American Journal of Plant Sciences, 2019, 10, 1871-1887

https://www.scirp.org/journal/ajps
ISSN Online: 2158-2750
ISSN Print: 2158-2742

Morphological Diversity and Distribution of

Garcinia kola Heckel (Clusiaceae) in Two
Agro-Ecological Areas of Côte d’Ivoire

Any Olivier Komenan1*, Kouamé Kévin Koffi1, Ahou Anique Gbotto2, Moussa Kone3,
Doffou Sélastique Akaffou2, Kouakou Laurent Kouakou1, Kouadio Ignace Kouassi1,
Bi Irié Arsène Zoro1
1
Phytotechnical Unit and Genetic Improvement (PUGI), Training and Research Unit (TRU) of Science of Nature,
Nangui Abrogoua University, Abidjan, Côte d’Ivoire
2
Agroforestry Training and Research Unit, Jean Lorougnon Guédé University, Daloa, Côte d’Ivoire
3
Science of Nature Training and Research Unit, Nangui Abrogoua University, Abidjan, Côte d’Ivoire

How to cite this paper: Komenan, A.O.,

Abstract
Koffi, K.K., Gbotto, A.A., Kone, M., Akaf-
fou, D.S., Kouakou, K.L., Kouassi, K.I. and The purpose of this study is to determine the morphological diversity and
Zoro, B.A. (2019) Morphological Diversity distribution of Garcinia kola Heckel (Clusiaceae) in two preferential agro-
and Distribution of Garcinia kola Heckel
ecological growth areas in Côte d’Ivoire, for the sustainable management of
(Clusiaceae) in Two Agro-Ecological Areas
of Côte d’Ivoire. American Journal of Plant the species. Ninety-four (94) trees of G. kola were sampled in Affery (south)
Sciences, 10, 1871-1887. and Biankouma (west) and characterized on the basis of 13 quantitative cha-
https://doi.org/10.4236/ajps.2019.1010132 racteristics of the fruit, the general appearance of a plant, leaves and seeds.
This study revealed the existence of very significant variability and differences
Received: August 25, 2019
Accepted: October 25, 2019
within the trees sampled for most characteristics. Very strong correlations
Published: October 28, 2019 were found among the characteristics of the fruits. On the basis of these cha-
racteristics, it emerged that the fruits of the Biankouma area are larger than
Copyright © 2019 by author(s) and those of Affery. Ascending hierarchical classification (AHC) structured the
Scientific Research Publishing Inc.
trees into three distinct phenotypic groups based on the following discrimi-
This work is licensed under the Creative
Commons Attribution International nating characteristics: fruit height (Hfr), seed mass (Mgr), trunk diameter
License (CC BY 4.0). (Dm), leaf width (largF), Height of first branch (HtF) and width of the petiole
http://creativecommons.org/licenses/by/4.0/ (larP). These parameters can be used as a basis for selecting and maintaining
Open Access the high variability of G. kola. Analysis of the geographical distribution of
trees, based on the nearest neighbour model, revealed an aggregate distribu-
tion in both areas.

Keywords
Garcinia kola Heckel, Morphological Variability, Geographical Distribution,
Discriminating Characteristics

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A. O. Komenan et al.

