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Ecology and ethnozoology of the three-cusped pangolin Manis tricuspis (Mammalia, Pholidota) in the Lama forest reserve, Benin

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The conservation of biodiversity and sustainable man-agement strategies must be based on an understanding of the structural and functional ecological traits of utilized species. Such basic data are still lacking for even the most endangered species in Africa. In this study, we combined ecological methods with ethnographic appro-aches to investigate the ecology and ethnozoology of three-cusped pangolin (Manis tricuspis) in the context of the Lama forest reserve (6855.89–58.89 N and 284.29–10.89 E) in Benin. We interviewed 79 people from various socio-professional groups to assess how important is three-cusped pangolin to indigenous people. We sur-veyed 15 strip transects (3 km=1 km) within 12 sites to seek for indices of pangolin presence and record-asso-ciated vegetation data in six plots of 900 m 2 (30 m=30 m). In total, 38 pangolins were observed in the forest. The density was 0.84 pangolins/km 2 during the dry season and the number of observations did not differ significantly between plantations and natural forest. The age ratio of juvenile/adult was 1:1. The preferred habitat was colonized by termite mounds in 62% of the sites. Pangolins were mostly found in holes of Dialium gui-neense and Ceiba pentandra in the closed natural for-ests. The distribution of pangolins in the Lama forest reserve suggests that the species is more sensitive to forest age than to its composition. Three-cusped pan-golin plays an important role as food, medicine, mythic and source of income for local communities around the Lama forest reserve. We also recommend future research guidelines important for the conservation of Manis tri-cuspis in Benin.
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Mammalia 72 (2008): 198–202 2008 by Walter de Gruyter Berlin New York. DOI 10.1515.MAMM.2008.046
Article in press - uncorrected proof
2008/46
Ecology and ethnozoology of the three-cusped pangolin
Manis tricuspis (Mammalia, Pholidota) in the Lama forest
reserve, Benin
Hugues A. Akpona*, Chabi A.M.S. Djagoun and
Brice Sinsin
Laboratoire d’Ecologie Applique´ e, Faculte´ des Sciences
Agronomiques, Universite´ d’Abomey-Calavi, 01 BP 526
Cotonou, Be´ nin, e-mail: akpona@gmail.com
*Corresponding author
Abstract
The conservation of biodiversity and sustainable man-
agement strategies must be based on an understanding
of the structural and functional ecological traits of utilized
species. Such basic data are still lacking for even the
most endangered species in Africa. In this study, we
combined ecological methods with ethnographic appro-
aches to investigate the ecology and ethnozoology of
three-cusped pangolin (Manis tricuspis) in the context of
the Lama forest reserve (6855.89–58.89N and 284.29–10.89
E) in Benin. We interviewed 79 people from various
socio-professional groups to assess how important is
three-cusped pangolin to indigenous people. We sur-
veyed 15 strip transects (3 km=1 km) within 12 sites to
seek for indices of pangolin presence and record-asso-
ciated vegetation data in six plots of 900 m
2
(30 m=30 m). In total, 38 pangolins were observed in the
forest. The density was 0.84 pangolins/km
2
during the
dry season and the number of observations did not differ
significantly between plantations and natural forest. The
age ratio of juvenile/adult was 1:1. The preferred habitat
was colonized by termite mounds in 62% of the sites.
Pangolins were mostly found in holes of Dialium gui-
neense and Ceiba pentandra in the closed natural for-
ests. The distribution of pangolins in the Lama forest
reserve suggests that the species is more sensitive to
forest age than to its composition. Three-cusped pan-
golin plays an important role as food, medicine, mythic
and source of income for local communities around the
Lama forest reserve. We also recommend future research
guidelines important for the conservation of Manis tri-
cuspis in Benin.
Keywords: Benin; ecology; ethnozoology; Manis
tricuspis.
Introduction
Wildlife conservation actions in Africa have been mostly
hindered by the lack of ecological data, even for the most
endangered species. Pangolins are largely nocturnal and
have adapted to a highly specialized diet of ants and
termites (Lekagul and McNeely 1988, Lim and Ng 2007).
Three-cusped pangolin (Manis tricuspis, hereafter
referred to as Pangolin) is one of the eight species of
pangolin and is native to Africa, distributed from Senegal
to western Kenya, and from southern Africa to Zambia
(Wilson and Reeder 1993). Despite its relatively wide dis-
tribution, recent data on the ecology, distribution and
conservation status of the species is rare (Coe 1975,
Rahm and Christiaensen 1996). Pangolin is reported in
few places in Africa, such as the eastern Zaire (Rahm and
Christiaensen 1966) and Mount Nimba in Liberia (Coe
1975). Most of the existing literature on Pangolin ecology
is reported by Page´ s (1975) and Stuart and Stuart (1997).
Pangolin meat is popular in West Africa (Angelici et al.
1999), where it is widely hunted (Sodeinde and Adedipe
1994). As a consequence, there is a decline in Pangolin
populations, even though commercial trade of all pan-
golin species is banned (Pangolin Specialist Group 1996).
