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Ecological diversity and conservation
of wild edible fruit trees species
in the Lama Forest Reserve in Benin
Photo 1.
Aerial view of the Lama Forest Reserve in Benin.
Photo S. Agbahoungba.
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3) 53
CONSERVATION D’ARBRES SAUVAGES À FRUITS COMESTIBLES / LE POINT SUR…
Symphorien Agbahoungba1, 3
Achille Ephrem Assogbadjo1
Flora Josiane Chadare2
Rodrigue Idohou4
Valère Kolawolé Salako4
Eric Etchikinto Agoyi1, 3
Romain Lucas Glèlè Kakaï4
1 University of Abomey-Calavi
Faculty of Agronomic Sciences
Laboratory of Applied Ecology
01 PO Box 526, Cotonou
Benin
2 University of Agriculture of Kétou
School of Sciences and Techniques
for Preservation and Processing
of Agricultural products
BP 114, Sakété
Republic of Benin
3 Makerere University
PO Box 7062, Kampala
Uganda
4 University of Abomey-Calavi
Faculty of Agronomic Sciences
Laboratory of Biomathematics
and Forest Estimations
04 PO Box 1525, Cotonou
Benin
Correspondance électronique avec les auteurs / Electronic correspondence with authors
rodrigidohou@gmail.com
RÉSUMÉ
DIVERSITÉ ÉCOLOGIQUE ET
CONSERVATION D’ARBRES SAUVAGES À
FRUITS COMESTIBLES DANS LA RÉSERVE
FORESTIÈRE DE LAMA AU BÉNIN
Une bonne connaissance des interac-
tions entre les forêts et les communautés
humaines voisines est indispensable à
la gestion participative des ressources
forestières. La présente étude s’est atta-
ché à déterminer la diversité écologique
des arbres sauvages à fruits comes-
tibles (ASFC) dans la Réserve forestière
de Lama, zone protégée dans le sud
du Bénin, et à définir des stratégies de
conservation adaptées. Un inventaire
des ASFC a été réalisé sur 53 placettes
systématiquement délimitées dans des
forêts denses typiques, de forêt dense
dégradée et de jachère dans la Réserve
forestière de Lama, ainsi qu’une enquête
ethnobotanique auprès de 136 partici-
pants. Les habitats des ASFC ont été iden-
tifiés à partir d’une Analyse des Corres-
pondances Simples sur les données de
densité. Une liste d’especes prioritaires
a été générée par la méthode des rangs
composés : Dialium guineense, Diospy-
ros mespiliformis, Drypetes floribunda,
Mimusops andogensis et Pterocarpus
santalinoides en forêt dense typique ;
Pancovia bijuga, Psidium guajava et
Lecaniodiscus cupanioides en forêt
dense dégradée ; Ficus capensis et Spon-
dias mombin dans les jachères. Contrai-
rement aux autres essences, la densité
moyenne de D. guineense, D. mespili-
formis, D. floribunda, L. cupanioides et
M. andongensis varie de manière signifi-
cative (P < 0,01) selon les types de végé-
tation. Les ASFC recensés sont principa-
lement utilisés dans l’alimentation et la
pharmacopée. Les essences à conserver
en priorité sont P. guajava, S. mombin,
F. capensis, P. santalinoides et P. bijuga.
Des efforts de protection renforcés sont
nécessaires en forêt dense dégradée et
dans les zones de jachère pour assurer
la conservation de ces essences. D’autre
part, il convient d’intégrer les ASFC prio-
ritaires dans des programmes de planta-
tion afin de réduire la pression des com-
munautés humaines voisines.
Mots-clés : conservation, réserve
forestière de Lama, diversité, arbres
sauvages à fruits comestibles.
ABSTRACT
ECOLOGICAL DIVERSITY AND
CONSERVATION OF WILD EDIBLE FRUIT
TREE SPECIES IN THE LAMA FOREST
RESERVE IN BENIN
Understanding interactions between
forests and neighbouring human com-
munities is crucial to participative mana-
gement of forest resources. This study
aimed to determine the ecological diver-
sity of wild edible fruit tree (WEFT) species
in the Lama Forest Reserve, a protected
area in southern Benin, and strategies
for their conservation. An inventory of
WEFT species was carried out in 53 sys-
tematically delimited plots in typical
dense forest, degraded dense forest and
young and old fallows in the Lama Forest
Reserve. An ethnobotanical survey was
conducted with 136 respondents and
the WEFT species habitats were iden-
tified by applying a Simple Correspon-
dence Analysis to the density data. A list
of priority species was generated using
the Compound Ranking System method.
The results revealed ten WEFT species,
including Dialium guineense, Diospy-
ros mespiliformis, Drypetes floribunda,
Mimusops andogensis and Pterocarpus
santalinoides in typical dense forest;
Pancovia bijuga, Psidium guajava and
Lecaniodiscus cupanioides in degraded
dense forest; and Ficus capensis and
Spondias mombin in fallows. Unlike the
other species, the average density of
D. guineense, D. mespiliformis, D. flori-
bunda, L. cupanioides and M. andongen-
sis varied significantly (P < 0.01) from one
vegetation type to another. The WEFT spe-
cies recorded were mainly used for food
and medicinal purposes. Top priority
species for conservation were P. guajava,
S. mombin, F. capensis, P. santalinoides
and P. bijuga. More protection efforts
should target the degraded dense forest
and fallow areas to ensure the conser-
vation of these species. Further, priority
WEFT species need to be included in
plantation programmes to reduce pres-
sure from riparian populations.
Keywords: conservation, Lama forest
reserve, diversity, wild edible fruit tree.
