fragmented populations of leopards in west-central africa: facing … · 2019. 9. 3. · fragmented...

Fragmented populations of leopards inWest-Central Africa: facing an uncertain future

Pauline Toni & Thierry Lodé*

UMR CNRS 6552, Université de Rennes 1, 35042 Rennes Cedex, France

Received 30 January 2012. Accepted 25 September 2013

During recent decades, most endangered species have suffered serious population declines.Little has been documented on leopards in West-Central Africa and as a result the efficiency ofprotection measures and wildlife managing practices can be questioned. Using 416 occurrencesof leopards, we investigated the relationship between different environmental factors andleopard distribution to establish a baseline distribution of this feline. Leopards are mainlypresent in two large populations: one in a forest habitat, the other in a savanna habitat.Leopard populations were found to be associated with lions and hyaenas but they avoidedhuman disturbances. Regarding potential breeding dispersal, the Gaussian representationshowed a clear fragmentation among populations, suggesting that long-term survival of thespecies could be threatened. We found no area to be exempt from threats. The leopard has,however, declined less than other carnivore species and still shows viable populations.Furthermore, occurrences were found to be significantly more numerous than expected inprotected areas, suggesting the relative efficiency of conservation.

Key words: conservation, dispersal distance, distribution, feline status, Panthera pardus,population fragmentation, threat, West-Central Africa.

INTRODUCTIONMost endangered species, and especially largefelids, have suffered serious declines in theirpopulations during recent decades. Hence theefficiency of the protection measures imple-mented can be questioned.

Knowledge of wildlife status is variable inWest-Central Africa. Some species, such as lions(Panthera leo), elephants (Loxodonta africana), andCercopithecus monkey species (e.g. Cercopithecussclateri), have been widely studied (e.g. Tchamba1996; Bauer 2003; Kümpel et al. 2008) whereasscientific literature is extremely poor concerningother species such as leopards (Panthera pardus),giraffes (Giraffa camelopardalis) or striped hyaenas(Hyaena hyaena) (Nowell & Jackson 1996, Henschel2008; IUCN/PAPACO 2009). Despite various pro-tection measures and a number of protected areas,some studies have shown that many species, includ-ing lions, wild dogs (Lycaon pictus), Derby elands(Taurotragus derbianus), and cheetahs (Acinonyxjubatus), are still declining (Woodroffe & Ginsberg1999; Bauer et al. 2003; Angwafo 2006; Haywardet al. 2007). Other species, such as the black rhinoc-eros (Diceros bicornis), have even been declaredextinct (Prouteau 2007). More data on species areneeded to set appropriate management guide-lines and to improve the existing ones.

Leopards may be the felids with the largestrange in the world, occupying almost all ofsub-Saharan Africa, and a great part of Asia(Nowell & Jackson 1996). Thus the species occupiesvarious habitats, from tropical forests to woodlandsavannas, and shows great adaptability to theenvironment (Nowell & Jackson 1996). Althoughsome researchers consider leopards as generalistfeeders (Hart et al. 1996; Ott et al. 2007), othersshow that they exhibit prey-size preferences (e.g.Hayward et al. 2006; Schwartz & Fischer 2006;Martins et al. 2011). Their hunting behaviour alsovaries according to the habitat: leopards arelargely nocturnal in an open environment (e.g.Stander et al. 1997; Bailey 2005), whereas theyshow diurnal and crepuscular habits in forestecosystems (Hart et al. 1996; Jenny & Zuberbuhler2005). Home-range size depends on the preferredprey availability in the habitat (Bothma et al. 1997;Steyn & Funston 2009). Although the specificconservation status is difficult to establish in thisvery elusive species (Norton 1990; Henschel 2008)it is commonly agreed that leopard populationsremain widely distributed.

In West-Central Africa, the distribution of thespecies is supposed to be semi-continuous fromthe south of Senegal to Burkina Faso and Benin,although the leopard could have a more discontin-uous range in Niger, Nigeria and Cameroon*Author for correspondence. E-mail: [email protected]

African Zoology 48(2): 374–387 (October 2013)

(IUCN 2010). However, the most recent analysis ofits status (Martin & Meulanaer 1988) remains verycontroversial (Norton 1990; Henschel 2008). Despitethe apparent consensus on the conservation status,the efficiency of protection measures has rarelybeen estimated (Rodrigues et al. 2004; Al-Johany2007; Visser et al. 2009).