Pangolin do not survive well in captivity, with a mortality
rate up to 71% in the first year of captivity (Wilson 1994),
and are believed to give birth to a single offspring each
year (Page´ s 1975, Kingdon 1997). Conservation and sus-
tainable management plans of a species must consider
both ecological constraints and human impacts on the
species. In this study, we documented knowledge of the
ecology and ethnozoology of the three-cusped pangolin
in the Lama forest reserve, which is located in the Daho-
mey Gap, an abrupt climatically induced rainforest frag-
mentation in West Africa during the late Holocene.
Specifically, (1) we tested the effect of vegetation struc-
ture and type on the abundance of the species, and (2)
assessed the importance of the species for local popu-
lations around the reserve.
Methods
Study area
This study was conducted in the Lama forest reserve
(6855.89–58.89N and 284.29–10.89E), the largest remnant
natural forest in Southern Benin. The forest is at one of
the lowest altitudes of the country ranging between 40 m
and 80 m. The climate is equatorial with two rainy sea-
sons (April to mid-July and mid-September to October).
The mean annual rainfall is above 1200 mm, the mean
temperature varies between 258C and 298C and the rel-
ative humidity between 69% and 97%. The prevailing soil
type is a black cotton soil (vertisol) rich in humus and
recent clay deposits.
The Lama forest reserve is subdivided in three main
zones: the natural forest (NF), also known as Noyau cen-
H.A. Akpona et al.: Ecology and ethnozoology of the three-cusped pangolin Manis tricuspis 199
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Figure 1 Outline map of the Lama forest reserve.
tral which is strictly protected from hunting; the teak
plantations (FP); and the bordering zone (BZ), where local
populations are allowed to settle and harvest non-timber
forest products from the forest (Figure 1). The NF repre-
sents 31% of the forest reserve and is made of a mosaic
of fallows of different age and successional stages invad-
ed in some places with neotropical pioneer Chromolaena
odorata, secondary forest and patches of primary forest.
It is surrounded by teak plantations established in the
early 1960s (1963–1965) and in 1985–1996 (Nagel et al.
2004).
Wildlife in the Lama forest reserve is represented by
threatened mammals: the mona (Cercopithecus mona),
the red-bellied monkey (Cercopithecus erythrogaster),
which is endemic to Benin (Nobime 2002), the vervet
monkey (Cercopithecus aethiops), the sitatunga (Trage-
laphus spekei), the royal antelope (Neotragus pygmaeus),
the black duiker (Cephalophus niger), the yellow-backed
duiker (Cephalophus silvicultor) and the Kitampo rope
squirrel (Funisciurus substriatus), an endemic species of
the Dahomey Gap (Refisch 1998, Sinsin and Assogbadjo
2002, Kassa and Sinsin 2003, Nagel et al. 2004).
Study design and data collection
Pangolin survey and habitat characterization We
collected data in the NF and FP. In total, 12 sites with
known pangolin presence were identified and 15 strip
transects (3 km=1 km) were surveyed. We surveyed pan-
golin both in the NF and in the FP using a point transect
method combined with opportunistic presence research.
The point transect method is recommended for closed
forested areas and has been successfully used in previ-
ous surveys in the Lama forest reserve (Kassa and Sinsin
2003). For each encounter of a pangolin, we recorded
the hour of observation, number of individuals, group
composition, age (yearling, sub-adult or adult), sex, tree
species that the pangolin was on and habitat character-
istics (i.e., vegetation type, vegetation cover: five classes
of 0–100%, presence or absence of termite mounds, ant
nests, tree termite mounds, holes in trees). To describe
the specific habitat of pangolin which is known to be an
arboreal species (Page´ s 1975), we installed six vegeta-
tion plots (three in FP and three in the NF) of 900 m
2
each
(30 m=30 m) to inventory the tree species.
Importance of the three-cusped pangolins for local
communities We conducted structured interviews
and focus group discussions in two local markets (Bohi-
con and Abomey) and five bordering villages (Agadjalig-
bo, Akpe´ , Gue´me´ , Massi and Zalimey). We interviewed
24 local people living in the forest border, 11 forest
guards, eight local hunters, 26 traditional healers and
skins sellers. Interviewees were selected according to
their ethnic groups (65% Holli and 35% Fon), socio-pro-
fessional groups, communities and position in relation to
the reserve. Questions asked were related to the different
uses (medicinal, food, ritual, etc.) of the species, the local
nomenclature according to different ethnic groups, the
socio-economic and the cultural role of the species, the
consumption preferences, the appreciation of the meat,
the reasons which motivate the hunting of pangolins and
the form of commercialization (alive, dead).
Results
Ecology and habitat of the three-cusped pangolin
in the Lama forest reserve
We observed a total of 38 pangolins (28 on trees and 10
on the ground; Figure 2). In total, 70% of the population
was observed in the wild forest (NF) and the remaining
was found in the plantations, specifically in relic island
forests or in old teak plantations. The number of pan-
golins observed in the reserve did not differ significantly
between plantations and natural forest (t-test, ps0.0671,
)0.05). During the data collection, anthills were found on
all sites where pangolins were observed (100%) and
termite mounds were found in 62% of the sites.