1. Introduction
Garcinia kola Heckel (Clusiaceae), known as “petit kola” in Côte d’Ivoire, is one
of the most important non-timber forest product (NTFP) generators. Indeed,
seeds are sought for their stimulating effects, aphrodisiacs, bad cholesterol
cleaners and liver protectors [1]. Seeds are also used in drugs to treat multiple
gastrointestinal and pulmonary conditions [2] [3]. Thus G. kola is used as a re-
medy for the treatment of diseases such as diarrhoea, laryngitis, gonorrhoea,
headaches and gastritis: Garcinia kola bark is also used as a purgative [4]. The
pulp is also consumed. The supply of minerals, vitamins and amino acids con-
tained in these fruits makes them complementary foods, sometimes essential,
during the lean season for local forest populations [1] [5]. About this species, all
parts (from the top to the root) are used by man. It, therefore, provides many
services to a large part of the rural population and provides an additional source
of income. G. kola is one of the forest species of socio-economic interest much
appreciated by local populations [6] because the plant has a good market value
in Côte d’Ivoire; the economic value of seeds per kilogram is between 1. 70 and
4. 25 USD on average [7].
As a result, there is strong anthropogenic pressure on this species. In Côte
d’Ivoire, Ahoussou et al. [8] have compiled a list of 35 useful but endangered
wildlife species. Among these is G. kola (Heckel) which is under permanent hu-
man pressure because it is multi-purpose. The threats to forest resources in Côte
d’Ivoire are worrying and linked to the expansion of agriculture but also to the
development of forestry [9]. In addition, the reforestation policy is aimed at spe-
cies with a high growth rate and major economic interest [8], such as teak (Tec-
tona grandis L. F), framiré (Terminalia ivoirensis A. Chev.), azobé (Lofira alata
Van Tiegh.), makoré (Tieghemella africana P.), etc. This has led to the disap-
pearance and scarcity of a significant number of forest trees, thus relieving Côte
d’Ivoire of its most beautiful species [9]. Thus, more scientific research studies
have recently been carried out in Côte d’Ivoire on this forest resource (G. kola).
The work mainly concerns technologies for the regeneration of the species [10]
and socio-economic interest [6]. Despite this work, several questions remain
unanswered, particularly regarding the structuring, distribution and variability
of G. kola Heckel. 1) Is there a morphological variability of certain characteris-
tics within the species? 2) What is the distribution of G. kola? The main objec-
tive of this study is to characterize Garcinia kola in two agro-ecological areas of
Côte d’Ivoire. The specific objectives are 1) to determine the most discriminat-
ing morphological parameters; 2) to group all individuals in the populations on
the basis of these parameters; and 3) to assess the distribution status of the two
natural populations.

2. Materials and Methods

2.1. Study Environment
This study was carried out in two agro-ecological areas of Côte d’Ivoire for three

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 A. O. Komenan et al.

years, from 2015 to 2018. One is in the west (Biankouma) and the other in the
south (Affery). The choice of these areas was made after several prospecting stu-
dies with wholesale and field merchants. The surveys revealed that most of their
supply of “petit kola” grains originated in these two areas. Biankouma is a de-
partment in the west of Côte d’Ivoire and is part of the Tonpki region. This lo-
cality is located 635 km from Abidjan between 7˚44'00'' North and 7˚37'00''
West. The villages in which the work was carried out are Kanta, Kabakouma and
Blagouin. Like the entire region, our study area is characterized by mountainous
relief, ferralitic and hydromorphic soils. During the year the temperature gener-
ally varies from 17˚C to 33˚C with an average of 24˚C. The rainfall varies be-
tween 1300 and 2400 mm per year and the vegetation consists mainly of humid
forest. Cocoa, rubber and oil palm are the main agricultural export resources,
including coffee from the region, which is very popular.
Affery, the second study area, is located in the south of Côte d’Ivoire in the
department of Adzopé, 101 km from Abidjan between 6˚18'54'' North and
3˚57'37'' West. The villages in which the work was carried out are Daguikoi,
Npokoi, Agbokia and Kossoa. They are located between 4 and 10 Km from Af-
fery. Like the whole region, Affery is located in a humid tropical climate zone, of
the Attiéen type. This climate gives it a relatively constant temperature which
oscillates around 27.5˚C with four seasons of uneven lengths. The annual rainfall
is 1300 mm on average. The town of Affery is characterised by the presence of
many hills whose average altitude does not exceed 100 metres. They are sepa-
rated by long valleys that look like precipices from which several marigots and
rivers sometimes leave. The vegetation is dominated by tropical rainforest. This
vegetation is composed of two types of forests: primary forest, which is similar to
classified forests, and secondary forest, which merges with the vast expanses of
fallow land resulting from shifting cultivation and intense logging. The soils are
mixed, sandy-clayey and suitable for growing coffee, cocoa, rubber and oil palm.
Humid hydromorphic soils suitable for the cultivation of plantain bananas,
sweet bananas, rice, etc. are also found there.