RESUMEN
DIVERSIDAD ECOLÓGICA
Y CONSERVACIÓN DE FRUTALES
SILVESTRES COMESTIBLES EN LA
RESERVA FORESTAL DE LAMA EN BENÍN
El manejo participativo de los recursos
forestales requiere un buen conoci-
miento de las interacciones entre bos-
ques y poblaciones aledañas. El objetivo
de este estudio es determinar la diver-
sidad ecológica de los árboles frutales
silvestres comestibles (AFSC) y definir las
estrategias de conservación adaptadas
en la Reserva Forestal de Lama, un área
protegida del sur de Benín. Se llevó a
cabo un inventario de especies de AFSC
en 53 parcelas sistemáticamente delimi-
tadas en zonas típicas de bosque denso,
bosque denso degradado y barbecho de
la reserva Forestal de Lama. Se realizó
una encuesta etnobotánica con 136 par-
ticipantes y se identificaron los hábitats
de AFSC mediante análisis de corres-
pondencias simples de los datos de
densidades. El método de rango de los
componentes permitió generar una lista
de diez especies prioritarias: Dialium
guineense, Diospyros mespiliformis, Dry-
petes floribunda, Mimusops andogensis
y Pterocarpus santalinoides en bosque
denso típico; Pancovia bijuga, Psidium
guajava y Lecaniodiscus cupanioides en
bosque denso degradado; y Ficus capen-
sis y Spondias mombin en barbechos.
A diferencia de las otras especies, la den-
sidad promedio de D. guineense, D. mes-
piliformis, D. floribunda, L. cupanioides y
M. andongensis varía de modo significa-
tivo (P < 0.01) según el tipo de vegetación.
La mayoría de AFSC inventariados tiene
un uso alimentario y medicinal. Las espe-
cies prioritarias para su conservación
son: P. guajava, S. mombin, F. capensis,
P. santalinoides y P. bijuga. Es necesario
reforzar los esfuerzos de protección en el
bosque denso degradado y las zonas de
barbecho para garantizar la conservación
de estas especies. Por otra parte, deben
incluirse los AFSC prioritarios en los pro-
gramas de siembra para reducir la pre-
sión de las poblaciones aledañas.
Palabras clave: conservación, reserva
forestal de Lama, diversidad, árboles
frutales silvestres comestibles.
S. Agbahoungba, A. E. Assogbadjo,
F. J. Chadare, R. Idohou, V. K. Salako,
E. E. Agoyi, R. L. Gièlè Kakaï
54
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3)
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Introduction
Non timber forest products (NTFPs) are the products
extracted from forest ecosystems and which are used for
food, animal feed, medicine, and income generation or
endowed of a social, cultural or religious signification.
These resources constitute an important income source
for local populations. In rural areas in Sub-Saharan Africa,
NTFPs contribute up to 46% of population’s income (Maha-
patra et al., 2005; Assogbadjo et al., 2009; Fandohan et
al., 2010).They include wild edible fruit tree (WEFT) species
which are treasured for their fruits, seeds, leaves, barks,
and roots (Belinda et al., 2013). These non-cultivated foods
are used as diet supplement, as coping mechanism in food
shortage times, and provide an important safety net for the
rural households (Takasaki et al., 2004) while contributing
to their food and nutritional security. Indeed, WEFT species
are one of the best sources of vitamin A, iron, calcium, cop-
per, magnesium, phosphorus, zinc, antioxidants and other
nutritive element useful for a balanced nutrition (Mahapatra
et al., 2012). As such they could help to substantially reduce
food insecurity in rural communities especially during food
shortage periods.
The high nutritional value, the easy access, the lack
of cares and the good organoleptic quality of WEFT species
have led to their overexploitation even in natural forests
which constitute the reservoir of WEFT species (Rigg et al.,
2009). Within the same context, FAO (2010) reported that
African forests’ coverage decreases at an alarming rate
due to human pressure. This forest degradation has been
pointed out as one of the major reason of the progressive
disappearance of WEFT species in developing countries (Kru-
sters et al., 2006). For instance, Benin has lost nearly one
third of its forest coverage since 1990, with the highest lost
rate (2.4% per year) between 2000 and 2005 (FAO, 2012).
Its protected areas represent one of the last refuges of WEFT
species, while some could be found on farms (Hermans et
al., 2004). Although, protected areas are subject to limited
human activities, species within these areas are sometimes
illegally exploited (Racquel, 2014).
Lama Forest Reserve located in the southern part of
Benin, is one of the last vestiges of natural dense forest
within the country. It is a mosaic of forests and fallows (young
fallow, old fallow, typical dense forest and degraded dense
forest) with different ecological patterns (types of soil and
diversity of species) (Bonou et al., 2009). The young fallow is
characterized by the predominance of Chromolena odorata
L. (75% of cover of the vegetation), whereas the old fallow
is characterized by the predominance of Ceiba pentandra
L., Ficus capensis Thunb. and Anogeissus leiocarpa (DC.)
Guill. The typical dense forest is dominated by Afzelia afri-
cana Sm., Ceiba pentandra L., Dialium guineense Willd.,
Diospyros mespiliformis Hochst. and Mimusops andon-
gensis Bruce. The degraded dense forest is dominated by
Cynometra megalophylla Harms (Bonou et al., 2009). These
vegetation types are protected since they are included in the
reserve, but more effort are concentrated on the degraded
dense forest and the fallows to recover the equilibrium with
the typical dense forest. Due to this diverse patterns, it is
expected that distribution and density of plant species differ
from one vegetation type to another. The management prac-
tice within this forest consists on a yearly planting of tree
species around the central core of the reserve. These spe-
cies often included are: Tectona grandis L., Gmelina arborea
Roxb., and Khaya senegalensis (Desr.) A. Juss. (Djego et al.,
2007). However, no edible fruit trees plantations have been
envisaged which may contribute to the reduction of human
pressure on WEFT species in the reserve. Because all species
cannot be conserved at once and do not receive the same
pressure and threats, it is necessary to inventory the diver-
sity of WEFT species in the Lama Forest Reserve and estab-
lish priorities for conservation. This could help to have a list
of species deserving conservation actions within the natural
forest and bring out those to target in management action.
