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Assessing local knowledge on the diversity and abundance of bushmeat species and hunting pressure in the fragmented forest islands of southern Benin (Dahomey Gap)



Illegal hunting of wildlife is one of the major issues in tropical ecosystems, especially when it occurs in highly degraded habitats with forest cover fragmentation. In this study, we assessed the impact of bushmeat hunting in a large forest patch (the Lama Natural Forest; LNF) and 11 nearby forest islands, using Traditional Ecological Knowledge from 240 interviewees across 16 villages. Thirty-five species belonging to nine orders of mammals, birds and reptiles were mentioned by local communities. Rodentia were significantly more observed in the forest islands, whereas medium-sized mammals belonging to Carnivora, Primates, Artiodactyla, Pholidota and Hyracoida were found predominantly in LNF. Approximately 57% of the species were reported to be rare in the forest islands, whereas c. 77% were listed as abundant in LNF, confirming the role of LNF as a refuge for forest species targeted by the bushmeat trade. Generalised linear models indicated that species sighting frequencies were positively correlated with perimeters of forest patches. We found hunting pressure to be greater in forest islands in the vicinity of LNF than those further away. Our results suggest that long-term conservation of wildlife in southern Benin may require a ‘mainland-islands’ approach including both LNF and its surrounding forest islands.
Afr J Ecol. 2021;00:1–10.
1© 2022 John Wiley & Sons Ltd
Received: 9 March 2021 
Revised: 7 Octob er 2021 
Accepted: 24 November 2021
DOI: 10.1111/aje.12955
Assessing local knowledge on the diversity and abundance
of bushmeat species and hunting pressure in the fragmented
forest islands of southern Benin (Dahomey Gap)
Chabi A. M. S. Djagoun1| Gilles Nago2| Akomian F. Azihou1|
Fifanou Vodouhê2| Auriane Agli1| Stanislas Zanvo1,3 | Bruno Djossa1,4 |
Achile Assogbadjo1| Brice Sinsin1| Philippe Gaubert3,5
1Labor atory of Applied Ecolog y, Faculty
of Agron omic Sciences, University of
Abomey- Calavi, Cotonou, Benin
2Labor atoire d’Ecologie, de Botaniqu e et
de Biologie végétale, Faculté d’Agronomie,
Université de Par akou, Parakou,
République du Bénin
3Unité de Recherche en Fores terie
et Conservation des Bioresso urces
(U/RFCBio), Uni versité Nationale
d'Agriculture de Porto- Novo, Porto- Novo,
4Labor atoire Evolution et Diversité
Biologique, UPS/CNRS/IRD, Universi
Paul Sabatier, Toulouse, France
5CIIMAR , Terminal de Cruzeiros Do Por to
de Leixoe s, Avenida Gener al Nor ton de
Matos s/n, Matosinhos, Portugal
Chabi A . M. S. Djagoun, L aboratory of
Applie d Ecolog y, University of Ab omey-
Calavi, Faculty of Agronomic Sciences ,
University of Abomey- Calavi, 01 B P 526,
Cotonou , Benin.
Funding information
program Jeune Equipe Associée à l’IRD
Illegal hunting of wildlife is one of the major issues in tropical ecosystems, espe-
cially when it occurs in highly degraded habitats with forest cover fragmentation. In
this study, we assessed the impact of bushmeat hunting in a large forest patch (the
Lama Natural Forest; LNF) and 11 nearby forest islands, using Traditional Ecological
Knowledge from 240 interviewees across 16 villages. Thirty- five species belong-
ing to nine orders of mammals, birds and reptiles were mentioned by local com-
munities. Rodentia were significantly more observed in the forest islands, whereas
medium- sized mammals belonging to Carnivora, Primates, Artiodactyla, Pholidota
and Hyracoida were found predominantly in LNF. Approximately 57% of the species
were reported to be rare in the forest islands, whereas c. 77% were listed as abun-
dant in LNF, confirming the role of LNF as a refuge for forest species targeted by the
bushmeat trade. Generalised linear models indicated that species sighting frequencies
were positively correlated with perimeters of forest patches. We found hunting pres-
sure to be greater in forest islands in the vicinity of LNF than those further away. Our
results suggest that long- term conservation of wildlife in southern Benin may require
a ‘mainland- islands’ approach including both LNF and its surrounding forest islands.
forest islands, poaching, protected areas, southern Benin, traditional ecological knowledge,
Wildlife trade
La chasse illégale de la faune sauvage est l'un des problèmes majeurs des écosystèmes
tropicaux, surtout lorsqu'elle se produit dans des habitats fortement dégradés avec
une fragmentation du couvert forestier. Dans cette étude, nous avons évalué l'impact
de la chasse de viande de brousse sur une grande aire forestière (la Forêt Naturelle
de la Lama; FNL) entourée de 11 îlots forestiers, en utilisant les connaissances
écologiques traditionnelles de 240 personnes interrogées dans 16 villages. Trente- cinq
(35) espèces appartenant à neuf ordres de mammifères, d'oiseaux et de reptiles ont
été mentionnées par les communautés locales. Les rongeurs ont été significativement
    DJAGO UN et Al.