Carnivores are usually found in quite low densi-ties, even in optimal conditions. They are sensitiveto every modification of their environment, partic-ularly those due to human activities that lead tohabitat deterioration and prey depletion. Forinstance, aquatic carnivores have to cope with thehuman alteration of freshwater ecosystems (Lodé2002; Bifolchi & Lodé 2005), while forest carni-vores have to face the influence of deforestationdue to forest exploitation and land reclamation(Hacker et al. 1998; Niedzialkowska et al. 2006).Habitat deterioration, which includes both habitatfragmentation and habitat loss, is fundamentallyrelated to species fragmentation, a critical threatfor endangered species (Fahrig 1997). Habitatpatches are generally interspersed in damagedenvironments, thus leading to a heterogeneousfragmentation within wildlife populations. If thedistance between continuous habitat patches istoo large, their connectivity becomes too low toallow a sufficient dispersal of individuals (Fahrig& Merriam 1994), altering both breeding exchangesand genetic diversity (Frankham 1995). As it maybe more difficult for individuals to mate (Comis-key et al. 2002; Rodriguez & Delibes 2003), frag-mentation might ultimately cause a decline inpopulations, and could lead to local extinctions,reinforcing the threatened status of those endan-gered species (Fahrig & Merriam 1994; Benderet al. 1998).

As a top predator, the leopard might be stronglyaffected by habitat degradation. In West-CentralAfrica, fragmentation of leopard populations hasbeen rarely documented (Henschel 2008), andlittle information is available concerning leoparddensities (Croes et al. 2011). However, the conser-vation of this emblematic species has important eco-nomic consequences (Verbelen 1999; Nemangaya2002) because wildlife represents a great source ofincome through tourism and trophy hunting(Kwabong 2008; Hemson et al. 2009; Lindsey et al.2012).

Recently, some attention has also been given tointeractions among predators (Creel et al. 2001;Odden et al. 2010). Leopards are sympatric withnumerous other predators in open habitats, and

such direct or exploitative competition could reducethe access to distinct resources. Nevertheless, it iscommonly admitted that where large carnivoresoccur sympatrically, their chances of persistenceare higher. The reason for this might be the greaterefficiency of multispecies conservation actions(Ray et al. 2005; Hess et al. 2006; Lindenmeyer et al.2006).

From a detailed investigation, the objective ofthis study was to analyze the ecological factors andmanaging practices related to leopard conserva-tion status in Cameroon. We analyzed the leoparddistribution and explored to what extent it wasaffected by environmental factors, interactionswith other species, and other threats such aspoaching, agriculture and forest logging. Thisstudy also provide an initial basis of leopard statusin Cameroon. Our study uses an original approach,combining a mathematical model of dispersalwith an evaluation of environmental deteriora-tion. This research should help highlight conser-vation priorities and provide advice for manage-ment of carnivores in West-Central Africa.

METHODS

Study areaCameroon has an area of 475 442 km². Human

population (18 894 406 inhabitants with a den-sity of 41 inhabitants/km²) is mainly concentratedin the cities (57% of the total population), while therest of the country is inhabited by local villagepopulations.

Climate varies from dry-tropical in the north, totropical-humid/equatorial in the south, with localvariations. Cameroon’s vegetation is of five maintypes: humid forest with either monomodal orbimodal rainfall, highland vegetation, Guineansavanna (deciduous woodland), and Sudano-Sahelian savanna (deciduous shrubland) (IRAD2009).

Leopard distributionWe documented the presence of leopard through-

out the Cameroon by field work, questionnaires,and bibliographical research. We recorded varioustraces that indicated the occurrence of leopards(faeces, pugmarks, carcasses and sightings) in theNational Park of Boubandjida (North province)from March to May 2010. We walked in dryriverbeds six days a week, the two first weeks ofeach month, looking for signs, from 6:00 to 10:00and from 16:00 to 18:00. While in the field, P.T. met

Toni & Lodé: Conservation of leopards in West-Central Africa 375

the hunting guides of 23 hunting zones in theBénoué National Park Complex (North province).During informal meetings they said they had seenleopards within their areas in the past year. Inorder to scientifically record these testimonies, weemailed questionnaires to them in May 2010. Thequestionnaires consisted of descriptive questionsabout the hunting zone (e.g. geographical infor-mation, wildlife species present, encounteredthreats), as well as the presence of leopard throughdetailed observation records. Only nine huntingguides answered. Data were also collected through32 published or unpublished papers or fieldworkreports (Table 1). We selected bibliographicaldata according to four criteria: 1) data from theses,unpublished data from extensive field research, orreports supervised by an international and inde-pendent organization (e.g. WWF, WCS), 2) thefieldwork was conducted in Cameroon after 1995,3) the results clearly show that leopard presencehas or has not been evidenced in the area for at leasta decade and 4) data are original. The reliability ofdata was assessed by personally discussing withthe authors their methods and results, and whenpossible, asking people we know in the area if theyhad observed leopards recently. From these andour field data collection we recorded the evidenceindicating presence or absence of leopard in thestudied areas.