This is crucial as limited funds are often available for species
conservation (Brehm et al., 2010; IUCN, 2011).
Ethnobotanical, food, cultural, economic and pharma-
cological aspects of WEFT species have been widely docu-
mented in Africa (Kebu and Fassil, 2006; Racquel, 2014)
and more specifically in Benin (Assogbadjo, 2000; Codjia et
al., 2003; Chadare et al., 2008; Fandohan et al., 2010). Most
of these studies focused on species in agroforestry systems.
Few studies had targeted natural populations of WEFT spe-
cies within protected areas. Such information is however
critical as it can help to design a sustainable management
strategies for these species even in protected areas. An
active conservation of WEFT species diversity, requires
the establishment of priority within species (Maxted et al.,
2006). But there is no single method to develop effective
strategies for biodiversity conservation. In the approach
described by Brehm et al. (2010), eight criteria are com-
monly used and four methods are generally applied to these
criteria to select the priorities species. Taking into account
that Lama Forest Reserve is a protected area, four criteria
were able to discriminate the WEFT species identified. These
criteria included: distribution (within the forest reserve),
ethnobotanical value, native status and threat assessment.
Compound Ranking System method was used in this study
to identify the priority species as it was the most suitable
among all four methods developed by Brehm et al. (2010).
Compound Ranking System method is easy to verify, flexible
and easy to apply for a large data set (Idohou et al., 2013).
This study aimed at assessing the ecological diversity
of WEFT species and generate a list of priority species for
further reforestation purpose in the Lama Forest Reserve
in Benin.
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3) 55
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56
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3)
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Material and methods
Study area
The Lama Forest Reserve is located between 6°55’ and
7°00’ latitude North and 2°4’ and 2°12’ longitude East in Sou-
thern Benin (White, 1983; figure 1). The forest was gazetted
by law in 1946 as forest reserve (Bonou et al., 2009). Lama
Forest Reserve covers 4,777 ha of which 1,900 ha of dense
forest, and the remaining being fallows (young and old fal-
lows) (Bonou et al., 2009, photo 1). The surrounding area of
the reserve is occupied by the plantations of T. grandis and
G. arborea (292 ha). The annual average rainfall fluctuates
between 1,000 mm and 1,200 mm and the relative humidity
averages 74.5%. Lama Forest Reserve is in a depression of
an average altitude of 60 m. The soil is clayey, more than
2 meters in depth (Akpona et al., 2008).
The local populations around the Lama Forest Reserve
belong mainly to Fon, Holli and at a lesser extent to Adja
sociolinguistic groups. Primarly, they settled inside the
forest, but later in 1984, they have been settled outside the
forest for the purpose of forest management plan. The main
activities around the forest are agriculture, animal rearing,
hunting and small-scale trading.
Sampling and data collection
A systematic sampling method was used following
the seven transect lines of the Lama Forest Reserve. This
method enabled to achieve a good precision by distribu-
ting the plots in all the vegetation communities (Rondeux,
1999). Square plots of 45m × 45m (Salako et al., 2013) were
systematically demarcated at each kilometer with first plots
installed at 15-20 meters away from each path. In total,
53plots (figure 1) were installed in the four vegetation types
of the forest reserve: 13 plots in the typical dense forest;
14 in the degraded dense forest; 14 in the young fallow; 8 in
old fallow and 4 in the surrounding plantations. The 4 plots
demarcated in the surrounding plantations were not consi-
dered in the analysis because of the absence of WEFT spe-
cies in these plots. Within each plot, the diameter at breast
height (dbh) of individual with dbh ≥ 5 cm was measured for
all WEFT species and non-WEFT species.
Figure 1.
Map showing the location of the Lama Forest Reserve in Benin and the plots.
Plots are square plots of 45m × 45m systematically demarcated at each kilometer following
the seven transect lines of the Lama Forest Reserve.
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3) 57
CONSERVATION D’ARBRES SAUVAGES À FRUITS COMESTIBLES / LE POINT SUR…
Data analysis
Assessment of the ecological diversity and structure
of wild edible fruit tree species in the Lama Forest Reserve
The exhaustive list of WEFT species within each inven-
tory plot was established in each plot by recording the name
of each species and the total number of its individuals.
A contingency table describing the number of each edible
fruit tree according to each vegetation type was then drawn.
This matrix was submitted to a Simple Correspondence Ana-
lysis (CA) using R 3.0.2 (Ihaka and Gentlman, 1996) to deter-
mine the preferred habitat of each WEFT species.
The tree density and Green index were calculated to
analyze the structure of the WEFT species. Tree density (D, in
trees/ha) is the average number of trees per hectare.
Means of Dij values were calculated for each vegetation
types and for each species. An analysis of variance was per-
formed on density values to test whether there is any signi-
ficant difference (at 5% significance level) between vege-
tation types with respect to each WEFT species. When the
differences were significant a Student Newman and Keuls
(SNK) test was applied to separate the means.
Green index (GI) was calculated to detect the spatial
configuration of each WEFT species by considering all indi-
viduals with a diameter at breast height (dbh) ≥ 5 cm of in
each vegetation type, using the formula (Green, 1966):
GI
=
S2
-
1
m
n - 1
(1)
In equation 1, m is the average density of a given WEFT spe-
cies; S2 is the variance of the densities in the different vege-
tation types and n is the total number of trees counted for
this species in a given vegetation type.
Values of Green index vary from 0 (random distribution)
to 1 (aggregative distribution). But this index could display
values that were superior to 1 in the case of large variability
in density values within the plots in the various vegetation
types or when the environment is disturbed (Green, 1966).