In Sub- Saharan Africa, illegal exploitation and trade in forest re-
sources (e.g. timber and bushmeat) have serious implications for bio-
diversity conservation and ecosystem services (Price & Gittleman,
2007). Such deleterious implications are exacerbated by current
trends of deforestation levels, as observed in West Africa where at
least 22% of the forest sur face is degraded by human activities (FAO
& UNEP, 2020). Uncontrolled deforestation leads to wildlife habi-
tat fragmentation and increased hunting activities, promoting local
faunal extirpation (Sreekar et al., 2015; Tilker et al., 2019). Romero-
Muñoz et al. (2019) showed that wildlife occurring in fragmented
areas tended to suffer more frequently from over- hunting than in
areas with continuous forest cover, as small forest patches are more
accessible to hunters.
In West Africa, the Dahomey Gap is a biogeographic barrier of
forest– savannah mosaics separating the Upper and Lower Guinean
forest blocks. The cumulated effect of Plio- Pleistocene climatic
oscillations and ancient agriculture have fragmented the area rain-
forests into sparse, isolated forest islands (Demenou et al., 2016;
Poorter et al., 2004). Benin, a countr y fully included in the Dahomey
Gap zone, provides a representative example of the superimposed
effects of climatic fluctuations and human activity on forest frag-
mentation and faunal depletion (Kokou et al., 2008). Such effects
are currently exacerbated by illegal forest exploitation including
hunting, harvesting, transport, processing, purchase and sale of
forest and wildlife products in violation of national laws (Djagoun
& Gaubert, 2009; Djagoun et al., 2018a; 2018b), turning southern
Benin into an extensive agricultural landscape (Siebert & Elwert,
2004). As a consequence, several vertebrate species have been ex-
tirpated from their natural habitats or have seen a decline in their
population status (Djagoun et al., 2018a; 2018b).
The bushmeat trade has been exhaustively investigated in West
and Central Africa, where this activity is thriving (e.g. Ingram et al.,
2021; Ziegler et al., 2016). Producing accurate estimates on the sus-
ceptibility of species to hunting in the tropics is a challenging task
requiring long- term surveys of both the bushmeat network and an-
imal populations, ideally across various study sites. However, it has
been suggested that local populations living in villages surrounding
protected areas often have important and long- standing Traditional
Ecological Knowledge (TEK) on forest biodiversit y (Uddin & Foisal,
2007). TEK is transferred from one generation to the next, repre-
senting cumulative local knowledge, and is amended following re-
cent experiences and observations (Fernandez- Gimenez, 2000). As
such, TEK is being sought by academics, scientists and policymakers
as a poten ti al sour ce of da ta fo r emerg ing mod els of ecos ystem man-
agement, conservation biology and ecological restoration (Ramos,
2018; Weiss et al., 2013). Consequently, as an alternative to direct
observations, TEK (Ramos, 2018) can be a useful approach to collect
indirect data on wildlife species diversity, distribution, abundance
and response to harvest pressure (Brinkman et al., 2009).
In Benin, several mammalian species went extinct because of
overhunting and habitat fragmentation, including the chimpan-
zee (Pan troglodytes; Djagoun et al., 2018b; Nombine et al., 2008),
the bongo (Tragelaphus eurycerus; Neuenschwander et al., 2011)
and probably the giant pangolin (Smutsia gigantea; Zanvo et al.,
2020). Our study focuses on southern Benin, where forest cover
is constrained into small to larger forest patches (0.26– 4777 ha;
Agonvonon, 2018), representing the last- standing shelters for a va-
riety of endemic and/or threatened species including the red- bellied
monkey (Cercopithecus erythrogaster erythrogaster). In southern
Benin, bushmeat activities are concentrated at a single, main bush-
meat market (Têgon) which is located at the periphery of the region's
major protected forest, namely the ‘Forêt classée de la Lama’ or
Lama Natural Forest (LNF; Djagoun et al., 2018a). LNF, the main-
land patch, is surrounded by several small forest patches (islands)
which forms a mainland– islands complex (Agonvonon, 2018). So
far, the levels of impact of the bushmeat trade across this complex
have remained unknown. Our objectives were to assess, using TEK
from local communities, (i) the diversity and population status of the
species hunted as bushmeat and (ii) the main drivers of bushmeat
hunting. According to the theory of island biogeography (Ewers &
plus observés dans les îlots forestiers, tandis que les mammifères de taille moyenne
appartenant à l’ordre des Carnivores, Primates, Artiodactyles, Pholidotes et
Hyracoïdes ont été trouvés principalement dans la FNL. Environ 57% des espèces
ont été signalées comme étant rares dans les îlots forestiers, alors que 77% d'entre
elles ont été pertoriées comme étant abondantes dans la FNL, ce qui confirme le
rôle de cette dernière en tant que refuge pour les espèces forestières ciblées par le
commerce de viande de brousse. L’analyse des modèles linéaires généralisés a indiqué
que les fréquences d'observation des espèces étaient positivement corrélées avec les
périmètres des forêt/îlots forestiers étudiés. Nous avons constaté que la pression de
chasse était plus forte dans les îlots forestiers situés à proximité de la FNL que dans
ceux qui en sont plus éloignés. Nos résultats suggèrent que la conservation à long
terme de la faune sauvage dans le sud du Bénin pourrait nécessiter une approche
"continent- îles" intégrant à la fois la FNL et les îlots forestiers environnants.
Didham, 2006), we hypothesise that (i) LNF (mainland) will harbour
mor e species and hea lt hier populations tha n small for es t patches (is-
lands). As a result, local communities will perceive greater species
abundance in LNF and that (ii) hunting activities are greater in non-
protected forest islands.