2.2. Device for Morphological Characterization

Morphological characterization was conducted on 94 adult trees in all two study
areas. These trees have produced fruit at least once. They are considered as
adults and therefore make it possible to record all the parameters of the study.
These trees were monitored throughout their breeding season, from flowering to
fruit harvesting, from April to November, to take into account all phenological
stages. From the remarkable flowering stage to the next flowering stage for two
cycles. 13 parameters were selected based on a work of Leakey et al. [11] [12]
Fofana et al. [13] and Towanou et al. [14]. It is about:
− Fruit parameters: Fruit height (HFr), fruit diameter (DiF), fruit mass (maF),
pericarp thickness (EpP), seed box cavity (CLG), seed mass (Mgr), number of
seeds per fruit (Ngr);
− The parameters of the shaft port: shaft diameter (Dm), height of the first fork

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A. O. Komenan et al.

(HtF);
− Leaf parameters: length (LgF) and width (largF) of the leaf, length (LgP) and
width (largP) of the petiole.
On each tree, a minimum of thirty fruits were collected. Given the importance
that owners attach to the tree, fruit is usually picked at the base of the trees.
Thus, to ensure that the fruits are actually from the collected tree, the selected
individuals are separated from each other by at least 25 m. All leaves were col-
lected at the end of the first branch, from the ground, from all trees. Measure-
ments were made on a minimum of thirty leaves per tree.
For data processing, EXCEL and R software were used. The means and analy-
sis of variance made it possible to assess the difference between the parameters
studied. The most discriminating variables and related species were identified.
The statistical tools that are the PCA (Principal Component Analysis) and AHC
(Ascending Hierarchical Classification) make it possible to achieve this objec-
tive. The classification carried out is a hierarchical bottom-up classification on
the principal component and covers all the parameters studied. This classifica-
tion was carried out according to the Ward method taking into account the Euc-
lidean distance matrix. The relationships between the different parameters were
studied on the basis of total correlations.

2.3. Methods for the Study of Distribution

Transects were constructed and an inventory of Garcinia kola trees was carried
out using GPS, in projected coordinate systems (x; y) and UTM (Universal
Tranvers Mercator). It consisted in noting the geographical coordinates of each
individual. Mobile sampling was conducted because the density per hectare of
the tree is low; less than three trees per hectare on average. It consisted of walk-
ing through the environment in all directions, noting all the trees. This invento-
ry made it possible to create a database.
Data analysis was performed using ArcGIS software. The Average Nearest
Neighbor (ANN) tool was used to study the distribution [15]. The ANN meas-
ures the distance between each tree and the location of the nearest neighbouring
tree. He then averages all these distances from the nearest neighbour. If the av-
erage distance is less than the average calculated for a hypothetical random dis-
tribution, the distribution of the entities analyzed is considered aggregated. If the
average distance is greater than the hypothetical random distribution, the enti-
ties are considered dispersed. The average of the nearest neighbour is equal to
the observed average distance divided by the expected average distance (the ex-
pected average distance is based on a hypothetical random distribution with the
same number of entities covering the same total area).
The nearest average neighbor, Average Nearest Neighbor, is given by the fol-
lowing relationship:
Do
ANN =
DE
where Do is the average distance observed between each tree and its nearest

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 A. O. Komenan et al.

neighbor

∑ di
n

Do = i =1
n
And DE is the expected mean distance for the characteristics given in a
random way
0.5
DE =
n A

In the above equations, di is equal to the distance between entity i and the
nearest neighbouring entity; n is the total number of individuals and A is the
area of the minimum rectangle encompassing all entities, or an area value speci-
fied by the user
The z-score of the nearest average neighbour for the statistics is calculated as
follows:
0.26136
SE =
n2 A

3. Study of Morphological Variability

3.1. Characteristics of Tree Fruits
Tables 1-3 present the mean values of the fruit parameters and the result of the
analysis of variance (ANOVA) for all trees sampled respectively within the pop-
ulation and between the two sites.

