Small carnivorans from southern Benin: A preliminary assessment of diversity and hunting pressure

Abstract

We conducted a preliminary study totalling nine-weeks between August 2007 and June 2008 in southern Benin to assess small car-nivoran diversity and the hunting pressure to which they are subject through interviews, surveys of local markets and occasional direct observations. We provide an 'Index of Rarity' (IR), expressed as the number of times a species is identified as 'rare' by interviewees / the number of times it is mentioned. Nine species or taxa (Genetta spp.) were identified through 86 interviewed hunters, representing a total 333 mentions. Genets, Cusimanse Crossarchus obscurus and Ichneumon Mongoose Herpestes ichneumon were by far the most sighted, the prevalence of such ecologically versatile species confirming that southern Benin constitutes an environmentally disturbed region. Other species were Spotted-necked Otter Hydrictis maculicollis, Marsh Mongoose Atilax paludinosus, White-tailed Mongoose Ichneumia albicauda, African Civet Civettictis civetta, Gambian Mongoose Mungos gambianus and African Palm-civet Nandinia bi-notata. Direct observations allowed us to confirm the presence of G. pardina / G. maculata, C. obscurus, H. ichneumon, and H. macu-licollis. Through market surveys and subsequent molecular identifications, we clearly distinguished among G. genetta, G. pardina / G. maculata and G. thierryi, raising to 11 the number of small carnivoran species present (or probably present) in southern Benin. During our interviews, Slender Mongoose Galerella sanguinea and African Small-clawed Otter Aonyx capensis were never positively identi-fied. The ubiquitous C. civetta was considered the rarest species (IR = 0.89), followed by H. maculicollis (0.72) and I. albicauda (0.69), whereas C. obscurus (0.01) was the commonest small carnivoran. Hunting techniques were mostly traditional guns, accompanied by dogs, and jaw traps. Despite the absence of selective hunting, small carnivorans are likely to represent a fair source of income for hunters, body parts being sold to fetish markets in 47% of the cases. Mean incomes range between US$ 2.5 and 5.4 per animal, with the notable exception of C. civetta (US$ = 14.6) and heads of H. maculicollis, reaching US$ 33.7. The fair proportion of small carnivorans observed on fetish market displays showed that hunting for animist practices might sustain a continuous hunting pressure in Benin. Our preliminary survey raises a number of questions as to the distribution of small carnivorans in southern Benin, the impact of heavily dis-turbed habitats on their survival and the level of sustainability of the hunting pressure they are subject to. Additional field surveys will be necessary for more precise characterisation of their status.

Full text

1
Small carnivorans from southern Benin: a preliminary assessment of
diversity and hunting pressure
Chabi Adéyèmi Marc Sylvestre DJAGOUN1 and Philippe GAUBERT2*
Abstract
We conducted a preliminary study totalling nine-weeks between August 2007 and June 2008 in southern Benin to assess small car-
nivoran diversity and the hunting pressure to which they are subject through interviews, surveys of local markets and occasional direct
observations. We provide an ‘Index of Rarity’ (IR), expressed as the number of times a species is identied as ‘rare’ by interviewees
/ the number of times it is mentioned. Nine species or taxa (Genetta spp.) were identied through 86 interviewed hunters, representing
a total 333 mentions. Genets, Cusimanse Crossarchus obscurus and Ichneumon Mongoose Herpestes ichneumon were by far the most
sighted, the prevalence of such ecologically versatile species conrming that southern Benin constitutes an environmentally disturbed
region. Other species were Spotted-necked Otter Hydrictis maculicollis, Marsh Mongoose Atilax paludinosus, White-tailed Mongoose
Ichneumia albicauda, African Civet Civettictis civetta, Gambian Mongoose Mungos gambianus and African Palm-civet Nandinia bi-
notata. Direct observations allowed us to conrm the presence of G. pardina / G. maculata, C. obscurus, H. ichneumon, and H. macu-
licollis. Through market surveys and subsequent molecular identications, we clearly distinguished among G. genetta, G. pardina / G.
maculata and G. thierryi, raising to 11 the number of small carnivoran species present (or probably present) in southern Benin. During
our interviews, Slender Mongoose Galerella sanguinea and African Small-clawed Otter Aonyx capensis were never positively identi-
ed. The ubiquitous C. civetta was considered the rarest species (IR = 0.89), followed by H. maculicollis (0.72) and I. albicauda (0.69),
whereas C. obscurus (0.01) was the commonest small carnivoran. Hunting techniques were mostly traditional guns, accompanied by
dogs, and jaw traps. Despite the absence of selective hunting, small carnivorans are likely to represent a fair source of income for
hunters, body parts being sold to fetish markets in 47% of the cases. Mean incomes range between US$ 2.5 and 5.4 per animal, with the
notable exception of C. civetta (US$ = 14.6) and heads of H. maculicollis, reaching US$ 33.7. The fair proportion of small carnivorans
observed on fetish market displays showed that hunting for animist practices might sustain a continuous hunting pressure in Benin. Our
preliminary survey raises a number of questions as to the distribution of small carnivorans in southern Benin, the impact of heavily dis-
turbed habitats on their survival and the level of sustainability of the hunting pressure they are subject to. Additional eld surveys will
be necessary for more precise characterisation of their status.