The average pangolin density was estimated at
0.84 pangolins/km
2
(38 individuals in 45 km
2
) during the
dry season. We observed a predominance of juveniles
and yearlings (39% and 23%, respectively; Figure 3) in
the pangolin populations. Only two adults were observed
in plantations, whereas 12 adults were recorded in the
wild forest. However, there were no significant differenc-
es in juveniles and yearlings between plantations and
wild forest (8 vs. 7 and 5 vs. 4, respectively).
The vegetation cover of the pangolin habitat varied
between 20% and 70% and sightings of the species
200 H.A. Akpona et al.: Ecology and ethnozoology of the three-cusped pangolin Manis tricuspis
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Figure 2 A three-cusped pangolin in the Lama forest reserve.
Figure 3 Age class of three-cusped pangolins in the different
forest types of the Lama forest reserve.
Figure 4 Tree species used as a support by Manis tricuspis in
the Lama forest reserve.
Figure 5 Human consumption rate of Manis tricuspis in the
Lama forest reserve.
were more abundant in closed areas (wild forest, forest
islands and old teak plantations), where 25 pangolins
were observed. Pangolins were found in tree holes of
species, such as Dialium guineense and Ceiba pentandra
(Figure 4). The most abundant tree species found in the
habitat of Manis tricuspis were Mimusops andogensis,
Dialium guineensis,Celtis integrifolia,Lonchocarpus seri-
ceus,Drypetes floribunda,Diospyros mespiliformis,Pou-
chetia africana,Ficus congensis,Malacantha alnifolia,
Cola gigantea,Terminalia superba,Lecaniodiscus cupa-
nioides,Holarrhena floribunda,Gmelina arborea,Antiaris
toxicaria,Mallotus oppositifolius and Milicia excelsa,
Dialium guineense and Ceiba pentandra.
Harvesting pangolins in the forest
Three-cusped pangolin is not considered as a divinity for
local people around the Lama forest reserve. Like all oth-
er small mammal species, three-cusped pangolins are
not protected by the legislation in Benin. Despite the
work being carried out by the ‘‘Office National du Bois
(ONAB)’’ to protect the reserve, illegal poachers still oper-
ate in the forest. Moreover, plantations where the species
occurs are not strictly protected. Traditional hunting took
place during the night in plantations, farms and natural
forest. Hunters used very basic materials: sticks, flash-
lights and bags for contention after capture. It is note-
worthy that hunters preferred yearlings and adult
pangolins for consumption.
The majority of pangolin hunters are farmers who hunt-
ed both for subsistence, medicinal and commercial pur-
poses. Three-cusped pangolin is known as ‘‘Iwo’’ by Holli
tribes and as ‘‘Akpakedje’’ or ‘‘Lihoui’’ by Fon tribes.
More than 78% of populations interviewed in each village
strongly appreciated three-cusped pangolin meat (Figure
5). Pangolins were sold on the main roads and the prices
varied from one place to another. For example, a sub-
adult or adult pangolin cost approximately US$4 (2000)
FCFA) in villages and US$8 (4000 FCFA) in city markets.
Three-cusped pangolins were sold alive or dead to tra-
dipracticians, healers, travelers and local populations of
the reserve. Tegon, a village of Zogbodomey district is
known as reference for the sale of this bushmeat. Some
organs, such as skin, heart, intestine and head, are used
for medicinal purposes to treat asthma, cardiovascular
and dermatologic diseases (Table 1).
Discussion
Ecology of the three-cusped pangolin in the
Lama forest reserve
Our results show that pangolins were observed more fre-
quently on trees than on the ground in the Lama forest
reserve. We expected this result considering that the
three-cusped pangolin often sleeps in trees (Page` s 1975,
Kingdon 1977). We expected a higher proportion of pan-
H.A. Akpona et al.: Ecology and ethnozoology of the three-cusped pangolin Manis tricuspis 201
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Table 1 Use of Manis tricuspis organs for medicine or ritual purposes around the Lama forest reserve.
Organs Diseases treated or virtues Quotation (%)
Skin Dermatosis 20
Heart Accelerate heart banging 38
Feet Normal growth and baby’s vigor 34
Intestine Digestive trouble 40
Tongue Asthma 16
Head Fever, hemorrhoid, introduction in wizard groups 48
Scale Prevention of accidents, invulnerability to rifle or stabs 60
golins in the natural forest than in the plantations,
because the natural forest is a well protected area made
up of a mosaic of secondary forest and patches of pri-
mary forest. However, our results did not support this
hypothesis in the Lama forest reserve. Instead, we found
that old teak plantations may offer similar or tolerable
conditions for the survival of the species with the natural
forest. The plantations border the natural forest in the
Lama forest reserve and they may serve as a buffer zone
that reduces the risk of illegal logging in the natural forest
(Nagel et al. 2004). In addition, plantations may also play
an important direct role for the conservation of biodiver-
sity in Lama forest reserve. Specifically, old plantations
(approximately 40 years old) may provide suitable habi-
tats even for rainforests, insects and extremely rare spe-
cies. For example, extremely rare species were found in
these plantations, as well as in isolated forest islands
(Attignon 2005).