The diameter size (diameter at breast height ≥ 5cm)
class distribution (SCD) of each WEFT species was also
established. The 3-parameters of theoretical distribution of
Weibull were then adjusted to the observed distributions
(Johnson, 1970). Its probability density function for a ran-
dom variable x is:
f(x) =
exp -
c
b
x - a
b
x - a
b
c - 1c
(2)
In equation 2, “a” is the location parameter (a = 5 cm, which
is the minimum value of dbh observed in the study area),
“b” the scale parameter, “c” the shape parameter, and exp
[ ] is the exponential function (the base of the natural loga-
rithm). The random variable x is the stem diameter at breast
height and is assumed to be a continuous random variable.
Identification of the most important WEFT species
for riparian populations of Lama Forest Reserve
The use value index (UV) was used to identify the most
important WEFT species. This index was estimated as the
sum of the number of uses mentioned by each informant
for each WEFT species divided by the total number of infor-
mants. For a WEFT species k, the use value UVk was calcu-
lated using the formula (Phillips and Gentry, 1993):
UV
K = ni
k
1
n
n
i = 1
(3)
In equation 3, n is the total number of respondents [n=136
determined using the formula of Dagnelie (1998)], nik is the
number of different uses mentioned by the informant i for
the WEFT species k.
The formula of Dagnelie (1998) is presented as follow:
n
= (1 - α/2)
U2
d
2
p(1 - p)
(4)
In equation 4, n is the total number of respondents, U1-α/2
is the value of the normal random variable for a probability
value of α = 0.05; U1-α/2 ≈ 1.96, p is the proportion of respon-
dents using WEFT species (p≈ 0.68), and d is the expected
error margin of any parameter to be computed from the sur-
vey, which was fixed at 0.08. Under these assumptions, the
sample size used, was n= 136 respondents.
Multipurpose species (high UV) were given priority
compared to less used species (with low UV).
Identification of the priorities Wild edible fruit tree species
for conservation within the Lama Forest Reserve
Compound Ranking System method was used to iden-
tify the priority species. The method uses individual criteria
to rank the species from the most priority to the least prio-
rity. The ranks obtained for each species (according to each
of the criterion) was then summed to obtain the total rank
from which the priority species were identified. Criteria used
included: ethnobotanical value (according to riparian popu-
lation), native status, distribution (within the forest reserve),
and level of threat. The formula and technique used to esti-
mate each of these criteria for the WEFT species are descri-
bed below:
b Distribution
The distribution of the WEFT species was assessed
using the ecological Importance Value Index (IVI). The eco-
logical Importance Value Index (IVI) of a WEFT species i was
computed as follows:
IVI
K
= RD
i
+ RF
i
+ RDoi
(5)
In equation (5) (Assogbadjo et al., 2009):
RDi is the relative density of the species i; RFi is the relative
frequency of the species i; RDoi is the relative dominance of
the species i;
Do
i =
a
i
x N
i
ni
Doi is the dominance of the species i; ai is the basal area of
the species i; Ni is the tree-density of the species i and ni is
the total number of individuals encountered for that species.
Values of IVI range from 0 to 3. The species which had
the lowest IVI was priority for conservation.
b Threats assessment
The threat status is the most used criterion to determine
conservation priority. Most threatened species received the
greatest attention. The red list of IUCN (www.iucnredlist.org)
and Benin red list were used to assess the threat of the WEFT
species. The different categories used were: Endangered
(EN), Vulnerable (VU), Near Threatened (NT), Least Concern
(LC), and Data Deficient (DD). The status of each WEFT spe-
cies was known by using the red list of IUCN and red list of
Benin. When the status of a WEFT species was not found in
the red list of Benin, we checked it in the red list of the IUCN.
Endangered species received greater attention than those
that are not under threat.
b Native status
Since the inventory has both native and introduced
species, priorities were given to native species.
b Ethnobotanical value
The natural forest of the Lama Forest Reserve is a pro-
tected area managed essentially for scientific researches,
but the local population harvest fruit from it either in com-
plicity with the security agents or clandestinely.
The ethnobotanical value of the species refers to the
importance of the WEFT species for the riparian population.
Here we consider the use value as previously developed in
equation 3.
Results
Diversity and ecology of WEFT species
in the Lama Forest Reserve
A total of 10 WEFT species belonging to 10 genera and
8 botanical families were encountered in the four vegeta-
tion types. These include: Dialium guineense Willd. (photo
2), Diospyros mespiliformis Hochst., Drypetes floribunda
Hutch., Mimusops andongensis Bruce, Pterocarpus santali-
noides L’Her. ex De., Pancovia bijuga Willd., Psidium gua-
java L. (photo 3), Lecaniodiscus cupanioides Planch., Ficus
capensis Thumb. and Spondias mombin L. (photo 4). The
Correspondence Analysis revealed that the two first axes
captured 84.6 % of the information contained in the con-
tingency table describing the relationship between WEFT
species and the four vegetation types. Correlations between
the two axes and the WEFT species indicated that all spe-
cies except F. capensis had better correlation (r > 0.50) with
the first axis which was more correlated to the second axis
(r = 0.68). Correlations between the axis and the different
vegetation types, indicated that typical dense forest and
degraded dense forest had better correlation (r = 0.99 and
r = 0.52 respectively) with the first axis whereas the old fal-
low and the young fallow were more correlated with the sec-
ond axis (r = 0.50 and r = 0.69 respectively).
Projection of the WEFT species and the vegetation types
in the same axis system defined by the two first axes (figure
2) showed on axis 1, an opposition between typical dense
forests which was associated with D. guineense, D. mespili-
formis, D. floribunda, M. andogensis and P. santalinoides; and
degraded dense forest associated with P. bijuga, P. guajava
and L. cupanioides. On axis 2, it was also observed an oppo-
sition between fallows which were associated with S. mombin
and degraded dense forest associated with F. capensis.