Keywords: bushmeat trade, distribution, ethnozoological survey, Herpestidae, Mustelidae, Nandiniidae, Viverridae
Les petits Carnivores du Sud Bénin: une évaluation préliminaire de leur diversité et de la pression de chasse
Résumé
Nous avons mené une mission de terrain étalée sur neuf semaines entre Août 2007 et Juin 2008 au sud Bénin, dans le but d’estimer la
diversité des petits Carnivores et la pression de chasse à laquelle ils sont soumis à travers des entretiens, des enquêtes sur les marchés et
des observations directes occasionnelles. Nous proposons un « Indice de Rareté » (IR), équivalent au nombre de fois qu’une espèce est
identiée comme « rare » par les interviewés rapporté au nombre de fois que celle-ci est mentionnée. Neuf espèces ou taxons (Genetta
spp.) ont pu être identiés sur la base de 86 chasseurs interviewés, représentant un total de 333 mentions. Les genettes, Crossarchus
obscurus et Herpestes ichneumon sont les plus cités, la prévalence d’espèces aussi versatiles d’un point de vue écologique conrmant
que le sud Bénin représente un « écosystème » perturbé. Les autres espèces sont Hydrictis maculicollis, Atilax paludinosus, Ichneumia
albicauda, Civettictis civetta, Mungos gambianus et Nandinia binotata. Nos observations directes nous ont permis de conrmer la
présence de G. pardina/maculata, C. obscurus, H. ichneumon et H. maculicollis. Grâce aux enquêtes de marché et à des identica-
tions moléculaires, nous avons clairement établi la distinction entre G. genetta, G. pardina/maculata and G. thierryi, établissant à 11
le nombre d’espèces de petits Carnivores présentes (ou probablement présentes) au sud Bénin. Au cours de nos entretiens, Galerella
sanguinea et Aonyx capensis n’ont jamais été mentionné. L’espèce ubiquiste C. civetta est considérée comme le petit Carnivore le plus
rare (IR = 0,89), suivie de H. maculicollis (0,72) et I. albicauda (0,69), alors que C. obscurus (0,01) est l’espèce la plus commune. Les
méthodes de chasse sont principalement la chasse aux fusils traditionnels accompagnée par des chiens, et les pièges à mâchoires. Mal-
gré l’absence d’une chasse sélective, les petits Carnivores représentent probablement une source non négligeable de revenus pour les
chasseurs, différentes parties de leur corps étant vendues sur les marchés des fétiches dans 47% des cas. Les revenus moyens oscillent
entre 2,5 et 5,4 US$ par animal, à l’exception de C. civetta (14,6 US$) et des têtes de H. maculicollis, pouvant atteindre 33,7 US$. La
proportion importante de petits Carnivores observés sur les marchés des fétiches montre que la chasse destinée à alimenter les pratiques
animistes implique très probablement une pression de chasse continue au Bénin. Notre étude préliminaire soulève un nombre de ques-
tions relatives à la répartition des petits Carnivores dans le sud Bénin, à l’impact des habitats fortement perturbés sur leur survivance et
à la durabilité de la pression de chasse à laquelle ils sont soumis, qui nécessiteront de nouvelles études de terrain an de tendre vers une
caractérisation plus précise du statut de ces mammifères peu connus.
Mots-clés: commerce de viande de brousse, enquête ethnozoologique, Herpestidae, Mustelidae, Nandiniidae, répartition, Viverridae
Small Carnivore Conservation, Vol. 40: 1–10, April 2009

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Djagoun and Gaubert
Small Carnivore Conservation, Vol. 40, April 2009
Introduction
Small carnivorans from western Africa have been little studied to
date. Despite their taxonomic diversity (Kingdon 1997, Wozen-
craft 2005), their variety of ecological functions — such as pest
controllers, seed dispersers, pollinators and an ambivalent prey/
predator condition — and their role as indicators of ecosystem
conditions (Kingdon 1977, Lack 1977, Charles-Dominique 1978,
Maddock & Perrin 1993, Duckworth 1995, Kingdon 1997, En-
gel 1998, Angelici et al. 1999a, Admasu et al. 2004a, De Luca
& Mpunga 2005, Martinoli et al. 2006, Dunham 2008), knowl-
edge on their natural history remains poor. As an example of the
paucity of the data available, distribution ranges for Viverridae,
Herpestidae, Nandiniidae and Mustelidae have remained specula-
tive for most of the subregion (Kingdon 1997). This is notably
because the available checklists (e.g. Rode 1937, Rahm 1961, Ae-
schlimann 1965, Kuhn 1965, Roche 1971, Happold 1987) are now
outdated and museum records are rare (Taylor 1989, Colyn et al.
1998, Grubb et al. 1998, Gaubert et al. 2002, Gaubert 2003). Sev-
eral publications dealing with the natural history of western Af-
rican small carnivorans have been made available more recently,
but their contribution remains anecdotal or episodic (Barnett et
al. 1996, Sillero-Zubiri & Bassignani 2001, Ziegler et al. 2002).
A recent study addressing the issue of low numbers of records
to reconstruct habitat suitability in some western African small
carnivorans showed that viverrids (such as Poiana leightoni and
several species of forest genets) were likely to be under threat
because of a high level of forest fragmentation in the subregion
(Papeş & Gaubert 2007). This sum of established facts thus sug-
gests an urgent reassessment of the conservation status of small
carnivorans from the subregion.
The status of small carnivorans in Benin is almost unknown.
Historical surveys are lacking and the museum record is virtually
empty (P.G. pers. obs.). Existing literature has focused on larger
mammals, including big carnivorans, and their conservation in
the faunistically rich biosphere reserves from the northern part of
the country (Sayer & Green 1984, Verschuren 1988, Boulet et al.
