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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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