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R E S E A R C H Open Access
Ethnozoological and commercial drivers of
the pangolin trade in Benin
Stanislas Zanvo
1,2*
, Sylvestre C. A. M. Djagoun
1
, Fortuné A. Azihou
1
, Bruno Djossa
1,3
, Brice Sinsin
1
and
Philippe Gaubert
2,4
Abstract
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.
Keywords: Ethnozoological knowledge, Spiritual use, Traditional medicine market, Wildlife trade, Pangolins, Benin,
West Africa
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* Correspondence: zanvostanislas@yahoo.fr
1
Laboratory of Applied Ecology, Faculty of Agronomic Sciences, University of
Abomey-Calavi, 01 BP 526, Cotonou, Benin
2
Laboratoire Evolution et Diversité Biologique (EDB), CNRS/UPS/IRD,
Université Toulouse III Paul Sabatier, Bâtiment 4R1, 118 route de Narbonne,
31062 Toulouse cedex 9, France
Full list of author information is available at the end of the article
Zanvo et al. Journal of Ethnobiology and Ethnomedicine (2021) 17:18
https://doi.org/10.1186/s13002-021-00446-z
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
Introduction
Bushmeat—i.e. the wild game from the tropics—consti-
tutes the main animal protein and income sources for
rural people in sub-Saharan Africa [1]. In the Congo
Basin, individual consumption amounts several dozens
of kg per year (e.g. [2,3]). Bushmeat often represents the
cheapest animal protein alternative for poor rural house-
holds [4]. Bushmeat hunting also stands among the
prime income-generating activities in rural areas of trop-
ical Africa, where the bushmeat trade can generate more
than 500 USD per year for a single household hunter
(e.g. [5]).
Bushmeat species also play a vital role in traditional
African medicine where animal-derived body parts
(items) are used for the treatment of diseases, ailments
and spiritual purposes (e.g. [6,7]). The specific markets,
mostly urban, where such items are sold add to the
bushmeat selling network already connecting rural to
main urban centres [8]. As a consequence, bushmeat
consumption and use, which occur at unsustainable
rates in Africa [9], have so far remained an intractable
issue, contributing at the same time to household
wealthiness and biodiversity extinction [10,11].
Pangolins (Pholidota, Mammalia), or scaly anteaters,
have recently emerged as the flagship taxon of the bush-
meat crisis. They are trafficked in unsustainable volumes
to feed the local and international—mostly driven by the
traditional Chinese medicine (TCM)—demands for both
their meat and the medicinal properties of their scales
and other items [12–14]. Pangolins have also been sug-
gested as the intermediate host responsible for the
COVID-19 pandemic [15], despite the lack of concrete
evidence for this claim. In Africa, pangolins have re-
cently seen their trafficking volumes and market prices
increase, in line with the trends observed for the Asian
species [16]. Effective trading networks are now connect-
ing Africa and Asia to feed the TCM demand for pango-
lin scales [14].
Among the four species of African pangolins, the
white-bellied pangolin (Phataginus tricuspis) is the most
frequently found on the bushmeat and traditional medi-
cine markets (e.g. [17,18]), notably in West Africa [6,7,
16,19]. Ethnozoological knowledge on the species shows
a diversity of uses by local communities involved in me-
dicinal and spiritual practices, to treat convulsion,
rheumatism, hiccups, healing wounds, woman unfaith-
fulness and impotence [18,20]. Scales are the most com-
monly used, although various items such as tongue,
bones and head are also regularly found on the trad-
itional medicine market (TMM [6,7];).
Benin, situated in the Dahomey Gap (West Africa) and
often designated as the cradle of the ‘Vodoun culture’,
harbours a vibrant market network for animal-based me-
dicinal and spiritual practices, likely to have deleterious
impacts on biodiversity conservation in the whole sub-
region [6,7]. Although the ethnozoology of pangolins
has received much attention in neighbouring countries
[6,17,20–22], the situation in Benin, where the white-
bellied 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. How-
ever, this study was restricted to southern Benin and in-
cluded only two ethnic groups (Hôli and Fon), despite
the larger extent of pangolin’s distribution in Benin [23].