2.1  |  Study area
The Lama Natural Forest (LNF) is located in southern Benin, be-
tween 6°55′ and 00′ N, and 2°04′ and 2°12′ E. The surrounding
small forest patches (islands) are within a 20- km radius. The latter
derive from extensive logging activities for agricultural development
and cover a total area of 53.34 ha (Figure 1, Table 1). LNF covers
4777 ha of natural forest called ‘Noyau Central’, strictly protected
and surrounded by teak plantations (Adomou, 2005). The ethnic
groups living around the forest patches include Holli, Fon and Aïzo
(INSAE, 2015). The climate is of dry Guinean type with mean an-
nual rainfall of 1200 mm and annual average temperature between
25 and 29°C (Kakpo et al., 2021). In the study area, strata of dense
forest are separated by fallow lands at various stages of evolution,
testifying the recent forest degradation by farmers. Rural commu-
nities from the two districts included in the study area represent
92,935 (Zogbodomey) and 101,535 ( Toffo) inhabitants (INSAE,
2015). Despite poaching activities, LNF is home to endemic mam-
malian lineages of high conservation concern, including the white-
bellied pangolin (Phataginus tricuspis; Gaubert et al., 2016; Zanvo
et al., 2020) and the red- bellied monkey (Houngbedji et al., 2012).
2.2  |  Data collection
This study was approved by the Public Forest Service through and re-
ce ive d the pr ior cons ent from lo c al au tho ritie s in eac h vil lag e. In total ,
we su r vey e d 16 vi l l ages surro undi ng LN F an d th e smal l for est patc hes
(islands) from July to November 2019. The villages were chosen ac-
cording to their proximity to the forests and the records of poaching
activ ities in th e area, as ex trac te d from previous stud ies (A ssogbadjo
et al., 2005; Djagoun et al., 2018a; Zanvo et al., 2021). Hunting in
Benin is limited to certain species and is regulated by law n° 2002- 16
(18 October 2004) on wildlife protection and decree n° 2011- 394
(28 May 2011). In the study area, bushmeat hunting is mostly done
illegally and remains a sensitive topic for rural communities (Djagoun
et al., 2009). Thus, we used a mixed- method approach to triangulate
FIGURE 1 Study area showing the Lama Natural Forest (mainland) and surrounding small forest patches (islands)
    DJAGO UN et Al.
findings, as recommended by Harris and Brown, (2010). This involved
using semi- structured interviews (containing open- ended questions)
to gather data on the TEK of bushmeat species such as abundance
versus rarity and the number of individuals hunted per species. Field
assistants conducted interviews in the main local language (holli).
Interviewers avoided sampling people congregated in one area (e.g.
around shops; King, 2014) as they may fear to respond in public
areas. The selec ted interviewees were at least 20 year s old and were
permanent residents for >5 years in the surveyed villages. In order
to efficiently locate long- term residents willing to participate in our
interviews, we asked community elders to establish initial groups of
interviewees. The subsequent interviewees were identified through
chain referrals, where each member of initial groups suggested the
names of fur ther interviewees (Huntington, 200 0).
In each of the 16 villages, we conducted 15 interviews on a vol-
untary basis, totalling 240 people distributed as follows: 74 hunters
and 38 former hunters, 108 farmers, 6 forest guards and 14 em-
ployees of local authorities. Prior to interviews, each par ticipant was
informed that (i) information would remain anonymous, (ii) all the
questions did not have to be answered and (iii) withdrawal was pos-
sible. They were then asked for permission to proceed. Interviews
were organised after trust- based relationships were built with key
informants representing selected villages. According to Knapp et al.,
(2010), self- reporting based on trust- developed relationships remains
the only way to collect sensitive data such as poaching activities.
This technique complements random samples like the randomised
response technique because it more effectively dispels fear in the
respondent, ensuring greater validit y in responses. It is assumed that
poachers who lacked adequate trust in the interview team will not
agree to being interviewed. From there we were able to establish the
approximate number of hunters involved in our study and develop a
collaborative relationship with them. Additionally, we had explained
the objectives and approach of our research and included the hunt-
ers as active informants. Finally, to ensure the quality of the data, the
same question regarding the number of hunted individuals within
a year was asked to the same inter viewee af ter a two weeks’ time
period, for cross checking. The aim of these interviews was to collect
quantitative estimates on bushmeat species in the study area (per-
ceptions on population status and hunting pressure) and qualitative
information on the drivers of bushmeat hunting. Photos of a series
of representative bushmeat species (as extracted from Assogbadjo
et al., 2005; Djagoun & Gaubert, 2009; Djagoun et al., 2018a) were
presented to the respondents to get their opinion on the presence/
absence and abundance of each species.
The questions (see details in Appendix 1) were designed fol-
lowing previous investigations on similar topics (Djagoun et al.,
2018a; Luiselli et al., 2016). These focused on: (i) existence or ab-
sence of the species in the targeted forest; (ii) abundance versus
rarity of the bushmeat species ( Wright & Priston, 2010); (iii) max-
imum number of species sightings within 1 year; (iv) numbers of
individuals hunte d per spe ci es within 1 year. We also collected th e
socio- demographic characteristics (including age, sex, household
size, education level, occupation and time of residency in the vil-
lage) of the interviewees.