To ensure the coherence of data we defined anindex calculated from the number of tracks ofleopards found in 10 areas of similar conditionsreported to a given surface. These data wereextracted from studies mentioning density esti-mates in their results section. We obtained aregression index of 105.03 (Fig. 1), and then definedour index as equal to one unit of presence orabsence per 100 km². The value of our index in onearea was thus equal to its size (in km²) divided by100. This seemed pertinent as the median homerange of leopard, calculated in previous studies(Jenny 1996; Stander et al. 1997; Grassman 1999;Marker & Dickman 2005; Balme et al. 2007;Henschel 2008; Simsharoen et al. 2008; Swanepoel2008), is equal to 102 km². The use of this indeximplies that leopards are uniformly distributedwithin the delimitated area where evidence oftheir presence has been recorded. We applied thisindex to the areas previously mentioned (Table 1)and obtained numbers of data points (or occur-rence) of presence or absence of leopard (n = 661).

In order to model potential breeding dispersal,leopard distribution was treated through a

Gaussian representation (see Lodé 2002). Themapping was smoothed out through a Gaussianblur of the conventional form

f j k

J k

( , )

(

= ⎛⎝⎜ ⎞

⎠⎟

− +⎛

⎝⎜⎜

⎞

⎠⎟⎟1

2

2 2

2 2

ps

s ,

where (j,k) represents the intensity at position (j,k)(Press et al.1992). Because home range size andbreeding dispersal occurred over about 100 km,the Gaussian blur illustrates the potential dispersalwithin a radius of 100 km around every occupiedsite. This representation does not illustrate thepopulation density, but it does illustrate thepatchy distribution and connectivity. When nu-merous population exchanges are possible, therange is represented with a darker shade.

Factors potentially related to leopard presenceWe defined five types of habitats, according to

the five main types of vegetation: humid forestwith monomodal rainfall in the western provinces,highland vegetation in the northwestern prov-inces, humid forest with bimodal rainfall fromthe south to the south of Adamawa province,Guinean savanna (with deciduous woodland) inthe Adamawa province, and Sudano-Saheliansavanna (with deciduous shrubland) in the north-ern provinces (Fig. 2).

We used three categories for human densities:less than 10 inhabitants per km² (meaning ruralareas), between 10 and 100 (rural areas with highlevel of exploitation), more than 100 (highly popu-lated areas).

This study focused on two types of interactions:competition and predation. In open areas, fourpredator species were identified as potential com-petitors: lions, hyaenas, wild dogs, and jackals,while no competitors occur in forest areas. Com-petitive interactions between leopards and otherlarge carnivores were tested only in open areas(i.e. highland vegetation, Guinean savanna andSudano-Sahelian savanna). Potential prey specieswere divided into four categories (Hayward et al.2006): small mammals (body mass <20 kg, e.g.oribi, duikers (Stuart & Stuart 2000)), medium-sized ungulates (20 kg< body mass <70 kg, e.g.bushbuck, reedbuck, kob, common warthog, andred river hog), large ungulates (>70 kg body mass,e.g. hartebeest, topi, waterbuck, sitatunga, giantforest hog) and monkeys and apes (baboon, chim-panzee).

We explored protection level through the presenceof potential threats and protection status of the

376 African Zoology Vol. 48, No. 2, October 2013


Tab

le1.

Rep

orts

,pap

ers

and

unpu

blis

hed

data

used

toas

sess

leop

ard

pres

ence

acro

ssC

amer

oon

(LC

S=

larg

eca

rniv

ore

surv

ey,L

MS

=la

rge

mam

mal

surv

ey,H

AS

=hu

-m

anac

tiviti

essu

rvey

,MP

=m

anag

emen

tpla

n;N

P=

Nat

iona

lPar

k,S

=S

anct

uary

,BR

=B

iosp

here

Res

erve

,FR

=F

auna

lRes

erve

,UF

A=

For

estr

yA

ppoi

ntm

entU

nit,

ZIC

=Z

one

ofC

yneg

etic

Inte

rest

;CT

=ca

mer

atr

appi

ng,S

C=

spoo

rcou

nt,L

T=

line

tran

sect

,RT

=re

cce

tran

sect

,Itw

=in

terv

iew

;ER

=en

coun

terr

ate,

DE

=de

nsity

estim

ates

,N

O=

num

ber

ofob

serv

atio

ns,P

res.

=pr

esen

ce,A

=an

imal

,T=

trac

e,P

=pi

ctur

e).