Investigating on the ethnozoology of pangolins should
help understand the causes and extent of the species de-
cline as related to the medicinal and spiritual practices
that prevail in Benin. In this study, we propose a
country-scale survey of the main ethnozoological drivers
of the pangolin trade encompassing four major Beninese
ethnic groups and incorporating the recently raised issue
of the international demand from local diasporas [14].
Our main objectives are to (i) assess the different uses of
pangolin items among ethnic groups in Benin and (ii)
identify the cohort of buyers involved in the pangolin
trade and related economic values along the chain, in-
cluding the local demand from the Chinese community.
Methods
Study area
The study took place in Benin from April 2018 to April
2019. Benin is a West African country located between
latitudes 6° 10′–12° 25′N and longitudes 0° 45′– 3° 55′
E (Fig. 1). Its vegetation is marked by a severely frag-
mented forest cover due to both drier climatic condi-
tions during the Holocene [25] and ever-growing human
(mostly agricultural) activities [26,27]. Benin is widely
known for its Vodoun culture, which is strongly linked
to a diversity of traditional practices (medicinal and spir-
itual) using various animal derivatives [6,7]. Vodoun
practices have driven the development of a dense TMM
(also called ‘fetish markets’) network in the country, with
at least 42 markets identified in southern and central
Benin (SZ, unpubl. data). Benin counts around 60 ethnic
groups for an estimated population of c. 10 M inhabi-
tants [28]. The ethnic group ‘Fon’is dominant in the
South whereas ‘Nagot’and ‘Bariba’are the most repre-
sented ethnic groups in central and northern Benin, re-
spectively [29].
Data collection
We targeted (i) 54 villages neighbouring all the potential
areas of occurrence of pangolins in Benin and (ii) the
five major TMMs in southern and central Benin (Avog-
bannan,Calavi,Dantokpa,Gbèdagba and Zobè) (Fig. 1).
The target local communities were identified after Zanvo
Zanvo et al. Journal of Ethnobiology and Ethnomedicine (2021) 17:18 Page 2 of 11
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et al. [23] from occurrence areas providing pangolin
scales. Focus group participants and vendors were adults
at least 18 years old. Focus group participants were vol-
unteers, identified through informal interviews with vil-
lagers whereas in the TMMs, availability and trust
towards the interviewer were the criteria of participation.
In each village, we organized a single focus group with
7–11 local people using a pre-established questionnaire
and a poster figuring pangolins. The questionnaire was
conducted as a semi-structured interview addressing the
names used for pangolins in local languages, the differ-
ent uses of pangolins and which items were involved,
the selling prices of pangolins, the categories of clients
and the time period at which the trade with each cat-
egory of clients started (Additional file 1). For each ques-
tion, all the answers were recorded without
discrimination among participants of the focus group. In
the TMMs, we generally had to use a modified approach
Fig. 1 Villages and traditional medicine markets (TMMs) in Benin surveyed as part of this study
Zanvo et al. Journal of Ethnobiology and Ethnomedicine (2021) 17:18 Page 3 of 11
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
focusing on the questions related to the commercial
value of pangolins. Such a strategy was necessary be-
cause we were not able to gather simultaneously several
vendors and the latter were not available for long-time
interviews, due to their activity. However, six inter-
viewees out of 35 were able to fully answer the question-
naire. Their responses on the ethnozoology of pangolins
were treated as a single focus group, as they all belonged
to the same Fon ethnic group.
Data analysis
We used the International Classification of Diseases
(ICD-11; version 09/2020) to group the recorded medi-
cinal uses into ICD categories for each item of pango-
lins. Given the important number of spiritual uses
recorded, we created an additional, dedicated category.
Each focus group was considered as a single observation
for the analysis. Descriptive statistics were used to assess
the frequency of citation of the items used, ICD categor-
ies mentioned by focus group participants and type of
clients buying pangolin items. We used the use value
index (UV; adapted from [30]) to assess the spectrum of
use of each pangolin item for medicinal and spiritual
purposes, as follows:
UV ¼
PUP
n
where Up represents the number of uses mentioned
by focus groups for each pangolin part and nthe total
number of focus groups.