2.3  |  Data analysis
Descriptive statistics were used to summarise answers, while per-
centages and graphs were plotted for better visualisation of the
results using R version 3.1.1 (R Core Team, 2006). We presented
data as the percentage of respondents confirming the abundance or
rarity of the bushmeat species and the percentage of species sight-
ings grouped per taxonomic order (Primates, Rodentia, Carnivora,
etc.). Chi- square (χ2) test s of goodness- of- fit were used to establish
whether or not the number of bushmeat species sighted between
LNF and forest islands was significantly different. We selected the
best predictor variables using generalised linear models (GLMs) in
R, following a two- step procedure. First, we generated univariate
GLMs to examine which predictor variables among forest patch
perimeter (Per), forest patch surface (Surf), nearest distance of the
forest islands to the mainland (NDist) and household size (HSiz)
Forest name Surface (ha) Perimeter (m) Nearest distance to LNF (km)
Lama Natural Forest 4777 74,8 0 0 0
Zounkidjazoun 28.98 2900 8.32
Kpofouonzoun- Tohiozoun 9.9 0 1700 7.9 2
Adjahountozoun 5.47 140 0 1.44
Agbogbozoun 4.60 1200 10.39
Vodounzoun 1.71 539 3.20
Gbékossozoun 0.65 330 6.94
Sozoun 0.65 361 2.44
Zounhouè 0.41 242 3 .41
Houéhounzoun 0.36 282 7.71
Lezoun- Tangnikpèhoué 0.36 293 7. 23
Tog odo zo u n 0.26 223 8.07
Abbreviations: ha, hectare; km, kilometer; m, metre.
TAB LE 1  Geographic details of the
forests studied in southern Benin. Lama
Natural Forest represents the ‘mainland’,
whereas the other 11 forests represent
the ‘islands’ (small forest patches)
were significantly associated (p 0.05) with the number of bush-
meat species and hunting pressure (response variables). Second, we
checked collinearity among variables in the model by calculating the
Spearman's correlation coefficient among pairs of variables in order
to include only variables with low correlations (rs ≤ 0.6).
Model selection was conducted following a backward stepwise
proced ure, using Akaike's Information Criter ion (Akai ke, 1998) to re-
move parameters stepwise (factors were excluded if it improved the
model fit by >2 AIC unit s) and select the most parsimonious model
with the best fit to the data (Burnham & Anderson, 2002). We fit-
ted a Bayesian regression to predict the probability of being close
to the mainland with the number of individuals hunted across all
species using the packages tidyverse (Venables & Ripley, 20 02) and
Psycho (Makowski, 2018) in R. As Bayesian models usually gener-
ate many samples (iterations), we computed the median of the pos-
terior prediction, as well as the 95% confidence interval for better
Semi- structured interviews gathered 240 respondents aging be-
tween 20 and 40 years old. All were unemployed male educated to
primar y and farmers. The household size average was around seven
individuals/household. The time of residency in the village was
11 years in average.
3.1  |  Proportion of sightings according to
taxonomic orders between the Lama Natural
Forest and forest islands
A total of 35 species and nine orders belonging to mammals, birds
and reptiles, were sighted by the interviewees (Figure 2). The num-
ber of sightings per order within a year varied significantly between
LNF and forest islands (χ2 = 378.06, df = 8, p < 0.001). According to
local perceptions, Rodentia were significantly more observed in the
fore st is la nds than in LNF (χ2 = 6. 3 3 7, df = 1 , p = 0.011). On th e ot her
han d, mediu m- s ized mammal s such as Ca rn ivora (χ2 = 32.467, df = 1,
p < 0.001), Primates (χ2 = 198. 02 , df = 1, p < 0.001), Artiodactyla
(χ2 = 252.59, df = 1, p < 0.001), Pholidota (χ2 = 35.504, df = 1,
p < 0.001) and Hyracoida (χ2 = 21.12, df = 1, p < 0.001) were signifi-
cantly predominant in LNF.
3.2  |  Local perceptions of the abundance of
bushmeat species
Eighteen species out of 35 were repor ted by ≥90% of the respond-
ents to be rare in the forest islands, including Carnivora (Civecttictis
civetta, Ichneumia albicauda, Herpestes ichneumon, Atilax paludinosus),
Artiodactyla (Tragelaphus spekii, Cephalophus silvicultor, Cephalophus
niger, Potamochoerus porcus, Philantomba walteri, Tragelaphus scrip-
tus), Primates (Cercopithecus mona, Colobus vellerosus, Erythrocebus
patas, Cercopithecus erythrogaster, Potto, Procolobus verus), Pholidota
(Phataginus tricuspis) and Squamata (Python sebae) (Figure 3). Only
four speci es were repor ted wit h simil ar rate s to be rar e in LNF, includ-
ing three species of C arnivora (C. civetta, I. albicauda, H. ichneumon)
and one species of Artiodact yla (C. silvicultor). A total of 23 species
were liste d as abu ndant in LNF by more than 50% of th e in te r vi ewees ,
the top 10 species representing Rodentia (Thryonomys swinderianus,
Cricetomys spp., Arvicanthis niloticus, Xerus erythropus), Carnivora
(Crossarchus obscurus), Ar tiodactyla (P. porcus), Primates (Galago sen-
egalensis), Lagomorpha (Lewis crawshayi), Galliformes (Francolinus
ahantensis) and Squamata (Bitis arietans). Only nine species wer e listed
as abundant in forest islands in more than 50% of the cases, including
Rodentia (T. swinderianus, Cricetomys spp., A. niloticus, X. erythropus),
Carnivora (C. obscurus), Lagomorpha (L. crawshayi), Galliformes (F.
ahantensis) and Squamata (Varanus niloticus, B. arietans).