Structural characteristics of the wild edible fruit
tree species in the Lama Forest Reserve
The analysis of variance revealed significant difference
in tree densities between the different vegetation types
(P < 0.01) in D. guineense, D. mespiliformis, D. floribunda,
L. cupanioides, and M. andogensis but not in F. capen-
sis, P. bijuga, P. guajava, P. santalinoides, and S. mombin
(P > 0.05) (table I). As it could be expected, trend in species
density was also in accordance with results of the Simple
Correspondence Analysis (see previous paragraph). Indeed,
a link was observed between species with low density and
their distribution around axis 2. Actually, the distribution of
S. mombin, P. guajava and F. capensis, on the periphery of
the graph (figure 2) is mostly explained by the low number of
individuals in these species (i.e. their tree density), in each
vegetation type.
In order to depict the spatial pattern of each of the
WEFT species, Green index was calculated. Values of this
index for each WEFT species in each of the different vege-
tation types showed that a majority of the WEFT species
roughly presented a relatively random distribution in the
Photo 2.
Fruit and tree of Dialium guineense collected
in the Lama Forest Reserve.
Photo S. Agbahoungba.
58
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four vegetation types. However some species including
D. guineense, M. andongensis, and P. bijuga presented in
the young fallow, a value of Green index greater than 1 (table
II). The same observation was made in the degraded dense
forest for L. cupanioides and P. guajava and in old fallow for
L. cupanioides and P. bijuga. An aggregative distribution
was observed for F. capensis in the young fallow and for
S. mombin in the typical dense forest (table II).
Distribution in stem diameter classes for each of the
WEFT species (figure 3) showed a right skewed (“inverted J”)
distribution, characteristic of multispecific forest stands or an
uneven-aged structure with the predominance of young indi-
viduals or trees with small diameter. The individuals of WEFT
species with a stem diameter between 6 and 15 cm were
the most represented per hectare, except M. andogensis
for which most individuals diameter was around 32 cm. The
highest difference between these structures were observed
on the scale parameter which fluctuated between 4.4 cm
(P. guajava) and 20.6 cm (M. andongensis). The stem diam-
eter classes of 12 cm and 30 cm were not recorded in P. gua-
java and P. bijuga respectively. The stem diameter class’s
distribution of P. santalinoides was not generated as very few
individuals of that species were recorded in the forest.
Photo 3.
Tree of Psidium guajava collected in the Lama Forest Reserve.
Photo S. Agbahoungba.
Photo 4.
Tree of Diospyros mespiliformis collected
in the Lama Forest reserve.
Photo S. Agbahoungba.
-1.0 -0.5 0.00.5 1.01.5 2.0
-1.5 -1.0 -0.5 0.00.5 1.0
Dim 1 (63.89%)
Dim 2 (20.74%)
Digu
Dime
Drfl
Fica
Lecu
Mian Pabi
Psgu
Ptsa
Spmo
Tdf
Ddf
Yf
Of
Figure 2.
Projection of the ten WEFT species and the vegetation types
on the two axes of the Simple Correspondence Analysis (CA).
Species (in black dots): Fica = Ficus capensis;
Pabi = Pancovia bijuga; Mian = Mimusosps andongensis;
Spmo = Spondias mombin; Psgu = Psidium guajava;
Lecu = Lecaniodiscus cupanoides; Ptsa = Pterocarpus
santalinoides; Drfl = Drypetes floribunda; Digu = Dialium
guineense; Dime = Dyospyros mespiliformis.
Vegetation types (in blue triangles): Tdf = Typical dense
forest; Ddf = Degraded dense forest; Yf = Young fallow;
Of = Old fallow.
60
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3)
FOCUS / WILD EDIBLE FRUIT TREES CONSERVATION
Most used wild edible fruit tree species
for riparian population of the Lama Forest
Reserve and priority species for conservation
D. guineense, P. guajava and S. Mombin
appeared to be the three most used WEFT spe-
cies as regards to their ethnobotanical Use Value
(UV), whereas the other species were less used
by the local populations (table III). WEFT spe-
cies inventoried, were mainly used for food and
medicinal purposes. The fruits of these species
were consumed crude. The leaves, the bark and
roots of D. guineense, P. guajava and only the
leaves of S. mombin were used by the popula-
tion to solve different health problems (table IV).
Assessment of threats on the WEFT species,
revealed that D. guineense and D. mespiliformis
were least concern (LC) species whereas D. flori-
bunda and P. bijuga belonged to data deficient
(DD) category. F. capensis, M. andogensis and
P. santalinoides belonged to the vulnerable (VU)
category. L. cupanioides and S. mombin were
near threatened (NT) species. P. guajava was the
only one endangered (EN) WEFT species (table III).
Assessment of the species origin revealed
that except P. guajava and S. mombin (which
originate from South America), and F. capensis
which is a pantropically distributed species, all
others were native to West Africa.
Table I.
Tree density of each Wild Edible Fruit Tree (WEFT) species per vegetation type in the Lama Forest Reserve.
WEFT species TDF DDF YF OF
Dialium guineense 120.4a ± 4.39 44.1b ± 3.99 17.6c ± 7.46 48.2b ± 4.33
Diospyros mespiliformis 83.0a ± 5.79 15.6b ± 2.50 4.6c ± 2.24 11.7b ± 3.73
Drypetes floribunda 64.2a ± 4.15 2.9b ± 1.86 3.2b ± 3.96 0.6b ± 1.75
Ficus capensis 0.4b ± 1.37 7.8a ± 6.21 0.7b ± 1.87 -
Lecaniodiscus cupanioides 15.2a ± 2.50 7.0b ± 3.35 2.1b ± 1.89 12.4a ± 5.34
Mimusosps andongensis 28.9a ± 3.51 6.6b ± 2.31 4.2b ± 3.61 0.7c ± 1.87
Pancovia bijuga 3.8a ± 1.30 1.7a ± 1.14 2.8a ± 3.26 -
Psidium guajava 0.4c ± 1.37 4.5b ± 3.45 1.1c ± 2.29 8.5a ± 4.09
Pterocarpus santalinoides 0.4 ± 1.37 - - -
Spondias mombin - 1.7a ± 1.61 1.4a ± 1.27 3.5a ± 2.13
TDF = Typical Dense Forest; DDF = Degraded Dense Forest; YF = Young Fallow; OF = Old Fallow.