The presence of termite mounds and ant hills within
the preferred habitat of pangolins in the Lama forest
reserve is due to their feeding preferences. Page´ s (1975)
reported that the exploitation of home range by Manis
tricuspis consists of terrestrial and arboreal displace-
ments in the search of food, the animal being largely noc-
turnal and adapted to a highly specialized diet made of
ants and termites (Lekagul and McNeely 1988). The dis-
tribution of three-cusped pangolins in the Lama forest
reserve could be discussed in relation to the exploitation
of home range, the hunting technique used by the spe-
cies and the protection status of each forest type. The
three-cusped pangolin foraged by exploring through the
termite mounds successively and not exhaustively
(Page´ s 1975). Research in Gabon, e.g., shows that the
abundance of three-cusped pangolin adult males is rel-
atively lower, and home ranges larger but overlapping
with the home ranges of several females. The females
only have small overlapping home ranges, and they often
overlap only at the edges (Page´ s 1975).
We found that adults were observed more in natural
forest and suggest that adults may take more precau-
tions than juveniles and yearlings by defining their home
range essentially in the natural forest which has a pro-
tected status. The relations between juveniles and year-
lings are relatively peaceful and the ranges of juveniles
have an amoeboid form with changing boundaries,
whereas yearlings are errant, change resting holes every
night and wander over large areas (Page´s 1975). This jus-
tifies the distribution of juveniles and yearlings both in
natural forest and in plantations. Species richness of ter-
mites was significantly higher in natural forest than in the
teak plantations, and the contrary was true for the mean
number of termite encounters per transect (Attignon
2005). However, very little is known on the prey prefer-
ences of Manis tricuspis in the reserve and it is not pos-
sible for us to state if there is a greater abundance of
prey preferred by the animal in the natural forest or in the
teak plantations. Our study recommends future investi-
gations on the dietary habits of M. tricuspis and exami-
nation of the relation between the availability of specific
ant or termite species in the different forest types and the
abundance and behavior of pangolins. Our estimate of
0.84 pangolins/km
2
in the Lama forest reserve is probably
under-estimated, given that we could not efficiently carry
out a census of pangolins at night. Pangolins are noc-
turnal and the periods of activity during which it is easier
to observe individuals may differ according to age and
sex (Page´ s 1975). The point-transect census method we
used supposes that no subject of interest escapes from
the observer. Therefore, we might have failed to observe
some individuals. The under-estimation of pangolins in
the Lama forest reserve is not as important as under-
standing the trend of the population over time using the
same method. There is a need to establish a monitoring
system of the species in this ecosystem.
Habitat characteristics
Habitat analysis showed that three-cusped pangolin pre-
ferred closed forest habitats (natural forest, forest islands
and old teak plantations). In other parts of Africa, Manis
tricuspis inhabits forests and gallery forests, but can also
survive in cultivated areas and forest mosaics (Haltenorth
and Diller 1980, Ansell 1982, Sodeinde and Adedipe
1994, Kingdon 1997). In the Lama forest reserve, three-
cusped pangolins were found dwelling in holes of tree
species, such as Dialium guineense and Ceiba pentan-
dra. The high representation of Dialium guineense and
Diospyros mespiliformis could be due to their higher den-
sity in the reserve (17 and 22 individuals/ha, respectively)
(Codjia et al. 2003). We suggest further investigations to
understand how the species chooses trees species.
Importance and sustainable use of Manis tricuspis
Wild meat harvests in African moist forests are presumed
to exceed production, even in the case of traditionalsoci-
eties still using rudimentary hunting methods (Fa et al.
2006). To understand the contribution of wildlife in the
subsistence of local populations, one should not consid-
er only the big game. Most of the meat consumed by
forest bordering populations in the Lama forest reserve
comes from small mammals, which can be captured at
any time of the year (Codija and Assogbadjo 2004,
202 H.A. Akpona et al.: Ecology and ethnozoology of the three-cusped pangolin Manis tricuspis
Article in press - uncorrected proof
Assogbadjo et al. 2005). Pangolin is considered a valu-
able bushmeat in West Africa (Angelici et al. 1999). In
Benin, it is an important source of protein, widely con-
sumed in both rural and urban areas around the Lama
forest reserve. Our study shows that more than 78% of
populations interviewed in each village around the Lama
forest reserve strongly appreciated three-cusped pan-
golin meat. Elsewhere, despite being officially listed as
endangered in Nigeria, pangolins are still hunted in Ogun
State, where deforestation has fragmented and reduced
their forest habitat; the species is becoming rare in the
region (Sodeinde and Adedipe 1994).
We found that organs of pangolins are used to treat
asthma, cardiovascular and dermatologic diseases. The
use of pangolins’ organs in traditional medicine is also
reported elsewhere. Pangolin scales are reported to be
used as antiseptics and to combat fever and skin disease
in Chinese culture. Scales are either used as is or ground
into a powder for use in potions, which are thought to
aid in treating venereal diseases. Pangolin skin is also
used for making boots and other leather goods (Fahey
1999).