One-way analysis of similarities (ANOSIM test) based
on 9999 permutations was carried out using the vegan
package in Rversion 4.0.0 [31] on a matrix of ICD and
spiritual categories including the ethnic groups parti-
tioned into geographic regions (South, Centre and North
Benin). The Rstatistics was interpreted according to
Clarke [32]: (i) R< 0.25 means no separation between
groups, (ii) 0.25 < R< 0.5 some level of separation be-
tween groups despite a degree of overlap, R> 0.75 well
separated groups, and R= 1 total separation between
groups. The ANOSIM test was used to assess variation
of knowledge among ethnic groups (recorded at least
four times: Bariba, Fon, Mahi and Nagot) and geo-
graphic regions using medicinal and spiritual use pat-
terns. We used non-metric multidimensional scaling
(NMDS) analysis to visualize in a Cartesian space the
dissimilarity between ethnic groups and geographic re-
gions. We removed meat from ANOSIM and NMDS
analyses due to the absence of medicinal and spiritual
uses for this specific pangolin item. We calculated the
average selling prices among local communities and in
the TMMs for the different categories of clients. We
used the ttest to compare average selling prices
according to the categories of clients in rural areas (local
communities) and in TMMs and between rural areas
and TMMs.
Results
Fifty-four focus groups were carried out across the oc-
currence zone of pangolins in Benin with 18, 22 and 16
focus groups in southern, central and northern regions
respectively. Thirty-five individual interviews were per-
formed in the TMMs. Focus group participants were
mostly men (84%), between 22 and 76 years (mean = 36
years), and included farmers (36%), farmer-hunters
(54%) and housewives (8%), whereas all the interviewees
from the TMMs were men, aged 26 to 54 (mean = 37).
Lihoui was the common name almost unanimously used
for pangolins among ethnic groups in southern Benin
whereas in central Benin, Nagot, Ifè and Idatcha named
pangolins either Aïka or Akikan. In northern Benin, the
common name of pangolins changed from an ethnic
group to another (Table 1). Pangolins were unanimously
cited by all the focus group participants for their use as
an animal protein source. Medicinal and spiritual uses
were cited by 65% and 37% of the focus groups, respect-
ively. Scales, tongue, head, bones, xiphisternum (a spe-
cific prolongation of the sternum to attach the tongue
muscles), blood and heart were the eight pangolin items
cited by focus group participants (Fig. 2). Scales (64%),
tongue (18%), bones (15%) and head (13%) were the
items most cited by the focus groups for medicinal and
spiritual uses. A total of 42 medicinal (n= 31) and spirit-
ual (n= 11) uses were recorded. The medicinal uses fell
into 15 out of 26 ICD categories. The scales were the
item of pangolins that had the highest use reported (56),
number of ICD categories (14) and use value (UV =
Table 1 Common names used for the white-bellied pangolin
among ethnic groups and geographic regions in Benin
Region Ethnic groups Local names
South Adja,Agouna,Aïzo,Gun Lihoui
Fon Lihoui,Houékin
Yoruba,Hôli Iwô
Centre Mahi Kosso Lihoui
Nagot,Ifè,Idatcha Aïka,Akikan
North Anii Sassabokourou
Bariba Kokowaka
Nagot loufiloufi
Lokpa Libilibi
Kotokoli Kaminakpara
Ditamaré Tempétakpara
N'tcha Akonkon
Yom Narm
Zanvo et al. Journal of Ethnobiology and Ethnomedicine (2021) 17:18 Page 4 of 11
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
1.56), followed by tongue, bones, head, xiphisternum,
blood and heart (Table 2). The spiritual use category
was the most mentioned by focus group participants
(69%), superior to any ICD categories (Fig. 3). Sexual be-
witchment was the spiritual use most frequently re-
ported by focus groups, followed by the control of
women’s infidelity. Among the 15 recorded ICD categor-
ies, neoplasms (ICD-2; 31%), traditional medicine condi-
tions (ICD-26; 25%), and certain infectious or parasitic
diseases (ICD-1; 22%) were the most cited ICD categor-
ies by focus group participants (Fig. 3). The ANOSIM
multivariate analyses showed slight differences of ethno-
zoological knowledge among the four major ethnic
groups (R= 0.55; p<0.001) and geographic regions (R=
0.48; p<0.001). All the Rstatistic values ranged between
0.25 and 0.5, indicating that the ethnic groups shared
some level of ethnozoological knowledge. The NMDS
plot showed some variation of the distance within and
between Fon,Nagot and Mahi ethnic groups located in
southern and central (Fig. 4). Conversely, the Bariba
were close to each other, and generally remote from and
opposite to all the Fon,Mahi and almost all the Nagot.