Sour

ceTy

peof

docu

men

tOb

ject

ives

Area

Tem

pora

lfra

me

Met

hods

Data

prov

ided

Leop

ard

Abou

dou

etal

.20

08M

.Sc.

thes

isLC

SBo

uban

djid

aNP

&ZI

C11

300j

in20

07CT

with

baits

+SC

ER(T

1P)

Yes

Angw

afo

2006

Ph.D

.the

sis

LMS

Béno

uéNP

&Fa

roNP

Aug

04–M

ay05

Itw+

LTER

(T)

Yes

deJa

ger2

007

M.S

c.th

esis

LCS

Béno

uéNP

Jan–

Jun

07CT

with

baits

+SC

ER(T

1P)

Yes

Asta

ras

(unp

ub.d

ata)

Ph.D

.res

earc

hLM

SKo

rup

NPFe

b–Ju

n06

Itw+

RTPr

es.

NoM

inEF

2007

NGO

repo

rtM

PKo

mNP

&M

enga

me

SNo

info

rmat

ion

LSPr

es.

NoM

inEF

1997

NGO

repo

rtM

PW

asa

NPNo

info

rmat

ion

LSPr

es.

NoNz

ooh-

Dong

mo

2003

NGO

repo

rtLM

&HA

SLo

béké

NPJa

n99

–May

03RT

+Po

nctu

alob

sDE

/ER

Yes

Ngau

djui

etal

.200

2Re

port

MP

Cam

po-M

a'an

NPJu

n00

–May

01RT

DE/E

RNo

Ngau

djui

etal

.20

03Re

port

MP

UFAs

21,2

4,25

Jun

00–M

ay02

RTDE

/ER

NoNc

hanj

i200

2Un

publ

.dat

aLM

&HA

SBa

nyan

g-M

boS

Noin

form

atio

nRT

DE/E

RNo

Halfo

rdet

al.2

002

NGO

repo

rtLM

&HA

SDj

aBR

Aug–

Sept

.02

Itw+

LTDE

/ER

(A&

T)No

Sund

erla

nd-G

rove

s&

Mai

sels

2003

Scie

ntifi

cpa

per

LMS

Taka

man

daFo

rest

Rese

rve

Noin

form

atio

nLT

ER(A

&T)

NoLu

cyet

al.20

03NG

Ore

port

LMS

Bom

boko

Fore

stRe

serv

eAu

g02

–Mar

03RT

NO(A

&T)

NoEn

o-Nk

u20

03NG

Ore

port

LMS

Ejag

ham

Fore

stNo

info

mat

ion

RTER

(A&

T)No

Ekob

o20

02NG

Ore

port

LMS

Mon

tCam

erou

nFo

rest

Oct–

Nov

03RT

ER(A

&T)

NoEk

inde

&W

ittin

ger2

005

NGO

repo

rtLM

SBa

ntak

pa&

Mbu

luFo

rest

2002

–200

5RT

ER(T

)No

Fom

ete

Nem

bot&

Tcha

nou

1999

NGO

repo

rtM

PAy

os,B

akos

si,D

oual

a-Ed

éaNo

info

rmat

ion

LSPr

es.

NoFo

met

eNe

mbo

t&Tc

hano

u19

99NG

Ore

port

MP

Koup

e,Lo

koun

dje

Nyon

gNo

info

rmat

ion

LSPr

es.

NoFo

met

eNe

mbo

t&Tc

hano

u19

99NG

Ore

port

MP

Man

engo

uba

Mw

ane

Noin

form

atio

nLS

Pres

.No

Fom

ete

Nem

bot&

Tcha

nou

1999

NGO

repo

rtM

PM

bam

&Dj

erem

,Nlo

nako

Noin

form

atio

nLS

Pres

.No

Fom

ete

Nem

bot&

Tcha

nou

1999

NGO

repo

rtM

PNt

a-Al

i,ok

u,Ri

ode

iRey

Noin

form

atio

nLS

Pres

.No

Fom

ete

Nem

bot&

Tcha

nou

1999

NGO

repo

rtM

PRu

mpi

,Tch

abal

Bado

,Yao

undé

Noin

form

atio

nLS

Pres

.No

Ekob

o19

98NG

Ore

port

LMS

Ngoi

la,N

kiNd

ongo

-Adj

ala

Noin

form

atio

nRT

ER(A

&T)

Yes

Ekob

o19

98NG

Ore

port

LMS

Boum

ba-B

ek&

NkiN

PNo

info

rmat

ion

RTER

(A&

T)Ye

sEk

obo

1998

NGO

repo

rtLM

SM

olou

ndou

-Mim

bom

imbo

Noin

form

atio

nRT

ER(A

&T)

Yes

Forb

oseh

etal

.20

07Sc

ient

ific

pape

rLM

SEt

inde

,Nkw

ende

Hills

,Mon

e20

02–2

004

RT+

LSER

(T)

NoNd

eh20

04NG

Ore

port

LMS

UFA

09-0

13No

info

rmat

ion

RTER

(A&

T)Ye

sEn

o-Nk

u20

04NG

Ore

port

LMS

UFA

09-0

21Ju

n00

–May

01RT

ER(A

&T)

Yes

Eno-

Nku

2004

NGO

repo

rtLM

SUF

A09

-024

Jun

00–M

ay01

RTER

(A&

T)No

Nzoo

h-Do

ngm

oet

al.20

04NG

Ore

port

LM&

HAS

UFA

10-0

13Oc

t–De

c03

RTDE

/ER

NoC

ontin

ued

onp.