Mean ± standard error of tree density (trees/hectare).
Number of replicates of D. guineense in TDF, DDF, YF and OF are 317, 116, 20 and 78 respectively. Number of replicates of
D. mespiliformis in TDF, DDF, YF and OF are 202, 41, 13, 19 individuals respectively. Number of replicates of D. floribunda in TDF,
DDF, YF and OF are 162, 7, 9, 1 individuals respectively. Number of replicates of F. capensis in TDF, DDF, YF and OF are 1, 19, 2 and
0 respectively. Number of replicates of L. cupanioides are 40, 18, 6 and 20. Number of replicates of M. andongensis are 80, 16, 8
and 1. Number of replicates of P. bijuga are 10, 8, 8 and 0. Number of replicates of P. guajava are 1, 11, 3 and 12.
Number of replicates of P. santalinoides are 1, 0, 0 and 0. Number of replicates of S. mombin are 0, 4, 8 and 12.
In the same row, means with different letters are significantly different (P < 0.05) using Student Newman Keuls’ test.
Table II.
Green index of each Wild Edible Fruit Trees (WEFT) species
per vegetation type in the Lama Forest Reserve.
WEFT species TDF DDF YF OF
Dialium guineense 0.16 0.35 2.46* 0.38
Diospyros mespiliformis 0.40 0.21 0.44 0.99
Drypetes floribunda 0.26 0.26 0.13 0.00
Ficus capensis 0.00 0.05 1.00* 0.00
Lecaniodiscus cupanioides 0.32 1.16* 0.29 1.92*
Mimusosps andongensis 0.39 0.28 1.44* 0.00
Pancovia bijuga 0.11 0.03 3.03* 1.60*
Psidium guajava 0.00 1.82* 0.50 0.00
Pterocarpus santalinoides 0.00 0.00 0.14 0.00
Spondias mombin 1.00* 0.08 0.14 0.72
TDF = Typical Dense Forest; DDF = Degraded Dense Forest;
YF = Young Fallow; OF = Old Fallow.
Green index greater than 1 are denoted by *.
Ecological importance value index (IVI) revealed
D. guineense and D. mespiliformis as the top two most eco-
logically important WEFT species (IVI = 1.424 and 0.794
respectively) whereas the least important was P. santalinoi-
des (IVI = 0.003) (table III).
All WEFTs species recorded in the Lama Forest reserve
were conserved in-situ.
The compound ranking system method applied on the
uses values of WEFT species revealed that D. guineense,
P. guajava, S. mombin, D. mespiliformis and D. floribunda
were defined as the five top priority species. According to
the native status, the five top species were D. guineense,
D. mespiliformis, D. floribunda, P. bijuga and L. cupanoides.
The compound ranking method applied on the importance
value index showed that the top five species were P. san-
talinoides, F. capensis, P. bijuga, S. mombin, D. floribunda.
The same method applied on the threat status showed that
P. guajava, F. capensis, M. andogensis, P. santalinoides,
L. cupanioides were the top priority species (table III).
The rank of each species (summing rank from individ-
ual criterion) revealed P. guajava, S. mombin, F. capensis,
P. santalinoides, and P. bijuga as the top priority WEFT spe-
cies for conservation in the Lama Forest Reserve (table III).
0
5
10
15
20
25
30
5 _ 15
15 _25
25 _ 35
35 _ 45
45 _ 55
55 _ 65
Frequency
Diameter (cm)
a = 5.00
b = 13.84
c = 1.13
= + 1.08
D. guineense
0
1
2
3
4
5
6
7
8
9
10
5 _ 15
15 _25
25 _ 35
35 _ 45
45 _ 55
55 _ 65
65 _ 75
Frequency
Diameter (Cm)
a = 5.00
b = 20.44
c = 1.29
= + 0.86
D. mespiliformis
0
2
4
6
8
10
12
14
16
5 _ 15
15 _25
25 _ 35
35 _ 45
45 _ 55
55 _ 65
Frequency
Diameter (cm)
a = 5.00
b = 6.76
c = 1.09
= + 3.54
D. floribunda
0
0,2
0,4
0,6
0,8
1
1,2
5 _ 15
15 _25
25 _ 35
Frequency
Diameter (cm)
Observed density
a = 5.00
b = 13.04
c = 1.90
= + 1.42
F. capensis
0
1
2
3
4
5
6
5 _ 15 15 _25 25 _ 35 35 _ 45 45 _ 55
Frequency
Diameter (cm)
a = 5.00
b = 7.88
c = 1.15
= + 2.28
L. cupanioides
0
0,5
1
1,5
2
2,5
3
3,5
4
5 _ 15
15 _25
25 _ 35
35 _ 45
45 _ 55
Frequency
Diameter (cm)
a = 5.00
b = 20.57
c = 1.75
= + 0.18
M. andogensis
0
0,2
0,4
0,6
0,8
1
1,2
1,4
5 _ 15
15 _25
25 _ 35
Frequency
Diameter (cm)
a = 5.00
b = 10.14
c = 1.31
= + 1.24
P. bijuga
0
0,5
1
1,5
2
2,5
3
5 _ 15
15 _25
25_35
Frequency
Diameter (cm)
a = 5.00
b = 4.40
c = 1.56
= + 0.98
P. guayava
0
0,1
0,2
0,3
0,4
0,5
0,6
0,7
5 _ 15
15 _25
25 _ 35
35 _ 45
Frequency
Diameter (cm)
a = 5.00
b = 16.81
c = 1.77
= + 0.68
S. mombin
weibull
Figure 3.