The conservation and sustainable use of pangolins in
the Lama forest reserve require monitoring of the capture
and the commercialization rates. The species has a lower
litter and our study suggests that it is easy to hunt using
cheap and very basic materials. The hunting rate is prob-
ably higher than we might estimate. IUCN (2007) classi-
fied the tree pangolins as ‘Lower Risk’ but suggests that
the species needs updating. Manis tricuspis might be
more endangered in Benin and in many other parts of
West Africa than is reported.
Acknowledgements
This study was conducted with the financial support of the Brit-
ish Ecological Society (BES) Overseas Bursary No. 904/1128.
We are also grateful to the ‘‘Office National du Bois’’ for logistic
support, to ‘‘Beyond the School’’ (BSc) association for orienta-
tions in research. We acknowledge Aubin Yolou, Guy Degue-
nonvo, Sylvain Bankole, Liamidi Konetche and Janvier Agbetou
for their field assistance, and Orou Gaoue, Norman Lim and
Gaubert Philippe for their comments on an earlier version of this
paper.
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... In the Dahomey Gap, the white-bellied pangolin (Phataginus tricuspis) and giant pangolin (Smutsia gigantea) have been previously reported (Sayer & Green 1984;Akpona et al. 2008;Akpona & Daouda 2011;Nixon et al. 2019;Pietersen et al. 2019). Benin remains the only country in the Dahomey Gap where the giant pangolin has been reported but without any direct or recent evidence (Sayer & Green 1984;Akpona & Daouda 2011;Amori et al. 2016;Nixon et al. 2019). ...
... Effective and modern wildlife conservation implementation requires informed decision-making (Rose et al. 2018;Ausden & Walsh 2020). In the Dahomey Gap, knowledge on pangolins was extremely poor with only one study on ecology and ethnozoology of the white-bellied pangolin in southern Benin (Akpona et al. 2008). The paucity of scientific knowledge makes challenging the conservation decision-making in the Dahomey Gap. ...
... The paucity of scientific knowledge makes challenging the conservation decision-making in the Dahomey Gap. Apart from the pioneer study in Lama forest reserve (Akpona et al. 2008), no participatory management plans, dissertation thesis and scientific papers did document the presence of pangolins in Benin and a similar gap of knowledge was observed in Togo before a very recent investigation on the distribution of the white-bellied pangolin (Segniagbeto et al. 2021 (Wu et al. 2005;Lim 2007;Irshad et al. 2015;Lee et al. 2017;Ashokkumar et al. 2017;Willcox et al. 2019;Sun et al. 2020;Karawita et al. 2020) and the Temminck's ground pangolin (Pietersen et al. 2016b) has been investigated using direct and indirect observations, none of these studies has reached a high taxonomic resolution identification of consumed preys. This data is essential to understand the role of pangolins in their wild habitats and for managed breeding programs. ...
Thesis
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Pangolins are on the top of conservation priority due mainly to their unprecedented level of harvesting. The four African species are under great gap of baseline data that hampers the set up of a well-designed, evidence-based strategy for tackling their alarming conservation status. My PhD dissertation focused on the biogeographically delineated Dahomey Gap region (West Africa) using the white-bellied pangolin as a case study to: (i) assess the geographic distribution and population trends of pangolin species across Benin, (ii) Model pangolin population extirpation rates using validated local ecological knowledge, (iii) assess the ethnozoological and commercial values of pangolins in Benin, (iv) investigate the population genetics of the white-bellied pangolin and set up the molecular tracing of their fine scale trade across the Dahomey Gap, and (v) assess the precise dietary composition of the white-bellied pangolin in Benin through eDNA metabarcoding. Local Ecological Knowledge revealed a 31 and 93 % contraction of the occurrence areas of the white-bellied and giant pangolins respectively, and highlighted habitat degradation and overexploitation as the main drivers of population decline for both species. Generalized linear model revealed that, abundance in 1998, distances from main roads and protected areas to villages, land use changes and deforestation over time were factors that significantly explained the current persistence of pangolins in Benin. Prediction models suggested decreasing trends for the white-bellied pangolin over the next two decades and a total extirpation of the giant pangolin whatever the scenario considered (deforestation versus no deforestation). Pangolins are valuable animals for local people and it occurs a regional trade network that involved Chinese diasporas. Pangolins in rural areas and traditional medicine markets are relatively cheaper in Benin compared to other countries in West and central Africa, but their price is 3-8 times higher when clients are from the Chinese diaspora. Microsatellites and mitochondrial DNA-based analyses suggested low genetic diversity, inbred populations, absence of clear geographic subdivision. Phylogenetic analysis suggested an endemic trade within the Dahomey Gap affecting the Dahomey Gap lineage of white-bellied pangolins and microsatellite markers pinpointed a long-distance trade within and between countries in the Dahomey Gap. The white-bellied pangolin preys mainly on ants and termites but also other insect taxa. The diversity of detected prey items was significantly higher (p<0.001) for the gut content (versus scat) samples. In conclusion, this dissertation revealed a high concern status of pangolins in the Dahomey Gap and provided substantial data that could be used as decision tools for the effective conservation of pangolins in the Dahomey Gap.