2004, Di Silvestre et al. 2004, Imorou et al. 2004, Assogbadjo et
al. 2005, Claro et al. 2006). One exception is the checklist of Hey-
mans (1984), where small carnivorans from northern Benin were
inventoried. Unfortunately, this is barely accessible grey literature
and, more signicantly, there is no indication about how species
identication was made and how distribution data were collected.
Although some recent ecological and ethnozoological sur-
veys have allowed the preliminary reassessment of the status of
a few small carnivorans (otters and mongooses) in southern Be-
nin, much of what is known remains anecdotal. Contrary to otters
(Hydrictis maculicollis, Aonyx capensis), which have become in-
creasingly rare in the area due to direct destruction and river dis-
turbance (Akpona 2004), some species of mongooses (Herpestes
ichneumon, Atilax paludinosus, Ichneumia albicauda, Crossar-
chus obscurus) seem to persist despite heavy anthropogenic pres-
sures (Djagoun et al. 2009).
In southern Benin, rainforest habitats are very patchily dis-
tributed into small forested islands and have been continuously
logged for agricultural development (Sayer & Green 1984). Since
small mammals from this region have been poorly surveyed, it
is not known whether the surveillance measures developed in
protected areas for larger mammals had a benecial effect on the
whole mammalian community. Despite a genuine will to promote
the farming of animals such as the Marsh Cane Rat Thryonomys
swinderianus as an alternative source of protein in Benin (Bap-
tist & Mensah 1980), factors such as demographic expansion and
widespread poverty, characterised by unemployment, few local
economic opportunities, and dependency on limited natural re-
sources contribute to the unstopped overexploitation of natural
ecosystems, including over-hunting of wildlife (Fa et al. 2003,
Ehui & Pender 2005, Bennett et al. 2006). In southern Benin, sev-
eral protected forested areas exist (such as ‘forêts classées’ and
‘forêts sacrées’) but do not benet from concrete conservation ac-
tions (Soury 2007). Small mammals are tempting targets for hunt-
ers, especially with the current monitoring of large game poaching
which will inevitably increase the pressure on less visible game
(Anadu et al. 1988, Codjia & Assogbadjo 2004). It is thus likely
that hunting activities have remained sustained, especially given
that small game in southern Benin represents most of the bush-
meat intake, to be used either for food consumption (Codjia &
Assogbadjo 2004, Assogbadjo et al. 2005) or as pharmacopoeia
(Tchibozo & Motte-Florac 2004) and fetish items.
Given that small carnivorans are usual constituents of the
small game trade in western Africa (Angelici et al. 1999b, Colyn
et al. 2004) and notably, as we are interested here, in Benin (As-
sogbadjo et al. 2005), our study intends to assess their diversity
and the hunting pressure to which they are subject in southern
Benin through interviews, surveys of local markets and occasional
eld observations. By doing so, we wish to provide a preliminary
update in the region from which better-informed management de-
cisions may be possible. Last, but not least, our survey should help
characterising the small carnivoran fauna of the southern Daho-
mey Gap area, a largely understudied stretch of forest–savannah
mosaic that separates the Upper and Lower Guinean rainforest
blocks.
Methods
Study area
This study was conducted in the southern part of the Republic of
Benin, encompassing a zone within 6°20′–7°40′N, 1°30′–2°50′E
(Fig. 1). The mean temperatures are constantly high (c. 25°C) with
daily amplitude below 5°C, and there is a great irregularity of an-
nual rainfall (mean = 1,200 mm; L’Hôte & Mahé 1996). Southern
Benin is located in the Dahomey Gap and has a subequatorial cli-
mate subdivided into four seasons of unequal length: two rainy
seasons (from April to July and September to October) and two
dry seasons (from November to March and end of July to Au-
gust). The vegetation is characterised by a great variety of frag-
mented phytocenoses resulting from a combination of climatic,
topographic and edaphic factors and human agency. The southern
part of Benin covers three ecological zones (Natta 2003): (1) a
plateau with Guineo–Congolian afnities representing semi-de-
ciduous forest, (2) the Lama depression, which is limited to the
North by the Zagnanado and Ketou plateaux and to the South by
the Allada and Sakete plateaux (this area is ooded during the
rainy season), and (3) gallery and swamp forests, patchily distrib-
uted along rivers. Dominant native trees in these zones are Ceiba
pentandra, Celtis mildbraedii and Dialium guineensis, whereas
exotic species mainly consist of Tectona grandis and Gmelina ar-
borea. Several emblematic species of mammals inhabit the forests
of southern Benin, including Mona Monkey Cercopithecus mona,
Vervet Monkey Cercopithecus aethiops, an endemic subspecies of

3
Small Carnivore Conservation, Vol. 40, April 2009
Red-bellied Monkey Cercopithecus erythrogaster erythrogaster,
and threatened ungulates such as Sitatunga Tragelaphus spekii,
Royal Antelope Neotragus pygmaeus, Black Duiker Cephalophus
niger and Yellow-backed Duiker C. silvicultor (Sinsin & Assog-
badjo 2002).