Diameter size class distribution (SCD) of each wild edible fruit tree.
a = 5 cm and represents the minimum value of the diameter at breast height (dbh) observed in the forest,
b is the scale parameter, c is the shape parameter and δ the standard deviation.
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3) 61
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Table III.
Results of Compound Ranking System of the Wild Edible Fruit Trees (WEFT) species inventoried in the Lama Forest Reserve.
WEFT UV Rank WEFT Native Rank WEFT IVI Rank WEFT Threats Rank WEFT Sum Total
species species status species species species of ranks
Dialium
0.64 1
Dialium
Native 1
Pterocarpus
0.003 1
Psidium
EN 1
Psidium
2+2+6+1 11
guineense guineense santalinoides guajava guajava
Psidium
0.49 2
Diospyros
Native 1
Ficus
0.073 2
Ficus
VU 2
Spondias
3+2+4+3 12
guajava mespiliformis capensis capensis mombin
Spondias
0.163 3
Drypetes
Native 1
Pancovia
0.107 3
Mimusosps
VU 2
Ficus
7+1+2+2 12
mombin floribunda bijuga andongensis capensis
Diospyros
0.04 4
Pancovia
Native 1
Spondias
0.178 4
Pterocarpus
VU 2
Pterocarpus
10+1+1+2 14
mespiliformis bijuga mombin santalinoides santalinoides
Drypetes
0.023 5
Lecaniodiscus
Native 1
Drypetes
0.224 5
Lecaniodiscus
NT 3
Pancovia
6+1+3+5 15
floribunda cupanioides floribunda cupanioides bijuga
Pancovia
0.017 6
Ficus
Native 1
Psidium
0.236 6
Spondias
NT 3
Dialium
1+1+10+4 16
bijuga capensis guajava mombin guineense
Ficus
0.01 7
Mimusosps
Native 1
Mimusosps
0.284 7
Dialium
LC 4
Drypetes
5+1+5+5 16
capensis andongensis andongensis guineense floribunda
Lecaniodiscus
0.01 8
Pterocarpus
Native 1
Lecaniodiscus
0.378 8
Diospyros
LC 4
Diospyros
4+1+9+4 18
cupanioides santalinoides cupanioides mespiliformis mespiliformis
Mimusosps
0.01 9
Psidium
Introduced 2
Diospyros
0.794 9
Drypetes
DD 5
Mimusosps
9+1+7+2 19
andongensis guajava mespiliformis floribunda andongensis
Pterocarpus
0.01 10
Spondias
Introduced 2
Dialium
1.424 10
Pancovia
DD 5
Lecaniodiscus
8+1+8+3 20
santalinoides mombin guineense bijuga cupanioides
UV = Ethnobotanical Uses Value; IVI = Ecological Importance Value Index.
Threats: EN = Endangered, VU = Vulnerable, NT = Near Threatened, LC = Least Concern, DD = Data Deficient.
Table IV.
Uses of the Wild Edible Fruit Trees (WEFT) species by the riparian populations of the Lama Forest Reserve.
WEFT species Organs Medicinal uses Food uses
Dialium guineense Fruit Eaten raw
Leaves To cure malaria
Bark To recover the dissolute menses
Root Against spot
Psidium guajava Fruit Eaten raw
Leaves Against dysentery
Bark Against cough
Root To correct bad breath
Spondias mombin Leaves To accelerate the childbirth with herbivorous
Healing of circumcision wound
To improve oligospermy
To cure measlves
To cure pimples
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3) 63
CONSERVATION D’ARBRES SAUVAGES À FRUITS COMESTIBLES / LE POINT SUR…
Discussion
Diversity, ecology and structure of the WEFT species
The study assessed the diversity of WEFT species within
the Lama Forest Reserve. The inventory showed 10 WEFT
species belonging to 10 Genera and 8 plant families in the
Lama Forest Reserve. Actually, the equilibrium disruption of
the Lama Forest Reserve due to the high degradation of its
structure in certain locations (degraded dense forest and
fallows), coming from human’s activities, explained the low
WEFT specific richness (10 species) in this forest (Djodjou-
win and Sinsin, 2007). The high nutritional value, the easy
access and the good organoleptic quality of WEFT species
may also explain their overexploitation in the Lama Forest
Reserve prior to the protection by law (Assogbadjo et al.,
2008; Debella et al., 2011). Results of WEFT species inven-
tory in other ecosystems in Benin revealed 52 species of
WEFT belonging to 45 genera and 37 families in the Pend-
jari Biosphere and 115 species of WEFT species belonging
to 92 genera and 48 families in Bassila Reserve (Vodouhê
et al., 2009; Segnon and Achigan-Dako, 2014). This rela-
tively low diversity of WEFT species in Lama Forest Reserve
could be explained by both the area covered by the forest
(less than Pendjari and Bassila reserve), and the deep clay
soil in the Lama Forest Reserve (>2m in depth), that causes
flood during rainy seasons and cracks in dry seasons, which,
according to Emrich et al. (1999), leads to a relatively poor
flora in forests. The variation in density of WEFT species
from one vegetation type to another may be explained by
the difference in ecological patterns (slope, disturbance,
moisture, soil characteristics) within each vegetation type.
For instance, the plant density increases in an environment
where the optimum conditions are met. The WEFT species
such as: D. guineense, D. mespiliformis, and M. andogen-
sis presented a high trees density (120, 83 and 64 trees/ha
respectively) in the typical dense forest because their eco-
logical requirements were met in that vegetation type. These
results are consistent with the findings of Assongba et al.
(2014) who found that the density of D. guineense is high in
the dense forest compared to the fallows and home gardens.
Similar results were obtained by Lokonon (2008) while eval-
uating the structure of D. guineense and D. mespiliformis
in the Lama Forest Reserve. In addition, the high density of
the WEFT species observed in the typical dense forest may
be explained by the non-degraded status of that vegetation
type which is scarcely disturbed. Unlike that, the density is
relatively low in the degraded dense forest, suggesting low
rate of forest regeneration.