378

different areas. We determined eight types ofthreats: poaching/disturbance due to game hunt-ing, agriculture, pasture, gathering/use of forestproducts (fruits, roots, fire wood, etc.), forest logging(exclusively in forest habitats), human–wildlifeconflicts, areas crossed by road infrastructures,and other threats. There are 24 protected areas(excluding Hunting Zones, Forest Reserves) in thecountry representing 3 755 602 ha (7.9% of thecountry’s area), 15 National Parks and nine FaunalReserves and Sanctuaries (Table 2) (RAPAC 2010;MinFOF & MinEP 2010).

We determined five types of protection statuses:National Parks, Reserves (including Faunal Re-serves, Sanctuaries, and Forest Reserves), Zones ofCynegetic Interest (ZIC), Exploited Forests (EF)gathering Units of Forestry Appointments (UFA)and Communal Forests (CF) and Other (mangroves,mountains, etc.). Leopards are integrally pro-tected in the whole country, and in National Parks,Reserves and ZIC, this protection is applied (hunt-ing restriction or interdiction, anti-poachingpatrols), whereas Sanctuaries concentrate on pro-tecting one species (gorilla in Mengame, chimpan-zee in Banyang Mbo). In Exploited Forests andother areas, no special measures are applied.

We assessed the relationship between factorsand leopard presence using chi-square (withYates’ correction) tests.

RESULTS

Environmental factors

Out of 661 data points, we evidenced 416leopard occurrences. Leopard distribution, as afunction of habitat type, was significant (c² (Yates’correction), npresence = 416, nabsence = 245, d.f. = 4, P< 0.001) (Fig. 3). Leopards were found to be morenumerous than expected in the Sudano-Saheliansavanna habitat (31.7%) and the humid forest withbimodal rainfall habitat (55.2%). However, noleopard occurrences have been reported in west-ern highland vegetation (0%) and few in humidforest with monomodal rainfall (8.8%) habitats.

Populations were subdivided into several demes(Fig. 2). There are two large populations in thenorth and in the south while three small popula-tions in the central part of the country were veryisolated. Regarding potential breeding dispersal,the Gaussian representation showed a clear frag-mentation among populations and leopard distri-bution greatly differed among areas. The southernpopulation exhibited some zones of fragility.


Tab

le1

(con

tinue

d)

Type

ofdo

cum

ent

Obje

ctiv

esAr

eaTe

mpo

ralf

ram

eM

etho

dsDa

tapr

ovid

edLe

opar

d

Nzoo

h-Do

ngm

oet

al.20

03NG

Ore

port

LM&

HAS

UFA

10-0

18No

v–De

c02

&M

ay03

RTDE

/ER

Yes

JMN

Cons

ulta

nts

2006

Repo

rtIm

pact

stud

yUF

A10

-030

,039

,041

,042

,044

Noin

form

atio

nRT

ER(T

)No

Eno-

Nku

2004

NGO

repo

rtLM

SUF

A11

-002

Noin

form

atio

nRT

ER(A

&T)

NoM

endo

mo-

Bian

g&

Nzoo

h-Do

ngm

o20

07NG

Ore

port

LM&

HAS

ZIC

1&

8M

ay–A

ug07

RTER

(A&

T)No

Ghog

ue20

07NG

Ore

port

LM&

HAS

ZIC

2Ap

r–Ju

n07

RTER

(A&

T)No

LFVe

ko20

07Re

port

LMS

ZIC

13Fe

b–M

ar07

LTDE

/ER

Yes

Boul

etet

al.20

08NG

Ore

port

LMS

ZIC

18,1

8B,1

5,03

,24

2005

–200

8Te

stim

onie

sNO

(A&

T)Ye

sM

ahop

2007

NGO

repo

rtLM

SZI

C19

Apr–

July

07RT

ER(A

&T)