We observed some overlapping between focus groups
among Bariba,Mahi and Nagot ethnic groups. More-
over, some focus groups among Mahi and Nagot ethnic
groups overlapped with Fon whereas others were remo-
tetly distributed in space.
Local communities cited five client types involved in the
trade of pangolins in Benin (restorers [i.e. managers of res-
taurants], traditional healers, traders of traditional medi-
cine markets, foreigners from West African countries and
Chinese community), while only two client types (trad-
itional healers and Chinese community) were reported by
TMM stakeholders (Fig. 5). Restorers, traders from TMMs
Fig. 2 Frequency of citations of pangolin items used by local communities
Table 2 Pangolin’s items used in traditional medecine and spiritual practices in Benin. Spiritual purposes are in bold. Use report
(UR), International Classification of Diseases (ICD) and Use Value (UV)
Items Conditions treated UR ICD UV
Scales Physical strength, oedema, cough, hiccups, rheumatism, convulsion of new born, pharyngitis, healing wound, snake
bite, measles, man dominance in household, healing fire burns, defeating the opponent in case of litigation,
facilitate delivery, infertility, scabies, sexual bewitchment, ringworm, joint pain, stomach aches, vigour of new born,
breast cancer, dizziness, heart palpitations, late infantile spinal muscular atrophy, osteoarthritis and asthma
56 14 1.56
Tongue Stomach aches, pneumonia, hip pain, developmental language disorders, protection against sexual bewitchment,
Epilepsy, incantation,defeating the opponent in case of litigation, sterility, thief sickness and stop women’s
infidelity
12 7 0.33
Bones Hiccups, cough, osteoarthritis, healing wound, ataxia, asthma, coxarthrosis, developmental language disorders and
late infantile spinal muscular atrophy.
11 6 0.31
Head Oedema, pharyngitis, epilepsy, defeating the opponent in case of litigation, vigour of new born, unnatural
power and luck
9 5 0.25
Xiphisternum Impotence,sexual bewitchment 4 2 0.11
Blood Asthma and convulsion of new born 3 2 0.08
Heart Man dominance in household 1 1 0.03
Zanvo et al. Journal of Ethnobiology and Ethnomedicine (2021) 17:18 Page 5 of 11
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and foreigners from West Africa (Togo and Ghana, ac-
cording to focus group participants) were cited by 74, 48
and 30% of the focus groups, respectively, as clients buy-
ing pangolins to local communities. Traditional healers
(4%) and Chinese community (9%) were the client cat-
egories less cited by local communities. Within the TMM
stakeholders, traditional healers (96%) and Chinese com-
munity (88%) were the only clients cited as buying pango-
lins. Local communities and the TMM stakeholders
highlighted that the trade of pangolins with local people
exists since the development of wildlife markets whereas
the trade with West African and Chinese communities
Fig. 3 Frequency of citations of the conditions treated with pangolin items according to ICD categories
Fig. 4 Spatial distribution of ethnic groups according to their ICD and spiritual categories-based knowledge. Euclidean distances between ethnic
groups and geographic regions (C—centre; N—north; S—south) reflect the degree of similarity of knowledge. Overlapping implies the sharing of
the same knowledge
Zanvo et al. Journal of Ethnobiology and Ethnomedicine (2021) 17:18 Page 6 of 11
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
started approximately one decade ago. Pangolin pricing
changed according to the category of stakeholders (local
community vs. trader in TMMs) and clients (local and
West African clients vs. Chinese community) (Table 3).
The average selling prices were higher in the TMMs than
in local communities. The Chinese community bought
only pangolins alive, while local and West African cus-
tomers both bought alive and dead pangolins or pangolin
items. Head and scales were only sold in the TMMs to
local and West African clients. The highest selling price
(73.38 USD) for a pangolin (alive) was recorded in a
TMM for a client from China. The average selling price of
pangolins sold on the TMMs to the Chinese community
was significantly higher (t=-34.089;p< 0.001) than the
selling prices to local and West African clients. The same
trend (t=−16.238; p< 0.001) was recorded at the local
community level with the Chinese community buying
pangolins alive at higher prices than local and West Afri-
can clients. There was no significant difference between
the average selling prices to Chinese community in local
communities and TMMs.