3.1.1. Fruit Height (HFr) and Mass and Fruit (maF)

The variables fruit height Hfr and fruit mass maF show a significant difference
for all trees in the two sites (Table 1). However, the maF does not show any
variation between sites (Table 3). This parameter has a coefficient of variation of
62.14%. The average weight of fruit in Affery is 146.18 g while in Biankouma it is
307.71 g. The HFr is significantly different from one site to another. The average
height of the fruit is 68.36 mm. The coefficient of variation between the sites for
fruit height is 20 for Affery and 20.48 for Biankouma.

Table 1. Mean tree performance for fruit dimensions and analysis of variance results (F).

Parameters Min Max Average Standard deviation CV (%) F

HFr (mm) 3.68 106.00 68.36 14.37 21.03 **
maF (g) 5.00 625.00 206.32 128.21 62.14 **
DiF (mm) 36.1 88.94 67.51 10.36 15.35 **
EpP (mm) 7.21 37.16 20.05 5.35 26.72 **
CLG (mm) 25.93 64.00 47.46 8.57 18.06 **
Mgr (g) 2.80 85.81 9.34 8.41 90.06 **
Ngr 1.00 4.00 2.08 0.89 43.01 NS

**: significant at the 5% threshold; NS: not significant at the 5% level. NB: Hfr: height of the fruit; maF:
mass of the fruit; DiF: diameter of the fruit; CLG: cavity of the seed box; Mgr: mass of seeds; Ngr: number
of seeds per fruit; EpP: thickness of the pericarp, CV: coefficient of variation.

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A. O. Komenan et al.

Table 2. Mean tree performance for leaf size, tree size, and variance analysis results (F).

Parameters Min Max Average Standard deviation CV (%) F

Dm (cm) 15.28 76.43 44.91 12.75 28.39 **
HtF (m) 1.13 14.00 5.27 2.64 50.24 **
LgF (cm) 6.81 17.42 11.84 2.35 19.89 **
largF (cm) 2.43 15.86 5.04 1.71 33.92 **
LgP (cm) 0.78 3.70 1.38 0.34 24.70 NS
larP (cm) 0.16 0.59 0.22 0.06 26.50 **

**: significant difference at the 5% level; NS: not significant at the 5% level. NB: Dm: diameter of the trunk;
HtF: height of the first branch; LgF: leaf length; largF: leaf width; LgP: petiole length; lgP: width of the pe-
tiole.

Table 3. Average performance of variables for each site and analysis of variance.

AFFERY BIANKOUMA
Variables Average CV(%) Average CV % P
Dm (cm) 48.56 25.35 38.76 28.75 0.07
HtF (m) 5.92 46.37 4.16 49.79 0.11
LgF (cm) 11.22 20.15 12.90 16.70 0.04
largF (cm) 4.60 24.50 5.79 38.28 0.07
LgP (cm) 1.34 15.95 1.45 33.12 0.03
larP (cm) 0.22 27.70 0.24 23.56 0.04
HFr (mm) 65.17 20.00 73.75 20.48 0.04
maF (g) 146.18 62.74 307.71 38.12 0.22
DiF (mm) 64.61 16.01 72.41 11.75 0.04
EpP (mm) 19.12 29.23 21.63 21.29 0.04
CLG (mm) 45.50 18.67 50.78 15.24 0.04
Mgr (g) 9.20 113.87 9.62 28.01 0.02
Ngr 2.03 40.50 2.17 46.75 0.02

NB: The underlined probability values are not significant at the 5% level.

3.1.2. The Thickness of the Pericarp (EpP) and the Diameter of the Fruit
(DiF)
The diameter of the fruit DiF and the thickness of the pericarp EpP are signifi-
cantly different at the 5% threshold for all trees and between sites (Table 1 and
Table 3). The average diameter of fruits DiF for both populations is 67.51 mm.
The coefficient of variation for this parameter is lower for the Biankouma zone,
11.75%, while that of Affery is closer to the general average of 16.01%. There is a
high variability for EpP between sites and for all trees (Table 1 and Table 3).