... Although the ethnozoology of pangolins has received much attention in neighbouring countries [6,17,[20][21][22], the situation in Benin, where the whitebellied pangolin's range has been contracted by 1/3 over the last two decades [23], remains understudied. Akpona et al. [24] found seven different items of pangolins used by southern communities for 13 medicinal and spiritual purposes, with scales as the most frequently cited. However, this study was restricted to southern Benin and included only two ethnic groups (Hôli and Fon), despite the larger extent of pangolin's distribution in Benin [23]. ...
... Relative to the sole study that had been conducted on the ethnozoology of pangolins in southern Benin [24], our investigations provide a deeper understanding of ethnozoological values across a diversity of ethnic groups in combination with the economic incentives possibly motivating the overexploitation of pangolins in Benin [23]. ...
... Pangolins are used for food, medicinal and spiritual purposes in Benin, in line with the literature record for tropical Africa [21,22,[36][37][38][39][40][41]. More specifically, pangolins in Benin constitute at the same time a bushmeat resource and an important input to traditional medicine and cultural practices [24]. Pangolin meat is unanimously consumed as food in Benin, whereas no medicinal and spiritual use is recorded, in line with Akpona et al. [24] but contrary to Boakye et al. [21,37] and Baiyewu et al. [36] who found that meat was used in traditional medicine in several countries of western and southern Africa. ...
Article
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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.
... The species seems to heavily rely on forest cover and old trees for its nocturnal activities [76], exploring its home range up to 1.8 km per night [77]. In the Dahomey Gap, WBP may occur in disturbed habitats including commercial plantations of teaks and palm trees, fallows and farmlands [78]. However, evidence of long-range dispersal is lacking and the general absence of structural connectivity among the remnant forest islands of the Dahomey Gap, especially in Benin (see [79,80]) does not support such a scenario. ...
Article
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Background: African pangolins are currently experiencing unprecedented levels of harvesting, feeding both local demands and the illegal international trade. So far, the lack of knowledge on the population genetics of African pangolins has hampered any attempts at assessing their demographic status and tracing their trade at the local scale. We conducted a pioneer study on the genetic tracing of the African pangolin trade in the Dahomey Gap (DG). We sequenced and genotyped 189 white-bellied pangolins from 18 forests and 12 wildlife markets using one mitochondrial fragment and 20 microsatellite loci. Results: Tree-based assignment procedure showed that the pangolin trade is endemic to the DG region, as it was strictly fed by the the Dahomey Gap lineage (DGL). DGL populations were characterized by low levels of genetic diversity, an overall absence of equilibrium, important inbreeding levels, and lack of geographic structure. We identified a 92-98% decline in DGL effective population size 200-500 ya-concomitant with major political transformations along the 'Slave Coast'-leading to contemporaneous estimates being inferior to minimum viable population size (< 500). Genetic tracing suggested that wildlife markets from the DG sourced pangolins through the entire DGL range. Our loci provided the necessary power to distinguish among all the genotyped pangolins, tracing the dispatch of a same individual on the markets and within local communities. We developed an approach combining rarefaction analysis of private allele frequencies with cross-validation of observed data that traced five traded pangolins to their forest origin, c. 200-300 km away from the markets. Conclusions: Although the genetic toolkit that we designed from traditional markers can prove helpful to trace the illegal trade in pangolins, our tracing ability was limited by the lack of population structure within the DGL. Given the deleterious combination of genetic, demographic, and trade-related factors affecting DGL populations, the conservation status of white-bellied pangolins in the DG should be urgently re-evaluated.
... Currently, some species like the White-bellied Pangolin suffer unsustainable harvest rates given the already low population densities. The population density of the White-bellied Pangolin ranges between 0.68 and 0.84 individuals/ km 2 (Cameroon and Benin) with a harvest rate of 30 per year by a single hunter in the province of Uíge (Akpona et al. 2008;Bobo et al. 2014). The decline of mammal biodiversity in the province of Uíge is significant and unique species like the White-bellied Pangolin and the endemic Southern Talapoin are likely to disappear in the near future. ...
Article
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Over-exploitation of wildlife especially bushmeat trade is the second most important threat to animal biodiversity. This also applies to Northern Angola but data on bushmeat and hunting techniques for this region are rare. Therefore, we study the most common hunting techniques, frequently captured species, and their economic value, and discuss the local resource use in relation to Angolan law and urgent global crises like the loss of biodiversity, the food supply in South African countries, and the risk of zoonoses. We recorded bushmeat hunting in 27 localities in the province of Uíge, accompanied hunters along their snare lines and interviewed additional 20 locals. Seven main types of snares and traps and their characteristics were defined. Hunters own on average 92 ± 128.7 snares and traps and capture about 25.3 ± 23.6 animals monthly. In total, respondents recognized 28 species of mammals of which one is considered as extinct and two as very rare. The majority of recorded species are hunted regularly. Rodents are most commonly caught followed by primates and duikers. Harvesting rates decrease with species' body size, leading to high economic value of and achievable prices for rare, large animals. Overall, our results document the hunting pressure on mammals and the persisting popularity of bushmeat in Northern Angola which poses an imminent threat to remaining mammal populations. Moreover, it endangers ecosystem integrity, rural livelihoods, and human health through the risk of new zoonoses. Our findings underscore the urgent need for sustainable solutions. The Angolan government should play a more active role in enforcing existing hunting legislation to reduce illegal bushmeat trade. Supplementary information: The online version contains supplementary material available at 10.1007/s10344-021-01541-y.