Discussion
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].
Lihoui was the common name for pangolins among
most of the ethnic groups in southern Benin, whereas
Aîka/Akika was used in central Benin. This uniformity
of common names among ethnic groups located within
Fig. 5 Citation frequencies of client categories within traditional medicine markets (black bars) and local communities (grey bars) in Benin. West
African foreigners originate from Ghana and Togo
Table 3 Variation in pangolin pricing between local communities and traditional medicine markets in Benin, after client categories.
African clients other than local originate from Togo and Ghana. Prices are given in USD (conversion: 30 July 2020). TMMs refers to
traditional medicine markets
Pangolin pricing (USD)
Local communities TMMs
Type of item Average Local and West African clients Chinese clients Local and West African clients Chinese clients
Alive Min 5.01 42.9 18.96 55.35
Max 7.84 60.05 24.83 73.38
Dead carcass Min 4.52 –18.96
Max 6.68 –24.83
Head Min ––3.88
Max ––4.73
Scales (whole body) Min ––8.62
Max ––12.06
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the same geographic region (Fon,Aizo,Adja and Gun in
South, and Nago,Ifè and Idatcha in Centre) could be
due to the genealogical relationships between the sam-
pled ethnic groups, which originate from the same an-
cestors and share similar languages [29]. The Fon ethnic
group is believed to be the initiator of the trade in ani-
mal derivatives since the Dahomey Kingdom (from c.
1600 to 1904 AD) and represents the vast majority of the
actors currently operating in this sector in Benin (SZ,
unpubl. data). The common name Lihoui could have
originated from this dominant ethnic group and through
the colonization of other ethnic groups have diffused
since centuries in all the TMMs of southern Benin (see
[33]). Conversely, northern Benin showed eight different
names used for pangolins. This is likely because (i) this
region has been colonized by ethnic groups from differ-
ent origins with socio-linguistic divergences [34,35] and
(ii) the TMM network is almost inexistent, thus prevent-
ing from any diffusion of a dominant pangolin name.
Pangolins are used for food, medicinal and spiritual
purposes in Benin, in line with the literature record for
tropical Africa [21,22,36–41]. More specifically, pango-
lins in Benin constitute at the same time a bushmeat re-
source and an important input to traditional medicine
and cultural practices [24]. Pangolin meat is unani-
mously consumed as food in Benin, whereas no medi-
cinal 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.
Among the eight recorded pangolin items, scales were
by far the most commonly used by local communities
for medicinal and spiritual purposes. Scales had the
highest use value (UV = 1.56) and were mentioned in 14
out of the 15 ICD categories recorded for 56 different
use reports. These results show that the scales possess a
great traditional value for local people in Benin, probably
justifying the storage of old scale samples in rural house-
holds [23] and the great number of scales present in the
TMM stalls (SZ, CD, PG; unpubl. data). Our results cor-
roborate those of previous studies in tropical Africa
pointing out the high use value and versatility of use of
pangolin scales in comparison with other items [21,37,
42]. Although the diversity of items for ethnozoological
use was generally lower in Benin than in other African
countries ([6,21,37]; but higher than in [24,43]), we
observed the use of a so far unreported pangolin item,
the xiphisternum. The latter is involved in impotence
and sexual bewitching treatments and could be a par-
ticular knowledge of Fon and Mahi ethnic groups.
A large proportion of the disease and ailment treat-
ments where pangolin items are involved had never been
described for Benin [24], although almost all of them
had already been recorded in West Africa [21,37]. The
ratio between the diversity of medicinal/spiritual uses
and items was higher in Benin (42 medicinal and spirit-
ual purposes for eight items) than what was recorded
from traditional healers and fetish markets in other
West African countries [6,21,37]. In Benin, spiritual
uses (69%) and the ICD categories neoplasms (31%) and
traditional medicine conditions (25%) were the most
cited by local communities. Overall, this indicates that
pangolin items are used against many diseases and ail-
ments generally uncovered by conventional medicine, re-
vealing the high endogenous value of pangolins in a
country where Vodoun practices are thriving [44].