3.1.3. The Cavity of the Seed Box (CLG)

The results of the analysis of variance indicate that there is a significant differ-
ence in the seed housing cavity between fruits from the same area and between
areas (Table 1). This character varies from 25.93 to 64 mm for the sites com-
bined. The average determined on all trees is 47.47 mm with a coefficient of var-

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 A. O. Komenan et al.

iation of 8.57 for Affery and 18.06% for Biankouma respectively. Between the
sites (Table 3) the cavity of the seed box varies from 45.50 (Affery) to 50.78 mm
(Biankouma). The coefficient of variation between the sites is from 15.24% (Af-
fery) to 18.67% (Biankouma).

3.1.4. Seed Mass (Mgr)

The mass character of the seeds (Mgr) differs significantly between sites and
within the entire sample (Table 1). It varies from 2.8 to 85.81 g for the two study
areas combined with an average of 9.34 g. The overall coefficient of variation of
90.06% is high. However, according to Table 3, the coefficient of variation is
higher in Affery (113.87%) than in Biankouma (28.01%).

3.1.5. The Number of Seeds per Fruit (Ngr)

Analysis of the variance analysis of the number of seeds per fruit (Ngr) for all
trees shows that there are no significant differences (Table 1). The number of
seeds varies from 1 to 4 per fruit for all trees. The average is 2.08 with a high
coefficient of variation of 43.01. The inter-site analysis (Table 3) of the variance
for this characteristic indicates a significant difference. The intersite averages are
between 2.03 (Affery) and 2.17 (Biankouma) with respective coefficients of vari-
ation of 40.50% and 46.75%.

3.2. Characteristics Tree Architecture

Table 2 presents the performance of the leaf and tree wearing variables and the
results of the analysis of variance. Table 3 presents the results of the descriptive
statistics and the analysis of inter-site variance.

3.2.1. The Diameter of the Trees (Dm)

For the trunk diameter characteristic measured at 1.30 m from the ground, the
analysis of variance shows that there is a significant difference for the entire
sample (Table 2). The diameter of the trunk is between 15.28 and 76.43 cm. The
overall average is 44.91 cm. However, the analysis of variance for each site
(Table 3) shows that there is no significant difference for this characteristic. The
coefficients of variation are close (Affery, 25.35% and for Biankouma 28.75%).

3.2.2. The Height of the First Branch (HtF)

The height of the first branch varies from 1.13 to 14 m with an average of 5.27 m
and a high coefficient of variation of 50.24%. The results of the analysis of va-
riance (Table 2) show that there is a significant difference for this characteristic
between trees while there is no difference between sites (Table 3). The coeffi-
cients of variation for the two sites are close (46.37% for Affery and 49.79% for
Biankouma).

3.3. Leaf Characteristics

3.3.1. Leaf Length (LgF) and Leaf Width (largF)
The analysis of variance (Table 2) for all trees shows that there is a significant

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A. O. Komenan et al.

difference for leaf length (LgF) and leaf width (largF). The length of the sheet va-
ries from 6.81 to 17.42 mm. The leaf width varies from 2.43 to 15.86 mm for all
trees and a high coefficient of variation of 33.92%. However, there is no signifi-
cant difference between the sites in terms of leaf width (widthF). For this cha-
racteristic, the values of the coefficient of variation are 24.50% for the minimum
and 38.2% for the maximum. The length of the inter-site sheets (Table 3) varies
slightly from 11.22 to 12.90 mm with a coefficient of variation between 16.70%
and 20.15%.

3.3.2. Petiole Length (LgP) and Petiole Width (largP)

The data in Table 2 show that the average petiole length (LgP) is 1.38 cm and a
coefficient of variation of 24.7%. The values for this characteristic range from
0.78 to 3.7 cm for all trees. There is no significant difference on the length of the
leaf petiole among all trees, unlike the width of the petiole (largP). In addition,
there is a difference between sites for petiole length (LgP). However, the width of
the inter-site petiole (largP) (Table 3) does not differ significantly. The coeffi-
cient of variation for petiole width (largP) is high for both zones combined
(26.5%) and inter-site (23.56% for Biankouma and 27.70% for Affery).