Surveys among hunters
An ethnozoological survey was conducted to gather information
on small carnivorans, representing a nine-week period spread be-
tween August 2007 and June 2008. Open interviews were conduct-
ed among 86 active hunters of 18 villages distributed in southern
Benin, representing about 2,500 km of road trip between Cotonou
(coast) and Dassa surroundings (206 km inland; Fig. 1). All the
people interviewed were men (hunting is not usual for women)
from seven ethnic groups (Fon, Ouémé, Idatcha, Mina, Adja,
Aïzo, Goun); their ages ranged between 23 and 67 years. Hunting
was a secondary activity, whereas agriculture was most generally
a primary occupation. A series of questions accompanied by illus-
trated identication sheets for each target species (dened follow-
ing Heymans 1984, Kingdon 1997, Wozencraft 2005) were asked
with the help of two local interpreters to collect information on
(i) the different hunted species of small carnivorans, (ii) among
them, which were considered rare, (iii) hunting techniques, and
(iv) the use made of the hunted animals (personal consumption,
trade, and related prices). As concerns point (ii), we calculated a
straightforward ‘Index of Rarity’ (IR) for each species, expressed
as the number of times a species is identied as ‘rare’ by inter-
viewed people / the number of time it is mentioned (because the
number of times a species can be identied as ‘rare’ depends on
the number of times it has been mentioned as present by hunters).
In summary, IR varies from 0 to 1, with higher values meaning a
higher level of rarity as expressed by hunters. Whenever possible,
the carcass, skin or other remains of the animals saved by hunters
were observed or collected to conrm taxonomic identication.
To minimise possible misidentications, species identity
was cross-checked during interviews with questions related to di-
agnostic morphological, behavioural and dietary traits (Table 1).
Despite that precaution, we realised that the three genets for which
we surveyed (Genetta genetta, G. pardina / G. maculata [see Gau-
bert 2003] and G. thierryi) were subject to potential confusion, the
different species being difcult to distinguish from our illustrated
sheets, but also for hunters in general (see Angelici et al. 1999a).
We thus treated all the mentions of genets under ‘Genetta spp.’.
Occasional survey of markets and road sellers
During August–July 2007, we twice had the opportunity to visit
fetish markets in Dantokpa (Cotonou) and Bohicon. Markets in
southern Benin seem not to be equivalent in size and purpose
to large bushmeat markets found in forested Africa (see Barnes
2002, Colyn et al. 2004, Edderai & Dame 2006, Fa et al. 2006,
Laurance et al. 2006, Albrechtsen et al. 2007, Bennett et al. 2007,
Willcox & Nambu 2007). Instead, bushmeat consumption and
trade, at least for small game (i.e. the most important wild source
of protein; Codjia & Assogbadjo 2004, Assogbadjo et al. 2005),
seems to be limited to local scales. The markets that we visited in
Dantokpa and Bohicon were selling animals as fetishes for tra-
ditional medicine and animist rites. The animals found on those
markets were mostly dried specimens (specimens may be left rot-
ten and drying at the feet of market displays), smoked heads, skins
or skulls and bones, which are conservable as such for months
or even years. Most displays spend the night unmoved, covered
with a piece of rough tissue. The rate of item renewal on market
displays appeared extremely low (at least for small carnivorans).
In Dantokpa, we noticed the presence of a wide variety of spe-
cies that suggested remote geographic origins of part of the sold
animals, including northern Benin but also neighbouring countries
(e.g. Leopard Panthera pardus, African Buffalo Syncerus caffer
and Hartebeest Alcelaphus buselaphus). Bohicon appeared to
Small carnivorans in southern Benin
Fig. 1. Map of southern Benin, showing the study sites.

4
house a more ‘local’ market, showing a less diversied taxonomic
assemblage, although species possibly from northernmost areas
were also represented.
In order to conrm species-level identications, we used nu-
cleotide sequencing (about 400 bp of cytochrome b) of what we
thought to be nine heads of genets (see Results) from the Bohicon
market, following protocols and primers used in previous studies
(Gaubert et al. 2004, Gaubert & Begg 2007).
Along the main road that borders the Lama classied for-
est (RNIE 2), we occasionally encountered points of ‘road sellers’
where fresh bushmeat was available. Such points were briey vis-
ited on four occasions during the same time period as our market
surveys.
Occasional sightings of small carnivorans made by one of us
(C.A.M.S.D.) during recent eld studies in the northern part of the
country were also reported.
Results
Interviews among hunters
Nine species or taxa (Genetta spp.) of small carnivorans were
identied through 86 interviewed hunters, representing a total
333 mentions (Fig. 2). Genets Genetta spp., the Cusimanse Cros-
sarchus obscurus and the Ichneumon Mongoose Herpestes ich-
neumon were clearly the most sighted species in southern Benin
(about 99%, 95% and 87% of the interviewees, respectively). The
Spotted-necked Otter Hydrictis maculicollis and the Marsh Mon-
goose Atilax paludinosus were mentioned by 34% and 27% of the
Djagoun and Gaubert
Small Carnivore Conservation, Vol. 40, April 2009
Table 1. Morphological, behavioural and dietary information on small carnivorans used to check species identication during
interviews.
Species Morphology Behaviour Diet
Genetta genetta spotted; tail long and annulated to the tip;
large ears
nocturnal; solitary; tree-climber carnivorous
Genetta pardina / G.