Our results suggested slight differences of ethnozoolo-
gical knowledge within and between ethnic groups and
geographic regions, and at the same time indicated par-
tial overlapping of ethnozoological knowledge among
the major ethnic groups (Fon,Nagots,Bariba and Mahi)
in Benin. Socio-cultural distances between ethnic groups
in Benin (see [34,35]) could be a factor explaining the
divergences of knowledge on medicinal and spiritual
uses of pangolins. For instance, the use of scales to treat
breast cancer was only cited by the Bariba ethnic group
whereas pneumonia, epilepsy, rheumatism and measles
were only recorded among Fon. The Nagot were the only
ethnic group that mentioned the use of pangolin items
for the treatment of ringworm, scabies and heart palpita-
tions. Thus, our results show a partial influence of ethni-
city on ethnozoological knowledge of pangolins, in line
with what was observed in Ghana [20]. However, Fon,
Mahi and Nagot ethnic groups also share ethnozoologi-
cal knowledge on pangolins such as healing wound,
treatment of sterility, hiccups, easy delivery, snakebite,
pharyngitis, sexual bewitchment, defeating the opponent
in case of litigation, and sterility. South and central
Benin count 42 TMMs (SZ, unpubl. data) mainly domi-
nated by the Fon ethnic group, whose beliefs are strongly
linked to Vodoun practices. Through this market net-
work, the Fon ethnic group has probably diffused or
gained medicinal and spiritual knowledge of pangolins
in the central region of Benin.
As part of the bushmeat species spectrum sold in Cen-
tral and West Africa, pangolins constitute a traditional
source of income for rural communities, restorers and
TMM vendors [6,16,19]. In Benin, we showed that 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 (from
alive to scales). Average selling prices for live and dead
pangolins were higher in TMMs (18.96–73.38 USD),
probably to write off the costs of being at the tip of the
supply chain and to benefit from the higher wealthiness
of urban households [4]. However, excluding buyers
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from the Chinese community, average selling prices of
pangolins in rural areas (4.52–7.84 USD) and in TMMs
(18.96–24.83 USD) were lower than those recorded in
the bushmeat markets from central Africa one decade
ago (mean = 34.27–35.66 USD [14]). The relatively
cheap cost of pangolins on the Beninese markets means
that the species is economically reachable by West Afri-
can consumers, which may not facilitate the mitigation
of the volumes of pangolins extracted each year in Benin
as part of the wildlife trade.
Our study highlighted the international component of
the pangolin trade in Benin through the diversity of for-
eign clients, involving West African and Chinese na-
tionals since c. one decade (probably also involving
Vietnamese nationals, after a case of recent seizure at
Cotonou airport [14]). Whether pangolins are consumed
locally or brought back to the clients’home countries is
unknown and would deserve further investigations. We
revealed different practices according to the type of cli-
ents, the Chinese community only buying pangolins alive
whereas local and West African clients would also buy
dead pangolins and various items (head, scales). The
average selling prices of pangolins to the Chinese com-
munity was 3–8 times higher than the selling prices to
local and West African clients, confirming that Chinese
diasporas in Africa has brought further economic incen-
tives to the endemic pangolin trade [16,19]. Further in-
vestigations are required to assess whether the Chinese
diaspora demand has modified the practices of wildlife
traders and is constitutive of an ‘overexploitation vortex’
of pangolins (see [45]) in Benin.
Assuming a mean body weight of 2.48 kg for the
white-bellied pangolin (2.36 ♂–2.6 ♀kg; in [46]) and a
1/4–1/3 contribution of the scales to the total weight of
the species [47], scales bought by local and West African
clients would cost 10.39–19.45 USD per kilogramme.
Because there is a possibility that the scales of pangolins
bought by the Chinese community in Benin end up feed-
ing the global pangolin trade (e.g. 513 kg of scales were
seized in Cotonou in 2018 [14]), we also estimated the
average price of pangolin scales per kilogramme if scales
were extracted from live pangolins (17.30–29.59 USD
per kg). Such prices confirm that pangolin scales consti-
tute a valuable source of income on West African mar-
kets [21]. Compared to the prices of pangolin scales sold
in China and Vietnam (485–759 USD per kg) [12], both
paths of acquiring scales in Benin (either directly or
from a live animal) would remain highly profitable to
traffickers that feed the illegal pangolin trade to—
mostly—China [48].