3.3  |  Forest patch characteristics predicting the
sighting frequency of bushmeat species
The most parsimonious GLM included predictors such as forest
patch perimeter (Per), forest patch surface (Surf) and the nearest
FIGURE 2 Relative percentages of
bushmeat sightings (per taxonomic order
and within a year) by local communities
between the Lama Natural Forest and
surrounding small forest patches (islands)
Rodentia CarnivoraSquamata Primates Galiformes Lagomorpha Artiodactyla Hyracoidea Pholidota
Proportional fauna event sighting (%)
Lama ForestForest Islands
    DJAGO UN et Al.
distance of the forest islands to the mainland (NDist), but showed
that only perimeter (Per) had a significant, positive influence on the
annual sighting frequency of bushmeat species (Table 2). In other
words, the larger the perimeter of a forest patch, the greater were
the sighting frequency of bushmeat species according to the local
communities (OR = 5%; p = 0.003).
3.4  |  Drivers of bushmeat hunting pressure in the
forest islands of southern Benin
The best GLM showed that household size (HSiz) and NDist were the
major drivers of the numbers of individuals hunted across all spe-
cies within a year in the forest patches of southern Benin (Table 3).
However, only NDist was significant (OR = 0 . 5 %; p < 0. 0 01) . Bay esi an
regression predicted that hunting pressure was greater in forest is-
lands at the vicinity of LNF than those further away (Figure 4).
We present the first exhaustive sur vey on the diversity and abun-
dance of the ver tebrate species hunted in the forests of southern
Benin. Bushmeat species richness within the study area reached 35,
which is higher than the number of species (20) previously identified
in LNF using a similar TEK- based approach (Djagoun et al., 2018a).
Our present study encompasses a larger, more heterogeneous net-
work of forest habitat s including LNF and small forest islands within
a 20- km radius, which could explain the higher species diversity ob-
served (habitat heterogeneity hypothesis; MacArthur & MacAr thur,
1961). The greater number of respondents who were considered in
our study (240 vs. 65 in Djagoun et al., 2018a) could also explain this
difference. According to Rist et al., (2010), TEK- based approaches
may compromise on accuracy for specific variables when using a re-
duced sample size. Because we were able to interview a large num-
ber of people across different local communities, we consider that
the data obtained in this study are accurate and reliable information
FIGURE 3 Local perceptions on the abundance of bushmeat species in the (a) Lama Natural Forest and (b) surrounding small forest
patches (islands)
TAB LE 2  Relative contributions of the forest characteristics
associated with the sighting frequency of bushmeat species
Variables Sign
97. 5%
CI p- value
Per +5.133 1.000 1.005 0.003*
NDist +0.98 4 0. 574 1.686 0.27 7
Surf +1.008 0.585 1.732 0.982
Abbreviations: CI, confidence interval; NDist, nearest distance of the
forest islands to the Lama Natural Forest; Per, perimeter; Sign, sign of
the correlation; Surf, forest patch surface.
*Significant at 0.05.
TAB LE 3  Main drivers of bushmeat hunting pressure in the
forests of southern Benin
Variables Sign
97. 5%
CI p- value
HSiz +0.985 0.963 1.009 0.233
NDist +0.579 0.492 0.683 0.000*
Abbreviations: HSiz, household size; NDist, nearest distance of the
forest islands to the Lama Natural Forest.
*Significant at 0.05.
(Berkes et al., 2000). Using a hunting recall period of 1 year can
be considered large for a TEK- based approach (Jones et al., 2008).
However, in order to prevent from this potential bias, we repeated
the interview with the same persons as accuracy is increased when
the same informants are questioned iteratively (Golden et al., 2013).
On the other hand, short recall periods are vulnerable to systematic
and response errors in the context of seasonal variation of the hunt-
ing events (Johansson et al., 2002).
Our study tends to confirm the role of LNF as a crucial refuge
for forest species targeted by the bushmeat trade, which in itself
justifies its national protection status (Nagel et al., 2004). A signifi-
cantly higher number of medium- sized mammals were sighted in
LNF than in forest islands. This fits with our first hypothesis based
on the theory of island biogeography, where mainland (LNF) har-
bours healthier populations when compared with islands (small
forest patches) (Ewers & Didham, 2006; Prugh et al., 2008; Sreekar
et al., 2015). Since its management by the ‘National Timber Office
of Benin— ONAB’ authority (ONAB, 2010), LNF has been secured
from uncontrolled occupation in its core area notably through the
translocation of local communities from illegally occupied zones.
Such actions allowed for the recovery of up to 4777 ha of natural
semi- deciduous forest named the Noyau Central’ (core zone), which
is only accessible to scientific research and biodiversity conservation
programmes. Rangers trained and employed by ONAB to secure the
‘Noyau Central’ are conducting anti- poaching activities on a regular
basis. Altogether, this may maintain the remarkable abundance of
bushmeat species repor ted in LNF (Assogbadjo et al., 2005; Djagoun
et al., 2009, 2018a), further testifying of the impor tance of LNF in
the conservation of the forest biodiversity of southern Benin (Nagel
et al., 2004). The fact that four species listed as of high conservation
concern on the IUCN Red List of Threatened Species (C. erythro-
gaster, P. Potto, C. vellerosus, P. tricuspis) were identified in 50% or
more of the cases as rare in LNF should call for further surveys to
characterise the population status of these taxa in the core area and
the potential factors behind this status (e.g. bushmeat trade, illegal
habitat degradation).