maculata
spotted; long annulated tail; dark distal part
of the tail; large ears
nocturnal; solitary; tree-climber carnivorous
Genetta thierryi smaller genet; spotted; long annulated
tail; dark distal part of the tail shorter than
above; large ears
nocturnal; solitary; tree-climber carnivorous
Civettictis civetta large size; spotted; shorter annulated tail
than genets; dorsal crest
usually nocturnal; solitary; not tree-
climber
omnivorous
Crossarchus obscurus uniform; short tail; small ears; long nose diurnal; gregarious; not tree-climber insectivorous
Atilax paludinosus large mongoose; uniform; short tail; small
ears
Mostly nocturnal; solitary; not tree-
climber; lives near rivers
piscivorous;
malacophagous
Herpestes ichneumon uniform; long tail with tip tapered in a dark
tassel; snake-like head
diurnal; solitary or in small families;
not tree climber; trotting with head
down
carnivorous (including
reptiles)
Ichneumia albicauda large mongoose; body usually grey-silver
with dark feet; long tail usually white
nocturnal; usually solitary; not tree-
climber
insectivorous
Mungos gambianus body grey with a white chest; short tail;
small ears
diurnal; gregarious; not tree-climber insectivorous
Galerella sanguinea small mongoose; uniform; long tail with tip
usually dark; eyes reddish
diurnal; solitary; may climb trees carnivorous;
insectivorous
Nandinia binotata dark and spotted; a pair of bright spots on
shoulders; long tail; feet with large pads
nocturnal; solitary; tree-climber frugivorous
Hydrictis maculicollis slender otter; throat with dark and bright
blotching; webbed toes
diurnal; usually solitary; not tree-
climber; lives in clear waters
piscivorous;
malacophagous
Aonyx capensis large otter with broad head; chest, chin and
cheeks whitish; unwebbed toes
diurnal; solitary; not tree-climber;
lives in a variety of water-related
habitats
mostly crabs (also
piscivorous and
malacophagous)
Fig. 2. Small carnivoran taxa identied
by hunters during interviews. White
bars correspond to the number of times
the taxa were mentioned (left scale),
and black bars represent ‘Index of Rar-
ity’ values (right scale, IR; see Meth-
ods). Because the Three-cusped Pango-
lin Manis tricuspis was also surveyed
during interviews, we here used it as a
reference value (see Results).

5
hunters, respectively, whereas the White-tailed Mongoose Ichneu-
mia albicauda represented 19%. Other, less mentioned, species
included the African Civet Civettictis civetta (10%), the Gambian
Mongoose Mungos gambianus (9%) and the African Palm-civet
Nandinia binotata (7%). During our interviews, the Slender Mon-
goose Galerella sanguinea and the African Small-clawed Otter
Aonyx capensis were never positively identied.
Civettictis civetta was considered the rarest species (IR =
0.89), followed by H. maculicollis (0.72) and I. albicauda (0.69)
(Fig. 2). To a lesser extent, A. paludinosus (0.57) and N. binotata
(0.5) were also identied as rare. Conversely, C. obscurus (0.01)
was considered the commonest species among the interviewed
people.
Comparing those results with a representative of another
mammalian family (Manidae) of similar body size, the Three-
cusped Pangolin Manis tricuspis was frequently mentioned by
hunters (69%), but was most of the time considered rare (IR =
0.76).
Hunting techniques were mostly traditional guns (52%), ac-
companied by dogs, and jaw traps (45%) (Fig. 3), which were
used in combination by 48% of the hunters. Hunting with packs of
dogs was a minor technique for small carnivorans (3%).
All interviewed people acknowledged that small carnivoran
meat was, partly or completely, used for personal consumption.
Nevertheless, in about 47% of the cases, body parts, such as skins,
heads and testes, were sold to fetish markets (Klouékanmè, Bo-
hicon), either directly or via resellers (Pahou, Cotonou, Taigon).
We managed to gather information on prices concerning sev-
en taxa sold to markets or resellers (Fig. 4). Civettictis civetta and
H. maculicollis were the most expensive species (mean = about
6,500 and 5,900 Francs de la Communauté Financière d’Afrique
[FCFA; US$ 14.6 and 13.3, as of August 2008], respectively). The
wide variation in the price value of H. maculicollis was due to
the fact that the species can be sold as body parts, including head
(3,000 to 15,000 FCFA [US$ 6.7 to 33.7]), skin (5,000 FCFA [US$
11.2]) or foot (1,000 to 2,000 FCFA [US$ 2.2 to 4.5]). Other taxa,
including genets, A. paludinosus, H. ichneumon and N. binotata,
were between 1,000 and 1,500 FCFA (US$ 2.2 and 3.4), whereas
C. obscurus represented a slightly greater source of income (mean
= 2,650 FCFA [US$ 6.0]). As a comparison, M. tricuspis was sold
at a mean of 2,250 FCFA (US$ 5.1).
Direct observations of animal remains allowed us to conrm
the presence of four species in areas neighbouring some inter-
viewed villages, including G. pardina / G. maculata (skins and
stuffed specimens, Fig. 5; Hon and Taigon), C. obscurus (anal re-
gion; Dèmè), H. ichneumon (stuffed specimen; Taigon), and H.
maculicollis (anal region and dried head; Kpomè).
Occasional surveys: fetish markets, road sellers and incidental
sightings
The fetish market in Dantokpa housed numerous dried heads of
three species of genets (G. genetta, G. pardina / G. maculata, G.
thierryi) and H. ichneumon, a few dried heads of H. maculicol-
lis, a few skulls and skins of C. civetta, and a few skulls of N.
binotata. It was remarkable to notice an unhidden, complete head
of Hippopotamus Hippopotamus amphibius together with some
rotting meat quarters. In Bohicon (Fig. 6), several dried heads and
skins of three species of genets (G. genetta, G. pardina / G. macu-
lata, G. thierryi) were present. Nucleotide sequencing of samples
taken from ve heads conrmed the identication of G. pardina
/ G. maculata and G. thierryi (data not shown). We also observed
numerous dried heads of H. ichneumon, very few dried heads of
H. maculicollis, and a couple of skulls of N. binotata. We also
noticed the presence of dried heads of what appeared to be domes-
tic dogs Canis lupus familiaris, and several skulls of unidentied
jackals Canis sp. After nucleotide sequencing, four dried heads of
what we thought to be, together with the main seller of the market,
G. genetta, actually belonged to a small canid not available in da-
Small Carnivore Conservation, Vol. 40, April 2009
Small carnivorans in southern Benin
Fig. 3. Local trap for small carnivorans, baited with palm nuts.