Studying client practices allowed us to delineate a
complex network for the pangolin trade in Benin, in-
cluding (i) the TMM that mostly supplies traditional
healers and the Chinese community and (ii) a less
urbanized and more diffused market network where
local and West African clients, together with restorers
and TMM stakeholders, would be the main buyers. The
low contribution of the Chinese community as men-
tioned by local communities (9%; only from the Lama
forest reserve, the main forest island from southern
Benin) may imply that intermediates collect pangolins
from more proximal sources than the TMMs, but this
will require further investigations.
Conclusions
Our study addressed the ethnozoology of pangolins
across Benin and its major ethnic groups and revealed
the importance of the traditional (medicinal and spirit-
ual) and economic values of pangolins for local commu-
nities and TMM vendors. Our results suggest that the
pangolin trade in Benin is based on endogenous prac-
tices now influenced by economic drivers (higher prices
and change of selling practices) from the local Chinese
demand and that a number of actors are involved in an
intricate, multi-scale network. Conserving pangolins in
Benin will require considering the multiple, cultural and
economic drivers of the market and engaging synergic
efforts against both endemic and international traffick-
ing. Long-term monitoring of offtake and trafficking net-
work in Beninese markets and targeted habitats,
together with higher law enforcement, will be essential
to reverse the decline of the white-bellied pangolin as
witnessed by Beninese rural communities [23].
Abbreviations
ANOSIM: Analysis of similarity; ICD: International Classification of Diseases;
NMDS: Non-metric multidimensional scaling; TMM: Traditional medicine
market; TCM: Traditional Chinese medicine; UR: Use report; USD: United State
dollar; UV: Use value
Supplementary Information
The online version contains supplementary material available at https://doi.
org/10.1186/s13002-021-00446-z.
Additional file 1. Questionnaire sur l’ethnozoologie et le commerce des
pangolins.
Acknowledgements
This study received the approbation of Public Forest Service and the prior
consent from local authorities in each district. We are grateful to these
different authorities and all the participants for generously giving their time
to complete this survey. Aubin Anago and Alfred Kakè played an important
role for data collection in the TMMs. We thank two reviewers for their useful
comments on the early draft of the manuscript.
Authors’contributions
ZS, GP, DS and SB designed the study. ZS conducted the data collection,
statistical analyses and wrote the first version of the manuscript, which was
read and improved by all the authors. All authors approved the final version
of the manuscript.
Funding
This research received financial support from the program Jeune Equipe
Associée à l’IRD (RADAR-BE). ZS is supported by a PhD grant ‘ARTS-IRD’.
Zanvo et al. Journal of Ethnobiology and Ethnomedicine (2021) 17:18 Page 9 of 11
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
Availability of data and materials
All the data generated and analysed during the current study are included in
this manuscript.
Declarations
Ethics approval and consent to participate
All the participants in the focus groups and interviews were volunteers who
gave their informed consent before interviews.
Consent for publication
The authors give their agreement for the publication of this manuscript.
Competing interests
The authors declare no competing interests.
Author details
1
Laboratory of Applied Ecology, Faculty of Agronomic Sciences, University of
Abomey-Calavi, 01 BP 526, Cotonou, Benin.
2
Laboratoire Evolution et
Diversité Biologique (EDB), CNRS/UPS/IRD, Université Toulouse III Paul
Sabatier, Bâtiment 4R1, 118 route de Narbonne, 31062 Toulouse cedex 9,
France.
3
Laboratoire de Foresterie et de Conservation des Bioressources
(LaFCBio), Ecole de Foresterie Tropicale, Université Nationale d’Agriculture,
Kétou, Benin.
4
Centro Interdisciplinar de Investigação Marinha e Ambiental
(CIIMAR), Universidade do Porto, Terminal de Cruzeiros do Porto de Leixões,
Av. General Norton de Matos, s/n, 4450-208 Porto, Portugal.
Received: 8 January 2021 Accepted: 9 March 2021
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