We found a positive correlation between forest patch perime-
ter and sighting frequencies of bushmeat species. This meets the
expectations from the theory of island biogeography, with larger
forest patches harbouring more abundant populations than smaller
patches (Palmeirim et al., 2018). Because of a lack of both law en-
forcement and long- term conservation policies, forest islands from
southern Benin have rapidly been degraded by logging and agricul-
ture activities (Nagel et al., 200 4). In line with the general decline
in wildlife observed in the fragmented forest habitats of southern
Benin (Assogbadjo et al., 2005; Djagoun et al., 2009, 2018a; Zanvo
et al., 2020), local communities from forest islands cited as rare many
medium- sized species of Artiodactyla (T. spekii, C. silvicultor, C. niger,
P. walteri, T. scriptus, P. porcus) and Primates (C. a. tantalus, C. mona,
C. e. erythrogaster, C. vellerosus, E. patas, P. verus). Those are mostly
habitat- specialised species with K- selected reproductive strategy
and dependent on large forest covers. As such, they remain the most
vulnerable to habitat degradation and hunting activities (Cowlishaw
et al., 2005). It is reasonable to suggest that a combination of habitat
degradation and reduced forested cover is the main factor explain-
ing the low density of medium- sized mammals in the forest islands
surrounding LNF (Bogoni et al., 2020; Tilker et al., 2019). Preferential
hunting for prized, medium- sized mammals which are the largest
mammals occurring in southern Benin could also have exacerbated
this tren d (P rice & Gi t tlema n, 20 07), bu t dat a to supp ort this hyp oth -
esis are currently lacking.
The ecology of the most abundant species sighted from the
forest islands tends to support a signature of habitat degradation
in those areas, which typically includes widespread species with
R- selected reproductive strategies (e.g. Rodentia, Lagomorpha,
Galliformes). Rodents, rabbits and francolins are known to success-
fully persist in agricultural landscapes subjected to hunting pressure
in the continental region of Equatorial Guinea (Kümpel et al., 2010).
The abundance of squamates such as B. arietans and V. niloticus in
degraded habitats could be expected as they also seem to success-
fully adapt to agricultural landscapes (Edgar et al., 2010).
Although the socioeconomic profile of tropical forest hunters
was shown to influence hunting efficiency (Nunes et al., 2020), we
could not detect any significant effect of socioeconomic character-
istics on the hunting pressure (number of individuals hunted across
all species within 1 year) around LNF. In the study area, all the active
hunters are from the same generation (between 20 and 40 years
old), and all were men and farmers. We found hunting pressure to be
greater in forest islands located at the vicinity of LNF than in forest
patches more distant from LNF. Because LNF has set up an efficient
anti- poaching strategy, poachers reported that they preferred to
operate in the nearby, unprotected forest islands that benefit from
the dispersal of the healthy populations preserved in LNF nearby.
FIGURE 4 Relationship between the numbers of individuals
hunted across all species within a year (on the x- axis) and the
probability of a small forest patch to be close to the Lama Natural
Forest as obtained from Bayesian regression
    DJAGO UN et Al.
Because of the link that we could establish between smaller forest
patches (islands) and low abundance of bushmeat species, popula-
tions from the forest islands close to LNF may critically be suffer-
ing from over- hunting. Small forest patches (islands) distributed in
a fragmented landscape can help maintaining functional diversity
and gene flow among populations (Magioli et al., 2016). In southern
Benin, hunting pressure on the remaining forest islands, where law
enforcement patrols do not operate, may be too high to enable such
a functional role. This brings to light the potential inappropriateness
of current conservation measures in the fragmented forest habitat
of southern Benin, as the effective conservation of the largest for-
est fragment (LNF) itself may not be sufficient to preser ve the areas
wildlife diversity in the long- term. The conservation role of forest
islands around LNF has likely been under- rated, and the abundance
of non- forest- dependent species suggests that ecosystem function-
ing may be highly disrupted (Lees & Peres, 2009; Michalski & Peres,
2007 for similar evidence in South America). Our results call for an
integrative approach to preserve the last fores ted habitat s of so uth-
ern Benin, where securing corridors to re- establish connectivity
among forest patches (Magioli et al., 2016; Michalski et al., 2008;
Prugh et al., 2008; Sreekar et al., 2015) should be a priorit y.
A substantial shift towards effective law enforcement, education
of rural communities and economic incentives to develop access to
alternative source s of protein are to be implemented in order to mit-
igate the effects of the bushmeat trade on forest wildlife in southern
Benin. There is also a need to quantify the scale of bushmeat hunting
activities, notably on the less abundant species as continued hunting
could lead to local extinctions.
This study was funded by the program Jeune Equipe Associée à l’IRD
(RADAR- BE). SZ was supported by a PhD grant ARTS- IRD. We are
grateful to all the participants for generously giving their time to
complete this survey. We thank Sean Heighton for improving the
English writing skill of this manuscript.
The co- authors have no conflict of interest to declare.
The data that support the findings of this study are available from
the corresponding author upon reasonable request.
Chabi A. M. S. Djagoun
Gilles Nago
Akomian F. Azihou
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How to cite this article: Djagoun, C. A. M. S., Nago, G.,
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Assogbadjo, A., Sinsin, B., & Gaubert, P. (2021). Assessing
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Questionnaire addressed to the rural communities
1. Village
2. Sex
3. Age (20– 40 years; 41– 60 years; >60 years)
4. Level of education (None; Primary level; At least Secondar y level)
5. Occupation (Employed or involved in agricultural activities;
1. What is your duration of stay in the village?
2. What is the size of your household?
3. How often do you see the bushmeat species within a year in the
4. May you list from the list that I am providing which species are
rare or abundant in the forest?