The jaw trap is buried at the entrance of a fence made of short
sticks. Leaves are put at the top of the trap to gure a small tree.
Fig. 4. Mean prices (in FCFA) of seven small
carnivoran taxa sold to markets or resellers.
Vertical bars indicate minimum and maximum
values. Again, the Three-cusped Pangolin is
used as a reference value (see Fig. 2).

6
tabases such as GenBank (http://www.ncbi.nlm.nih.gov/Genbank/
index.html; data not shown). After comparison of the prepared
skulls with photographic material (see Acknowledgements), we
conclude that those canids represent the Sand Fox Vulpes pallida.
Our sporadic surveys of road-side sellers bordering the Lama
forest failed to detect any species of small carnivorans. During
the eld period, we only happened to observe sporadic selling of
Marsh Cane Rats, Giant Pouched Rats Cricetomys gambianus,
Maxwell’s Duikers Cephalophus maxwelli, and one Bushbuck
Tragelaphus scriptus.
Incidental sightings made by one of us (C.A.M.S.D.) dur-
ing eld investigations in northern Benin in 2007 included a skin
of Zorilla Ictonyx striatus (Pendjari Biosphere Reserve: 10°30′–
11°30′N, 0°50′–2°00′E) and a tail belonging to C. civetta (Bétérou:
9°11–47′N, 1°58′–2°28′E). According to the local perception of
the populations living in neighbouring villages of the Pendjari, the
Wild Cat Felis silvestris, the Ratel Mellivora capensis, the Sand
Fox and N. binotata have been observed by hunters (C.A.M.S.D.,
pers. obs.).
Discussion
Diversity of small carnivorans in southern Benin
Our study does not intend to provide a denitive check-list of
small carnivorans from southern Benin. Rather, by combining
various sources of evidence from interviews, direct observation,
nucleotide sequencing identication, and incidental observations,
we aim at a refreshed, preliminary, assessment of their diversity
in this understudied region. Such a reassessment will have to be
further consolidated by eld studies and a more exhaustive cover-
age of sites, which would be benecial to extend to the northern
part of the country.
Nevertheless, some interesting results on the diversity of
small carnivorans from southern Benin were found (Table 2). A
total of 11 species, representing four families, Viverridae (four
species), Herpestidae (ve), Nandiniidae (one) and Mustelidae
(one), were identied by hunters and our molecular identica-
tions (genets). Remains of four of them were directly observed
in villages during our interviews (G. pardina / G. maculata, C.
obscurus, H. ichneumon, and H. maculicollis), those correspond-
ing to the most mentioned taxa by hunters (Fig. 2). Other species,
including G. genetta, G. thierryi, and N. binotata, were sighted in
the ‘local’ fetish market of Bohicon (but see below). The species
C. civetta was only observed in the larger fetish market of Dantok-
pa (Cotonou), but recent, direct observations of skins made by one
of us (C.A.M.S.D. pers. obs.) conrmed the natural occurrence of
the species in southern Benin. The presence of A. paludinosus,
although not directly observed here, was recently conrmed in the
Lama and Niaouli forests (Djagoun et al. 2009), as was also the
case for C. obscurus and H. ichneumon.
Two species of mongooses, I. albicauda and M. gambianus,
were never observed directly. Interestingly, Djagoun et al. (2009)
reported the presence of I. albicauda in southern Benin from a
low number of hunters’ mentions, but could not make any direct
observations of the species.
The commonest small carnivoran taxa appeared to be, ac-
cording to hunters, genets Genetta spp., C. obscurus and H. ich-
neumon (Fig. 2). The apparently high occurrence of the two spe-
cies of mongooses may be due to their ability (i) to occupy a wide
spectrum of habitats (Kingdon 1997, Palomares in press), as was
Djagoun and Gaubert
Small Carnivore Conservation, Vol. 40, April 2009
Fig. 5. Remains of a freshly killed G. pardina/maculata found in Hon (26°59′N, 02°18′E). The skin (a) is maintained a few days under
ashes, subject to a soft smoking technique. Parts of the body are separated for supposedly different uses, including the tail (b) and the
head (not shown).
Fig. 6. One of the displays of the Bohicon fetish market. Small
carnivorans, including genets and mongooses, are on the front
left.

7
observed for C. obscurus, and (ii) to live around crop elds and
to feed on poultry, as for H. ichneumon (Djagoun et al. 2009, and
comments of hunters from this study). In the case of genets, a
combination of both factors (genets were reported to kill poultry
frequently) may be the reason for their high occurrence (Admasu
et al. 2004a, Gaubert et al. 2008, Delibes & Gaubert in press). The
prevalence of such versatile species indicates that our survey zone
represents an environmentally disturbed area, where only small,
ubiquitous or prone-to-commensality species would be able to
maintain their populations.