... The species seems to heavily rely on forest cover and old trees for its nocturnal activities [76], exploring its home range up to 1.8 km per night [77]. In the Dahomey Gap, WBP may occur in disturbed habitats including commercial plantations of teaks and palm trees, fallows and farmlands [78]. However, evidence of long-range dispersal is lacking and the general absence of structural connectivity among the remnant forest islands of the Dahomey Gap, especially in Benin (see [79,80]) does not support such a scenario. ...
Preprint
Full-text available
We conducted in the Dahomey Gap (DG) a pioneer study on the genetic tracing of the African pangolin trade. We sequenced and genotyped 189 white-bellied pangolins from 18 forests and 12 wildlife markets using one mitochondrial fragment and 20 microsatellites loci. Tree-based assignment procedure showed the ‘endemicity’ of the pangolin trade, as strictly fed by the lineage endemic to the DG (DGL). DGL populations were characterized by low levels of genetic diversity, an overall absence of equilibrium, inbreeding depression and lack of geographic structure. We identified a 92-98% decline in DGL effective population size 200-500 ya –concomitant with major political transformations along the ‘Slave Coast’– leading to contemporaneous estimates inferior to minimum viable population size. Genetic tracing suggested that wildlife markets from the DG sourced through the entire DGL range. Our loci provided the necessary power to distinguish among all the genotyped pangolins, tracing the dispatch of same individuals on the markets and within local communities. We developed an approach combining rarefaction analysis of private allele frequencies and cross-validation with observed data that could trace five traded pangolins to their forest origin, c. 200-300 km away from the markets. Although the genetic toolkit that we designed from traditional markers can prove helpful to trace the pangolin trade, our tracing ability was limited by the lack of population structure within DGL. Given the deleterious combination of genetic, demographic and trade-related factors affecting DGL populations, the conservation status of white-bellied pangolins in the DG should be urgently re-evaluated.
... The international trade in pangolins was banned when they were transferred to Appendix I of the Convention on International Trade of Endangered Species of Wild Fauna and Flora (CITES;, and all African pangolins are classified as threatened on the IUCN Red List (International Union of Conservation Nature [IUCN], 2021). In terms of local use within African countries, research has shown that in West Africa, nearly all parts of the animal have been reported to be used in African traditional medicine, purported to treat a plethora of different spiritual and physical ailments including rheumatism, convulsions, asthma, and cardiovascular and dermatological problems (Akpona et al., 2008;Boakye et al., 2014;2015;D'Cruze et al., 2020;Djagoun et al., 2012;Soewu & Adekanola, 2011;Soewu & Ayodele, 2009). Whilst research efforts on pangolins have increased, there is still limited information on pangolin ecology and local uses of pangolins, particularly in Central Africa. ...
Article
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Information about the presence and population status of pangolins, and the threats they face, remains limited in many parts of Cameroon, a country that is home to three species of pangolin and considered to be a global hub of pangolin trafficking. Local communities living in rural areas can provide valuable information on species presence, local uses of wildlife, and possible threats, that is useful for prioritising conservation actions. Using interview surveys in 20 villages surrounding Mbam and Djerem National Park, we investigated local peoples’ knowledge of pangolin presence, perceptions of population trends, cultural importance, consumptive and non-consumptive uses, and hunting of pangolins. Our results showed that most people recognised the white-bellied and giant pangolins, but only 10% recognised the black-bellied pangolin. Ethnolinguistic group significantly affected the likelihood of respondents recognising and having seen a pangolin before. Giant pangolin populations were perceived to be declining, particularly by older respondents. We found evidence of local use of pangolins for meat, but few respondents reported uses of scales. Cultural significance was reported by few respondents, but when it was reported it mostly referred to giant pangolin. White-bellied pangolins are reportedly hunted using bare hands for local consumption most frequently, whilst giant pangolins were mainly hunted for local consumption and income generation using wire snares. Overall, our study shows the possible value of local knowledge for planning and prioritising conservation actions for pangolins. We highlight the urgent need to monitor pangolin populations, and assess the possible impacts to pangolins from threats such as hunting.
... The pangolin (Manidae), a protected family of scaly anteaters have in the last five years become the most globally trafficked vertebrates [16,17]. Despite being prohibited from trade in most parts and being listed in Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) [18,19], all pangolin species which have been described, four from Sub-Saharan Africa and four from Asia [20][21][22], are all threatened by continuous demand for illegal trade largely from Southeast Asian markets [23,24]. Pangolins are presently experiencing astronomical decline across its ranges and face extinction risk due to habitat degradation, hunting, and sustained trafficking for international trade, which threatens their survival. ...
... Questions on the different uses of these suids, their socio-economic and cultural importance were asked to all socio-professional groups considered without distinction of gender and age [30]. Data collection was carried out using semi-structured interviews based on a questionnaire developed for this purpose. ...