Yes

LFVe

ko20

07Re

port

LMS

ZIC

31Ju

n07

LTDE

/ER

Yes

LFVe

ko20

07Re

port

LMS

ZIC

38Ju

n07

LTDE

/ER

Yes

Nzoo

h-Do

ngm

oet

al.20

02NG

Ore

port

LMS

ZIC

GC2,

3,8

Mar

01–A

pr02

RTER

(A&

T)No

Nzoo

h-Do

ngm

oet

al.20

02NG

Ore

port

LMS

ZIC

GC1,

9M

ar01

–Apr

02RT

ER(A

&T)

Yes


Fig. 2. Distribution and fragmentation (Gaussian representation) of leopard populations in Cameroon. Numbers referto different habitats: 1, Sudano-Sahelian savanna; 2, Guinean savanna; 3, western highland vegetation; 4, humidforest with monomodal rainfall; 5, humid forest with bimodal rainfall

Fig.1.Regression between size (km2) of study area and number of leopard records (data obtained from 10 areas).


In 88.2% of occurrences, evidences of leopardswere found in places with very low human density.Leopard occurred significantly less frequentlywhere human density was above 10 inhabit-ants/km² (c², npresence = 416, nabsence = 245, d.f. = 2,P < 0.001) (Fig. 4): 11.8% of occurrences werefound in areas of medium densities and no occur-rence in densely populated areas (>100 inhabi-tants/km²).

Interactions with other species

Occurrences of leopards were not significantlyrelated to prey species presence (Fig. 5, c², npresence

= 416, nabsence = 245, d.f. = 4, n.s.). By contrast,there were significantly more occurrences of leopardpresence than leopard absence where other pred-ators occur (savanna habitats) (c², npresence = 150,nabsence = 36, d.f. = 4, P < 0.001). Lions and hyaenaswere the species most frequently associated withleopard occurrence (43% of leopard occurrenceswere associated with lion presence and 39% tohyaena presence, Fig. 6).

Protection level

Leopard distribution was significantly related to

protection statuses (c², npresence = 416, nabsence = 245,d.f. = 4, P < 0.001) (Fig. 7). Leopards are presentmore in National Parks and ZIC than in reserves,Exploited Forests, and Other areas. Actually 49%of the evidence of the occurrence of leopards wasfound in National Parks, 29% in ZIC, 15% inReserves and 7% in Exploited Forests.

Numerous threats were recorded within leopardranges, with poaching/disturbances linked tohunting (92% of considered areas) and logging(89% of considered areas) being nearly omnipresent.The number of occurrence of leopard presencewas significantly related to the presence of thedifferent types of threats (c², npresence = 416, nabsence

= 245, d.f. = 7, P < 0.001, Fig. 8).

DISCUSSIONLeopards were evidenced in West-Central Africa,but the species exhibits fragmented populations,revealing important conservation challenges.

Habitat factors

Leopards are regarded as generalists in terms ofhabitat (Nowell & Jackson 1996; Bailey 2005).Accordingly, we found that leopards were widely

Table 2. Protected Areas in Cameroon (NP = National Park, BR = Reserve of Biosphere, FR = Faunal Reserve, S =Sanctuary).

Name Status Surface (ha) ‘Province’

Kalamaloué NP 4 500 extreme NorthMozoko-Gokoro NP 1 400 extreme NorthWaza NP 170 000 extreme NorthBoubandjida NP 220 000 NorthBénoué NP 180 000 NorthFaro NP 330 000 NorthMbéré NP 77 760 AdamawaKorup NP 125 900 SouthwestMbam et Djerem NP 125 000 CentreMpem et Jim NP 100 000 CentreCampo-Ma’an NP 264 064 SouthLobéké NP 217 854 EastBoumba-Bek NP 238 225 EastNki NP 309 362 EastMefou NP 1 044 CentreDja BR 526 000 EastMbi Crater FR 370 NorthwestKimbi FR 5 625 NorthwestDouala-Edéa FR 160 000 LittoralLac Ossa FR 4 000 LittoralSantchou FR 7 000 WestMonts Bakossi FR 29 300 SouthwestBanyang Mbo S 66 000 SouthwestMengame S 96 010 South

spread in savanna areas and in rainforest habitats.However, they seem to be absent from Cameroo-nian rainforests; forests are widely exploited inCameroon (Halford et al. 2002), causing habitatloss for leopards and for their prey populations(Verbelen 1999). Moreover, industry and trade arehighly developed, in particular around Douala,which is the biggest city in the country and has alarge port. Additionally, forest habitats are in-creasingly used for agriculture (Halford et al. 2002),which causes the active presence of humans. Thehigh human density increases the pressure on theenvironment and often leads to high levels ofbushmeat trade (Fa et al. 2006). Here we found fewoccurrences of leopards in areas with a densityhigher than 10 inhabitants per km², a densitymuch lower than the density of 900 inhabitants/km² predicted as critical by Woodroffe (2000), or

the density of 80 inhabitants/km² characterizingareas where leopards have become extinct (Fa et al.2006). However, Henschel (2008) found no leop-ards in one of his study sites where human densitywas only 1–2 inhabitants/km². Avoidance of evenbarely populated areas might be due to threatspresented by humans (hunting/poaching of leop-ards and/or prey species) more than to humandensity itself (Linnell et al. 2001; Halford et al.2002).