Other small carnivorans, including C. civetta, H. maculicol-
lis, I. albicauda, A. paludinosus and N. binotata, were considered
rare by hunters. This may reect habitat specialisation affecting
survival in the context of intensive crops and teak plantation, as
found in southern Benin (Ganglo & de Foucault 2005, Barthès et
al. 2006), for species such as N. binotata (dependent on rainfor-
est canopy; Charles-Dominique 1978, Angelici et al. 1999a), and
H. maculicollis and A. paludinosus (dependent on forest galler-
ies and water; Kingdon 1997, Ray 1997). Although known to be
a ubiquitous species (Ray 1995, Kingdon 1997), C. civetta was
considered as the rarest small carnivoran, which may reect the
intense depletion of large- to mid-size game that has occurred in
the region. The high IR of the ecologically versatile I. albicauda
(Rosevear 1974, Kingdon 1997, Admasu et al. 2004b), together
with the absence of direct evidence for its presence, remains un-
explained. A possible restriction to a specic habitat such as low-
land forest in the particular case of southern Benin was previously
evoked by hunters (Djagoun et al. 2009). The scarcity of mentions
of M. gambianus, which may be locally common (IR = 0.38), also
remains unexplained. Confusion by hunters with C. obscurus may
have occurred (see Rosevear 1974), but ecological factors may
be a better, yet unknown, source of explanation if the species is
genuinely uncommon in the region.
One species of mongoose, G. sanguinea, presumed to oc-
cur in southern Benin (Kingdon 1997), was not cited by the
interviewees. Although Voglozin (2003) listed the species for
the Lama forest reserve, it has never been directly observed
there (C.A.M.S.D. pers. obs.). Despite the fact that G. sanguinea
may occupy a wide range of habitats, including forest swamps
(Kingdon 1997), it seems to avoid forested areas in southern
Benin but to occupy northern savannahs, where it causes con-
icts with people by preying on poultry (Djagoun & Sinsin
2007). Similarly, A. capensis, considered by Kingdon (1997) as
occurring in southern Benin, was not reported by interviewed peo-
ple. On the other hand, A. capensis may still be present in northern
Benin, where it was sighted near the Pendjari river (1978), and
recently around the Bali pool in the Pendjari Biosphere Reserve
(2002–2003; Di Silvestre et al. 2004).
Hunting pressure and trade
Traditional guns and jaw traps were predominantly used by hunt-
ers from southern Benin to catch small carnivorans. The use of
jaw traps and the lack of mentions of wire snares is a remarkable
feature here, since the majority of published studies on small- to
mid-game hunting in Africa showed the co-dominance of guns and
wire snares as the most usual techniques (Noss 1998, Willcox &
Nambu 2007). Snares may be suitable for more densely forested
regions than southern Benin, and the use of jaw traps may be a
more efcient technique in open habitats such as degraded forests,
savannah–forest mosaics and croplands.
Nearly all the species of small carnivorans present in
southern Benin were hunted. However, from the hunters’ com-
ments, and despite the fact that there was no selective hunt-
ing, small carnivorans were not a priority because of their small
size and low income values compared with large herbivores and
rodents (Codjia & Assogbadjo 2004, Assogbadjo et al. 2005).
Several bushmeat surveys in the African rainforest showed that
small carnivorans generally constitute a small but regular pro-
portion of the bushmeat intake (Anadu et al. 1988, Wilkie et al.
1992, Colyn et al. 2004, Fa et al. 2006; but see Angelici et al.
1999b). In southern Benin, this trend also seems to apply (Cod-
jia & Assogbadjo 2004). In our case, the majority of hunters
Small Carnivore Conservation, Vol. 40, April 2009
Small carnivorans in southern Benin
Table 2. Synthetic table of small carnivoran species diversity in Benin, compiling literature and inputs from the present study.
Species Heymans (1984) Kingdon (1997) Wozencraft (2005) This study3
Genetta genetta N Benin Benin Benin S Benin4
Genetta pardina / G. maculata N Benin Benin [taxonomic confusion1] S Benin
Genetta thierryi Benin Benin S Benin4
Civettictis civetta N Benin Benin Benin S Benin
Crossarchus obscurus S Benin Benin [under platycephalus]S Benin
Atilax paludinosus N Benin Benin S Benin
Herpestes ichneumon N Benin Benin S Benin
Ichneumia albicauda N Benin Benin S Benin
Mungos gambianus Benin S Benin
Galerella sanguinea N Benin Benin Benin N Benin
Nandinia binotata N Benin Benin Benin S Benin4
Hydrictis maculicollis N Benin Benin Benin S Benin
Aonyx capensis N Benin Benin Benin N Benin
Ictonyx striatus N Benin N Benin Benin N Benin
Mellivora capensis N Benin Benin [erroneous range2] N Benin
1 See Genetta pardina and genettoides (p. 556)
2 Wozencraft only mentioned “South Africa” as for the African distribution of the species (p. 612)
3 Our study includes two categories of species, namely those for which the occurrence in southern Benin (S Benin) could be ascertained (whether or
not they are present in northern Benin), and those that could not be found in southern Benin but that are present in northern Benin (N Benin)
4 But see Discussion

8
(94%) declared that their preferred meat was the Marsh Cane
Rat (similarly in Nigeria; Anadu et al. 1988), and mongooses were
sometimes reported by interviewees to be little valued because of
the strong, musky, smell from their anal glands. Nevertheless, sale
of small carnivorans is likely to represent a non-negligible income
for hunters. Indeed, whereas Marsh Cane Rats and Giant Poached
Rats were sold at US$ 8–10 and US$ 2–4, respectively, around the
Lama forest (Assogbadjo et al. 2005), mean incomes from small
carnivorans in southern Benin ranged between US$ 2.5 and 5.4
per animal for most of the species. Two exceptionally valued ani-
mals were H. maculicollis (US$ 13.3, with maximum price (head)
reaching US$ 33.7), and C. civetta (US$ 14.6). Thus, we suspect
that hunting pressure, although not specically targeted on this
group, will remain continuous.