5. In your opinion, how many individuals were hunted per species
within a year?
6. Why are these wild animals hunted?
7. In your opinion, what are the main reasons for people to engage
into illegal hunting activities in the area?
8. Are there any strategies in place to minimize illegal hunting in the
are a?
9. Are there any other information/comments that you would like to
share with us regarding illegal hunting activities in the area?
Full-text available
Background Pangolins are trafficked in unsustainable volumes to feed both local and global trade networks for their meat and the medicinal properties of their derivatives, including scales. We focus on a West African country (Benin) to assess the medicinal and spiritual values of pangolins among different ethnic groups and identify the cohort of buyers involved in the pangolin trade and related economic values along the chain, notably from local diasporas. Methods We organised 54 focus groups in villages surrounding occurrence habitats of pangolins across Benin and conducted 35 individual interviews with vendors from five major traditional medicine markets (TMMs). Our questionnaire addressed the different uses of pangolins, the commercial value of pangolin items, the categories of clients and the related selling prices. Results Pangolin meat was strictly consumed as food. Scales, head, bones, tongue, blood, heart and xiphisternum were the items used by local communities as part of medicinal (65% of the focus groups) and spiritual (37%) practices. Scales were the most frequently used item (use value index = 1.56). A total of 42 medicinal and spiritual uses, covering 15 International Classification of Diseases (ICD) categories, were recorded among ethnic groups. The ICD and spiritual categories-based analyses of similarity showed a partial overlapping of ethnozoological knowledge across Benin, although knowledge was significantly influenced by ethnicity and geographic location. The pricing of pangolins both varied with the category of stakeholders (local communities vs. stakeholders of TMMs) and clients (local and West African clients vs. Chinese community) and the type of items sold. The Chinese community was reported to only buy pangolins alive, and average selling prices were 3–8 times higher than those to West African clients. Conclusions Our results confirm that pangolins in Africa are valuable and versatile resources for consumption and medicinal / spiritual practices. The pangolin trade in Benin is based on an endogenous and complex network of actors that now appears influenced by the specific, high-valued demand from the Chinese diaspora. Further investigations are required to assess the growing impact of the Chinese demand on the African wildlife trade.
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Neotropical mammal diversity is currently threatened by several chronic human-induced pressures. We compiled 1,029 contemporary mammal assemblages surveyed across the Neotropics to quantify the continental-scale extent and intensity of defaunation and understand their determinants based on environmental covariates. We calculated a local defaunation index for all assemblages—adjusted by a false-absence ratio—which was examined using structural equation models. We propose a hunting index based on socioenvironmental co-variables that either intensify or inhibit hunting, which we used as an additional predictor of defaunation. Mammal defaunation intensity across the Neotropics on average erased 56.5% of the local source fauna, with ungulates comprising the most ubiquitous losses. The extent of defaunation is widespread, but more incipient in hitherto relatively intact major biomes that are rapidly succumbing to encroaching deforestation frontiers. Assemblage-wide mammal body mass distribution was greatly reduced from a historical 95th-percentile of ~ 14 kg to only ~ 4 kg in modern assemblages. Defaunation and depletion of large-bodied species were primarily driven by hunting pressure and remaining habitat area. Our findings can inform guidelines to design transnational conservation policies to safeguard native vertebrates, and ensure that the “empty ecosystem” syndrome will be deterred from reaching much of the New World tropics.
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Although critical in understanding human societies relying on natural game stocks, little attention has been paid to how socioeconomic traits can influence hunter behaviour. Our research focuses on whether village size, household size and age, and hunter age and monetary income affect hunting efficiency (catch-per-unit-effort) and catchment areas of traditional Amazonians. In collaboration with 13 volunteer hunters from six villages, we assessed social traits, and identified hunting areas with GPS and animal kills over six months. Contrary to expectations from central-place foraging, hunters in larger villages used smaller catchment areas, potentially because cassava-associated game species are more common near larger villages. Older hunters were more efficient, emphasizing the role of experience gained through time. Catchment areas increased with hunters' income, but apparently the hunting efficiency did not. Overall results support the notion that a spatial arrangement of fewer large villages, rather than many small ones, maximizes hunting efficiency and minimizes catchment areas.
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Although pangolins are considered to be one of the most trafficked wild mammals in the world, their conservation status remains uncertain through most of their ranges, and notably in western Africa. Using local ecological knowledge in combination with direct occurrence evidence, we assessed the distribution and abundance of the white-bellied pangolin (Phataginus tricuspis) and the giant pangolin (Smutsia gigantea) in Benin over the last two decades (1998–2018). We organised focus groups with local hunters in 312 villages within sampling units of 25 km × 25 km covering the whole country. Participatory maps of past and current geographic ranges of pangolins combined with direct evidence (96 collected scales) suggested that the white-bellied pangolin had a wide distribution and reached higher latitudes than previously known, whereas the giant pangolin could be restricted to a single forest and has been absent from the northern part of the country over at least the last two decades. Local hunters perceived a significant decline of the white-bellied pangolin (range contraction = 31% in 20 years) and almost unanimously an extirpation of the giant pangolin (93%), the latter being restricted to the Alibori forest reserve (northern Benin). Protected areas were identified as the main potential occurrence zones for pangolins: 77 and 100% of the potential geographic ranges of the white-bellied and the giant pangolins, respectively, overlapped partially or totally with the protected area network of Benin. Using a generalized linear model, we showed that the probability of reporting recent observations of the white-bellied pangolin was significantly higher for villages near protected areas and distant from main roads. The forests perceived as having sustainable populations of white-bellied pangolins were the Lama Forest reserve (southern Benin) and the complex comprising Monts Kouffé - Wari Maro - Ouémé supérieur (central Benin), a view that was supported by the great number of direct occurrence evidence collected in those areas. Overall, our study highlights an important geographic range contraction of the white-bellied pangolin and a possible entire extirpation of the giant pangolin over the last two decades in Benin.
