carnivora), diet assessed through scat-analysis: a comparison

and critique of different methods. Folia Zool. 52, 23–30.

(Manuscript accepted 27 April 2014)

doi: 10.1111/aje.12150



feeding ecology of the ethiopian wolf in the simien mountains national park, ethiopia

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