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Background Despite the number of wild animals that are useful to the local population and have served as a basis for ethnozoological research in Benin, others such as Phacochoerus africanus and Potamochoerus porcus remain poorly documented according to their uses. They are already facing threats to their proper conservation in ecosystems where they are found in metapopulations as a result of the fragmentation effects. In this study, the ethnozoological knowledge of these suids in southern Benin was inventoried. Methods To achieve this objective, structured interviews were conducted with 138 people randomly selected in nine districts. The use values of these suids parts and the fidelity level were assessed through global ethnozoological value. Non-parametric tests were carried out to analyse the uses differences according to gender, age and ethnic group. Two Correspondence Analyses (PCA) were carried out to describe use categories within districts and ethnic groups. A Generalized Linear Model (GLMs) of binomial families was used to relate the suids practice hunting to districts and distance from the village. Results More than two use categories (food, medicinal and magical) were identified for both species. With regard to the most commonly used parts, we have the mane, teeth, horns for the common warthog and the skull, teeth, nose and testicles for the red river hog. Ten illnesses were cited to be treated by their parts. Significant differences were observed between ethnic groups regarding the suids medicinal uses but not for gender. The ethnozoological use value of suids was almost similar for adults and the elderly. Conclusion Strong pressure continues to be exerted on these two suids and their habitats despite their high zootechnical potential compared to exotic pigs. After the creation of income-generating activities, future research needs to be carried out to evaluate their budgets activities, genetic study and diet through metabarcoding approaches to promote their domestication. This could help to capitalise data that can be used in breeding programmes in order to limit threats on the few species individuals found in the natural environment, as well as to participate in reintroductions if necessary.
... The pangolin (Manidae), a protected family of scaly anteaters have in the last five years become the most globally trafficked vertebrates [16,17]. Despite being prohibited from trade in most parts and being listed in Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) [18,19], all pangolin species which have been described, four from Sub-Saharan Africa and four from Asia [20][21][22], are all threatened by continuous demand for illegal trade largely from Southeast Asian markets [23,24]. Pangolins are presently experiencing astronomical decline across its ranges and face extinction risk due to habitat degradation, hunting, and sustained trafficking for international trade, which threatens their survival. ...
Article
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We summarize and characterize the emerging role of Nigeria in transcontinental illegal pangolin shipments from Sub-Saharan Africa to Asia from 2012 to 2019 using public online news reports data. The findings indicate that 57 seizure incidents were predominantly pangolin scales as reported from the findings and the equivalent of 462,092 individual pangolins trafficked from Nigeria. Also, findings reveal that China and Vietnam constitute 65 % of pangolin incidents reported. A peak was reached in 2018 for pangolin incidents, and Nigeria also serves as a transit route to Cameroon. We concluded that Lagos plays a crucial role in pangolin shipments to Asia.
Article
Background and research aims Cameroon hosts three species of pangolins and has recently been identified as a hub of pangolin trafficking. However, information on threats to pangolins needed to guide conservation efforts remains scarce, notably on consumption and trade patterns, and the prevalence of individuals trading pangolins. Local communities sharing the same habitats with pangolins can provide such information, which is useful to better target interventions. Methods Based on a snowball sampling approach and using interview surveys in 20 villages surrounding Mbam et Djerem National Park, we investigated the consumption of pangolins and its drivers, parts sold, selling prices, places sold and buyers, and employed the nominative technique to estimate the percentage of people within the surveyed population engaged in selling pangolins. Results Our results showed that both giant and white-bellied pangolins are locally consumed, mainly for their taste, and traded for meat and scales. Distance to the capital city Yaoundé, ethnolinguistic group, and education significantly affected the likelihood of consuming or trading white-bellied and giant pangolins. Selling and prices of giant pangolin meat and scales were significantly affected by distance to Yaoundé. The prevalence of people selling white-bellied and giant pangolins were higher in Tibati and Yoko compared to other municipalities. Implications for conservation Overall, our study provided information necessary for effective law enforcement and research-oriented decision-making for pangolin conservation. We recommend the establishment of consumption reduction campaigns focusing on taste preference, investigation of the impacts of the newly constructed national road on pangolin supply chains and trafficking, and increasing the involvement of local communities in the management process of Mbam et Djerem National Park.
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Here we present the first field report on the ecology of a single female Sunda pangolin and her young in their natural habitat on an offshore island of Singapore. Observations of this event were made possible by the use of radio-telemetry and infrared-triggered camera traps. Only one offspring was recorded from this birth event and the period of maternal care was approximately 3 to 4 mo. A total of 3 natal dens were used throughout the monitoring period, and hollows of large trees (>50 cm diameter at breast height, DBH) were associated with all dens. The 100 and 95% home range estimated by minimum convex polygons is 6.97 and 5.63 ha, respectively. The daily active period of the pangolin was 127 ± 13.1 min, and peak activity levels were between 03:00 and 06:00 h. The findings of this report are discussed in relation to the conservation challenges confronting this species.
Article
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Article
This field guide begins with a checklist. The main part of the volume consists of entries for each species. Each entry provides information on common names, measurements, recognition, geographical distribution (plus map), habitat, diet, behaviour, adaptations and conservation status. Illustrations are also included. Brief notes are also provided on the African environment (physical, climate and vegetation) and palaeoecology (habitats and species). Finally a short section examines African wildlife conservation.