Spacing patterns are mainly governed by thebreeding system and by the general pattern ofdispersal. Nevertheless, patterns of dispersal inbreeding felids are expected to proceed accordingto a continuous or quasi-continuous distributionin which sub-population exchanges are favouredin contiguous zones (Rodriguez & Delibes 2003).As a result, the quality of contiguous habitats


Fig. 3. Presence/absence of leopards according to habitat type.

Fig. 4. Presence/absence of leopards according to human density.

has proved to be decisive for populations andsubdivision may directly affect the breeding andviability of small isolated populations because ofthe low number of breeding adults (Rodriguez &Delibes 2003). The ‘Allee effect’ hypothesis (Alleeet al. 1950) predicts that poor or deteriorated habi-tats should result in home-ranges that are so ex-tensively scattered that low densities preventmost females from finding mates. In the currentstudy leopard populations showed a very patchydistribution, avoiding numerous areas so thatbreeding dispersal already seems to be highlyaffected. Thus, in West-Central Africa, leopardsexhibited widely spaced territories, disturbingthe social system. Many leopard adults mayfail to breed, which supports the Allee effecthypothesis.

Associated species

The leopard has been assumed to be an opportu-nistic predator, showing a generalist diet (Hartet al. 1996), although small to medium-sized preyrepresent the best compromise between energycosts and benefits (Sunquist & Sunquist 2002; AlJohany 2007), and have been identified as preferredprey in several studies (Henschel et al. 2005; Hay-ward et al. 2006; Schwartz & Fischer 2006). Leopardstend to be found less where there are monkeysand apes. This may suggest that the presence ofthose species is not a shaping factor of leoparddistribution (Hayward et al. 2006).

In areas where sympatric predators occur, leop-ards are present more where other large carnivorespecies are found. Similarly, across Africa, popula-tions of leopards are chiefly recorded where they


Fig. 5. Occurrence of leopards according to prey species (small mammals (body mass <20 kg, e.g. oribi, duikers(Stuart & Stuart 2000), medium-sized ungulates (20 kg < body mass < 70 kg, e.g. bushbuck, reedbuck, kob, commonwarthog, red river hog), large ungulates (>70 kg body mass, e.g. hartebeest, topi, waterbuck, sitatunga, giant foresthog) and monkeys and apes (baboon, chimpanzee)).

Fig. 6. Presence/absence of leopards according to associated carnivore species.

occur sympatrically with other large carnivores,especially lions and hyaenas (Nowell & Jackson1996). It may be that competition is reducedbecause the leopard forages individually. As aresult, dietary overlap between leopards andother large carnivores is not that important (Mills& Biggs 1993; Hayward & Kerley 2008). Conserva-tion efforts are therefore usually more effectivewhen addressed to an ecosystem than to a singlespecies (Ray et al. 2005; Hess et al. 2006; Linden-meyer et al. 2006). Thus, management strategieshave to be improved by considering species assem-blage and not only the requirements of one species.

Protection level

One of the major points of our study is that wedid not find any areas where leopards are notthreatened. The decline of large felids is commonlyattributed to habitat degradation, and huntingand poaching of both predators and their prey(Bauer et al. 2003; Marker & Dickman 2005; AlJohany 2007). Forestry logging had a negativeimpact on leopard distribution, although we werenot able to quantify the pressure this threat repre-sented within each place. The influence of defor-estation is not only due to its extent but also to itspattern (Rodriguez & Delibes 2003; Bender et al.


Fig. 8. Occurrence of leopards according to different types of threats or the absence of threats.

Fig. 7. Presence/absence of leopards according the conservation status of an area. (NP = National Park, R =Reserves (Faunal Reserve/Sanctuary/Forest Reserve), EF = Exploited Forest (UFA = Forestry Appointment Unit/CF= Communal Forest), ZIC = Cynegetic Interest Zone, Other = mangroves, mountains, etc.).

1998), thus affecting population fragmentation.Domestic species may become a substitute for

wild prey (Ogada et al. 2003; Kolowski & Holekamp2006; Holmern et al. 2007; Kissui 2008; Gusset et al.2009). Livestock owners may then remove leopardsas retaliation (Dickman 2005; Holmern et al. 2007;Kissui 2008; Chapman & Balme 2010). Such retalia-tory killing, added to widespread agriculture, maydirectly threaten leopard populations (Camp 2002;Ott et al. 2007). However, we found the leopardspresent more often than expected in areas wherethere were human–wildlife conflicts. This could bebecause these habitats still had numerous sheltersfor leopards or because we labelled conflicts asevery kind of human–wildlife interaction, withoutspecifying those involving leopards.