Similar to what Djodjouwin et al. (2011) found,
D. guineense, M. andongensis, and P. bijuga presented a
value of Green index greater than 1 in the young fallow. This
suggests a large variation in densities among plots within
this vegetation type. This could be imputable to the change
in floristic composition and change in environmental condi-
tions. The value of the Green index less than 1 in the typ-
ical dense forest, reflects a relatively homogeneous distri-
bution of the WEFT species in that vegetation type. Similar
spatial pattern was observed by Hounkpèvi (2010) in the
Lama Forest Reserve, where a random distribution pattern of
D. mespiliformis, M. andogensis and Lonchocarpus sericeus
(Poir.) Kunth ex DC. was noticed while evaluating the eco-
logical and structural characteristics of plant communities
within the forest. From the conservation point of view, the
predominance of young individuals, suggested that the spe-
cies are not overexploited by the riparian populations and
they are more preserved in this protected forests (Assongba
et al., 2013).
Among the ten WEFT species, D. guineense and
D. mespiliformis were the most abundant and found together
in the same plot within the different vegetation types. This
may be an indication that these two are co-occurring spe-
cies. However, further investigations in other forests are
needed to test for this hypothesis.
Most used wild edible fruit tree species
Ethnobotanical use values indicated that D. guineense,
P. guajava and S. mombin were the most used WEFT species.
This is consistent with the earlier investigations of Ambé
(2000) in Seguela (Ivory Coast), whereby, D. guineense and
S. mombin were the most frequently used among the 55
WEFT species classified. This could be imputable to their
sweet taste and their availability. The diverse uses of the
abovementioned species clearly explains their high use
values as mentioned earlier by Oryema et al. (2010) and
Belinda et al. (2013). Unfortunately, these species mostly
used by the populations are collected from the major part
of the reserve. Further inclusion in plantation programmes
and in agroforestry systems could help to reduce human
pressure on the Lama Forest Reserve. Besides, these spe-
cies fulfill the three main reasons (contribution as food,
medicinal uses and perceptions on availability) identified by
Assogbadjo et al. (2012) which support peasant ambition to
conserve or grow WEFT species in farmland.
In addition, inclusion of these species in the home gar-
dens, the botanical gardens throughout the country could
also contribute to their ex-situ conservation.
Priority species deserving active conservation
P. guajava, S. mombin, F. capensis, P. santalinoides,
P. bijuga were defined as the top priority WEFT species for
conservation using Compound Ranking System method. In
fact, these species are highly used by the local populations
and are lowly represented in the Lama Forest Reserve. That
conditions confer to them the highest rank among the WEFT
species recorded in this forest. This priority list provides a
strong basis for sustainable management at affordable cost,
as handling a huge number of species may not be cost effi-
cient (Maxted et al., 2006). However, the presence of P. gua-
java and S. mombin, native from South America, in the list of
the WFET species recorded in the Lama Forest Reserve may
be explained by their introduction in the forest by the local
populations who were living in the reserve before the protec-
tion status of the forest. This statement was verified by their
high density found in the fallows that represent the living
64
BOIS ET FORÊTS DES TROPIQUES, 2016, N° 329 (3)
FOCUS / WILD EDIBLE FRUIT TREES CONSERVATION
areas of the local populations. Hence, we recommend more
protection efforts on the fallows and the degraded dense for-
ests since these five WEFT species are predominantly pres-
ent in the fallows and degraded dense forests. In addition
these species could be selected by the forest managers in
the plantation and the Lama’s reserve enrichment program.
The Lama Forest managers are encouraged to promote the
introduction of these species in the local population agro-
forestry system to improve their availability near the popu-
lation, hence less pressure on them in the protected areas.
The ranking approach used in this study is in accordance
with those used by Lawrence et al. (2005) in Cameroun, and
Vodouhê et al. (2009) in Northern Benin. Compound Ranking
System method is easy to verify, flexible and easy to apply
for a large data (Idohou et al., 2013). N’Danikou et al. (2011)
recommended the use of the Compound Ranking System
method for studies of this type. Yet, the method used in the
current study can be updated whenever new information is
collected. As data were not always available for the IUCN sta-
tus of species and conservation strategies, it is strongly rec-
ommended that priorities be reassessed and refined when
more detailed information is available. Brehm et al.(2010)
argue that prioritization of species should be a dynamic pro-
cess and as noted by Heywood (1995), the success of any
method of prioritization will depend, in large part, on the
inclusion of results in conservation activities, and especially
the support of the international community (hardware and
financial) in the preservation of biodiversity.
Conclusion
The present study revealed that Lama Forest Reserve
has ten (10) Wild Edible Fruit Trees (WEFT) species belonging
to 10 genera and 8 botanical family. The ecological inves-
tigation, showed that dense forests (typical and degraded
dense forests) sheltered the majority of the WEFT species.
The prioritization conducted, revealed that the priority WEFT
species were: Psidium guajava, Spondias mombin, Ficus
capensis, Pterocarpus santalinoides, and Pancovia bijuga
and these species are predominantly present in the fallows
(young and old) and degraded dense forest. We recommend
more protection efforts on the fallows and degraded dense
forest to ensure the conservation of these species. In addi-
tion, these species should be included in the enrichment
program of the reserve, as well in the areas surrounding the
central core. But Dialium guineense, P. guajava, and S. mom-
bin would be introduced in the plantation program and the
population’s farms in order to reduce the human pressure
on the reserve. Permanent plots should also be established
in this ecosystem in order to follow the dynamics of these
species’ populations. Together with the current study, addi-
tional studies will help to improve knowledge about these
species’ habitat according to the climatic zones of Benin.
Acknowledgements
Our acknowledgment goes to the University Cooperation for
Development-Belgium (CUD) who financed this study and to the
local population for their help during the data collection phase.
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