Despite the fact that small carnivorans were not especially
prized as game, we could notice clear differences in the use/
value of some species, differences that could impact the level of
hunting pressure to which they are subject. The most remarkable
case is that of H. maculicollis, which appeared to be the most
prized small carnivoran because of the signicant income relat-
ed to its resale, mostly as body parts, to the fetish market. Some
hunters considered that H. maculicollis destroyed shing nets and
pots, and that the prices that could be obtained from its sale com-
pensated the damage to shing material. Similarly, the relatively
high resale price of C. civetta could increase hunting pressure on
that species, which may be vulnerable in southern Benin given its
large size and its level of rarity as estimated by hunters.
Mongooses have an ambivalent status in hunters’ percep-
tions, since (i) their strong smell seems to make them a less val-
ued game and, as insect- and snake-eaters (C. obscurus, H. ich-
neumon), they may represent ‘positive’ animals, but on the other
hand, (ii) their reputation as poultry killers (H. ichneumon) and
their use in specic animist rituals (A. paludinosus) may promote
their active hunting (also see Djagoun et al. 2009).
According to hunters, fetish markets were the main way in
which small carnivorans were sold. Although we do not pretend
to draw denitive conclusions from our episodic surveys, the low
representation of mongoose diversity on displays (only H. ichneu-
mon was sighted), compared with all the species of Nandiniidae,
Viverridae and Lutrinae reported by hunters, may suggest a higher
level of personal consumption (i.e. mongooses are little sold) that
would have gone undetected during our interviews, or simply a
lower rate of mongoose intake during the hunt (for reasons men-
tioned above).
The markets of Dantokpa and Bohicon exhibited a similar
set of small carnivoran remains, with the exception of C. civetta,
which was only found in Dantokpa. Given the broader geographic
origins suspected for the animals exhibited in the latter, a propor-
tion of C. civetta items may originate from northernmost regions
and/or neighbouring countries. In Bohicon, which was supposed
to be a local market, the presence of Vulpes pallida also suggests
that at least a part of the remains displayed come from animals
taken from northernmost areas (see Kingdon 1997). Thus, remains
of G. genetta, G. thierryi and N. binotata sold in Bohicon (see
above) are not 100% reliable evidences that those species inhabit
southern Benin.
The presence of a fair proportion of small carnivorans on
fetish market displays showed that hunting for animist practices
constituted a regular pressure on populations (for a similar situ-
ation in Nigeria, see Angelici et al. 1999b). It remains, however,
unknown whether the hunting, either for personal consumption
or resale, is sustainable. Remarkably, the rate of item renewal on
market displays appeared extremely low (but our survey period
was short).
Conclusions
Because of great agricultural pressure and deforestation, natural
habitat destruction is almost complete in southern Benin (Siebert
& Elwert 2004). Very reduced patches of almost unprotected for-
ests are now scattered across the area, and the impact of the pre-
dominance of disturbed / cropland habitats on the survival of small
carnivorans is totally unknown. In conjunction with this, hunting
pressure, although opportunistic, on those taxa seems sustained
and may thus be a serious threat to the survival of some species
already affected by habitat destruction, especially since national
protection does not exist for small carnivorans.
Despite the fair diversity of West African small carnivorans
in southern Benin found on this preliminary survey, questions re-
main open relative to (i) the absence of direct observations of two
species of mongooses (I. albicauda and M. gambianus), and (ii)
the ecological status of all species in the region. Unlike the more
conspicuous large carnivorans, populations of small carnivorans
can decrease with little notice (Buskirk & Zielinski 2003). Thus,
there is an urgent need for additional eld surveys in Benin in
order to test the hypotheses of distribution and status built from
the interviews of hunters. In return, a better characterisation of
small carnivoran diversity, population history and ecology should
enlighten our understanding of the Dahomey Gap as a key factor
in the structuring of the West African fauna.
Acknowledgements
We wish to thank the representation and staff of the Institut de Re-
cherche pour le Développement in Cotonou, Benin, for providing the
necessary infrastructure to our eld trip. We are grateful to S. Tchibo-
zo and R.S. Gantoli for assistance in initiating the project in southern
Benin. We thank H. Akpona for fruitful exchanges on the Beninese
biodiversity. We are grateful to all the interviewees who accepted to
share some of their time and experiences with us. C. Voisin prepared
the material for taxonomic identication. Jim Dines (Natural Histo-
ry Museum of Los Angeles County, CA), Linda Gordon (Smithsonian
Institution, National Museum of Natural History, Washington, D.C.),
Frieder Mayer (Museum für Naturkunde, Berlin, Germany) and Paula
Jenkins (Natural History Museum, London, U.K.) kindly provided pho-
tographs of Vulpes pallida and Rüppell’s Fox V. rueppellii. Extrac-
tions and PCRs (Polymerase Chain Reactions) were done by P.G. at
the Service de Systématique Moléculaire, Muséum National d’Histoire
Naturelle, Paris (France). Sequencing was conducted by Genoscreen,
Lille (France). This study was partly funded by the Société des Amis
du Muséum national d’Histoire naturelle et du Jardin des Plantes, Paris
(France).
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1Laboratoire d’Ecologie Appliquée, Faculté des Sciences
Agronomiques, Université d’Abomey-Calavi, 01 BP 526
Cotonou, Benin.
Email: sylvestrechabi@gmail.com
2UR IRD 131, Département Milieux et Peuplements
Aquatiques, Muséum National d’Histoire Naturelle, 43 rue
Cuvier, 75005 Paris, France.
Email: gaubert@mnhn.fr
*Corresponding author