Nonetheless, hunting and poaching were com-mon in the great majority of the areas covered bythis study. Unexpectedly we found more evidenceof leopard presence in areas where people huntand/or poach. It could be hypothesized that bothhunters and leopards choose areas where herbi-vores are abundant (Marker & Dickman 2005; AlJohany 2007; Hayward et al. 2009; Martins et al.2011). Moreover, these areas are not highly popu-lated, and Fa et al. (2006) and Henschel (2008)showed that human density and proximity resultedin higher pressure on wildlife and lower ungulatedensities. Thus, leopards would prefer habitatswith low human densities, regardless of both thethreat and competition posed by hunters (Jorgenson& Redford 1993; Henschel 2008). Nevertheless,hunting and poaching were present in more than90% of the sampled areas. This means there arefew places where leopards can occur but poachingand hunting do not. Fortunately, as killing leopardsis forbidden, this species is probably little threat-ened by direct hunting, and as a discrete animal,leopards require too much effort to be poached.Finally, leopards are sacred animals in mostCameroonian ethnic groups (Wilcox & Nambu2007; Astaras, pers. comm.). Therefore, leopardsare more likely to be threatened by habitat degra-dation and prey depletion, than directly by poach-ing (Karanth & Stith 1999). As a result, leoparddistribution is determined as much by anthropo-logical factors (human density) as by ecologicalones (prey densities, habitat type) (Marker &Dickman 2005; Al Johany 2007; Hayward et al.2009).

Unsurprisingly, we found that leopard distribu-tion is highly related to the protection status of thestudied area. The species is widely spread in

National Parks, where the protection level is thehighest (Bauer 2003; MinFOF 2005): human pres-ence is strictly limited, habitations are forbiddeninside the area, as are any kind of anthropogenicmodifications (cutting trees, pasturing etc.,) andcollections (of dead wood, carcasses etc.). There-fore, it seems that measures of protection are rela-tively efficient, when willingly applied. Leopardpresence is also important in ZIC. These ZICs aremostly close to National Parks, additionally, hunt-ing is scrupulously regulated since quotas aredefined and hunters must have a license (MinFOF2005), thus reducing human presence. By contrast,there is little evidence of leopard presence infaunal reserves, despite the fact that they areunder strict protection rules (MinFOF 2005).Generally, the explored reserves were smallerthan national parks. We could hypothesize thatthere is less wildlife in such areas, where conserva-tion measures are less strict than in National Parks(Nowell & Jackson 1996). As home range size iscorrelated to prey abundance in the leopard, theyshould require larger home ranges in such areas,resulting in lower numbers (Rodrigues et al. 2004;Marker & Dickman 2005; Ray et al. 2005). On thewhole, evidence of leopards is rare outside pro-tected areas, and this would lead to populationfragmentation.

It seems that leopards persist in small isolatedpopulations, but are threatened by extinction.Moreover, within the southern population wefound several zones of fragility, which suggeststhat breeding exchanges have been affected.

Our study established a baseline overview ofavailable data on the leopard in establishing theconservation status of the feline. The decline ofleopards in Cameroon is less alarming than otherspecies, especially other big carnivores. Its widepresence in protected areas suggests that effortsmade in these zones are efficient but should beimproved because of the fragmentation of leopardpopulations.

We also highlighted the paucity of data on thespecies in Cameroon. There is an urgent need forfurther studies to propose management guide-lines to improve conservation efforts based onaccurate data. For instance, detailed faunal inven-tories should be carried out regularly in order topromote better management of animals insideand outside protected areas. Additionally, huntingpolicies should be modified in order to ensurebetter management of faunal resources, while tak-ing into account local realities, and efforts to



strengthen the anti-poaching cause shouldcontinue and be strengthened. The importance ofthis iconic animal in local beliefs, combined withits economic potential through its value for tourism,could be a starting point for future conservationefforts.

ACKNOWLEDGEMENTS

We would like to express our gratitude to PaulBour, without whom this study would not havebeen possible and to Maïmounatou Bilkissou forher help. We would also like to thank all the peoplewho helped us in this study including theConservateur André NDjidda, the anti-poachingguards, the hunting guides (Pável Redondo Marti-nez, David Fine, Charles Dogas de la Boissonny,Vincent Citerne), and all the people who providedus with information we needed (Barbara Croes,Beskreo Waga). Thanks are due to two anonymousreferees. Finally, special thanks to those who gaveus financial and affective support.

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Responsible Editor: J.H. van Wyk


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