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African teak (Milicia excelsa (Welw.) C.C. Berg) is an endangered multi-use species. Understanding the impact of climate change on the distribution of this species may improve the ability to anticipate or recognize its decline or expansion and to take appropriate conservation measures if necessary. Ecological niche modeling was projected in geographical space to study the current and future distribution of M. excelsa in Bénin. MaxEnt was used to estimate the potential geographic distribution of the species under two Representative Concentration Pathways (RCP). Miroc 5 summaries and two RCP 4.5 and RCP 8.5 scenarios were used as predictor variables for projections of the geographic potential of this species. The performance of the model was assessed by the area under the curve (AUC), true skill statistics (TSS) and partial receiver operating characteristics (Partial ROC). From the results, M. excelsa was more a secondary species in the Guinean climatic zone and part of the Sudanian-Guinean and Sudanian climatic zone. The projections show a significant decrease in suitable habitats for the species from the two RCP scenarios. Only a part of the Guinean climatic zone remained suitable and few protected areas will conserve in situ M. excelsa. For the sustainable conservation of M. excelsa, it is essential to strengthen the protection of sacred forests located in the Guinean climatic zone.
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Habitat degradation and hunting have caused the widespread loss of larger vertebrate species (defaunation) from tropical biodiversity hotspots. However, these defaunation drivers impact vertebrate biodiversity in different ways and, therefore, require different conservation interventions. We conducted landscape-scale camera-trap surveys across six study sites in Southeast Asia to assess how moderate degradation and intensive, indiscriminate hunting differentially impact tropical terrestrial mammals and birds. We found that functional extinction rates were higher in hunted compared to degraded sites. Species found in both sites had lower occupancies in the hunted sites. Canopy closure was the main predictor of occurrence in the degraded sites, while village density primarily influenced occurrence in the hunted sites. Our findings suggest that intensive, indiscriminate hunting may be a more immediate threat than moderate habitat degradation for tropical faunal communities, and that conservation stakeholders should focus as much on overhunting as on habitat conservation to address the defaunation crisis.
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Aim Understanding how habitat loss and overhunting impact large carnivores is important for broad‐scale conservation planning. We aimed to assess how these threats interacted to affect jaguar habitat (Panthera onca) between 1985–2013 in the Gran Chaco, a deforestation hotspot. Location Gran Chaco ecoregion in Argentina, Paraguay and Bolivia. Methods We modelled jaguar habitat change from 1985–2013 using a time‐calibrated species distribution model that uses all occurrence data available for that period. We modelled habitat as a function of resource availability and hunting threats, which allowed us to separate core (high resource availability and low hunting threat), refuge (low resources but safe), attractive sink (high resources but risky) and sink (low resources and risky) habitat for 1985, 2000 and 2013. Results Jaguar core areas contracted by 33% (82,400 km²) from 1985–2013, mainly due to an expansion of hunting threats. Sink and attractive sink habitat covered 58% of the jaguar range in 2013 and most confirmed jaguar kill sites occurred in these areas. Furthermore, habitat loss and hunting threats co‐occurred in 29% of jaguars’ range in 2013. Hunting threats also deteriorated core areas within protected areas, but 95% of all core areas loss occurred outside protected lands. About 68% of the remaining core areas in 2013 remained unprotected, mostly close to international borders. Main conclusions Our study highlights the synergistic effects that habitat loss and hunting threats exert on large carnivores, even inside protected areas, emphasizing the need to consider the geography of threats in conservation planning. Our results also point to the importance of areas along international borders as havens for wildlife and thus the urgent need for cross‐border planning to prevent the imminent extinction of jaguars from the Chaco. Opportunities lie in reducing jaguar mortality over the widespread attractive sinks, particularly in corridors connecting core areas.
Several hundred species are hunted for wild meat in the tropics, supporting the diets, customs, and livelihoods of millions of people. However, unsustainable hunting is one of the most urgent threats to wildlife and ecosystems worldwide and has serious ramifications for people whose subsistence and income are tied to wild meat. Over the past 18 years, although research efforts have increased, scientific knowledge has largely not translated into action. One major barrier to progress has been insufficient monitoring and evaluation, meaning that the effectiveness of interventions cannot be ascertained. Emerging issues include the difficulty of designing regulatory frameworks that disentangle the different purposes of hunting, the large scale of urban consumption, and the implications of wild meat consumption for human health. To address these intractable challenges, we propose eight new recommendations for research and action for sustainable wild meat use, which would support the achievement of the United Nations Sustainable Development Goals. Expected final online publication date for the Annual Review of Environment and Resources, Volume 46 is October 2021. Please see for revised estimates.