3.4. Structuring Morphological Variability

Correlations between parameters
The correlation matrix between the parameters studied (Table 4) shows that
there are many positive and significant correlations between the fruit parame-
ters. Some correlations between the fruit parameters are as follows:

Table 4. Correlation matrix on all variables studied in Garcinia kola H.

Variables Dm HtF LgF largF LgP larP HFr maF DiF EpP CLG Mgr Ngr

Dm 1

HtF 0.04 1

LgF −0.23 −0.08** 1

largF −0.04 −0.08 0.57**** 1

LgP −0.06 −0,05* 0.30** 0.27** 1

larP −0.14 −0.16* 0.35*** 0.33** 0.07 1

HFr −0.02 −0.13* 0.22* 0.27** 0.06 0.27** 1

maF −0.07 −0.15 0.29** 0.25* 0.13 0.13 0.50**** 1

DiF −0.03 −0.14 0.25* 0.25* 0.09 0.14 0.55**** 0.74**** 1

EpP 0.03 −0.08* 0.19 0.23* 0.10 0.01 0.39*** 0.39*** 0.56**** 1

CLG −0.05 −0.12 0.19 0.16 0.05 0.17 0.42**** 0.65**** 0.86**** 0.06 1

Mgr −0.01 0.04 0.01 0.08 0.04 0.01 0.03 0.05 0.09 0.03 0.09 1

Ngr −0.11 −0.16 0.01 −0.09 −0.08 0.03 0.07 0.30 0.34 0.06 0.38 0.06 1

NB: Underlined values indicate variables between which there is a strong correlation, p < 0.0001 “****”, p < 0.001 “***”, p < 0.01 “**”, p < 0.05 “*”, r value
without “*” are not significative.

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 A. O. Komenan et al.

− Diameter of the fruit (DiF) and the mass of the fruit (maF), with a correlation
coefficient r = 0.74 and p value < 0.0001;
− Seed compartment cavity (CLG) and fruit mass (maF) with a correlation
coefficient r = 0.65 and p value < 0.0001;
− Diameter of the fruit (DiF) and the cavity of the seed box (CLG) with a cor-
relation coefficient r = 0.86 and p value < 0.0001;
− Fruit diameter (DiF) and pericarp thickness (EpP) with a correlation coeffi-
cient r = 0.56 and p value < 0.0001;
− Diameter of the fruit (DiF) and height of the fruit (HFr) with a correlation
coefficient r = 0.55 p value < 0.0001;
− Fruit height (HFr) and fruit mass (maF) with a correlation coefficient r =
0.50 p value < 0.0001;
− Fruit height (HFr) and cavity of seed box (CLG) with a correlation coefficient
r = 0.42 and p value < 0.0001.
The only correlation between the morphological parameters of the leaves is
that between the width (largF) and length (LgF) of the leaf with a correlation
coefficient r = 0.57 with p < 0.0001. No significant correlations are established
between tree parameters, fruit parameters, seed parameters and leaf parameters.

3.5. Multivariate Analyses

Principal Component Analysis (PCA)
The principal component analysis (PCA) covers 08 parameters. Indeed, given
the correlations between the parameters, 5 variables were eliminated to avoid
redundancy. Table 5 presents the main axes of the PCA with their own value.
The results of the principal component analysis (PCA) showed that three axes
have eigenvalues greater than 1. Axis 1 (23.56% of the total variation) is largely
explained by leaf width (0.70), petiole width (0.64) and fruit diameter (0.62).
These parameters are all positively correlated with axis 1. This axis can be de-
fined as the growth axis of leaves and fruits. It makes it possible to distinguish
trees with strong foliar growth and producing large fruits. Axis 2 expresses
15.59% of the total change. The parameters that contribute to the formation of
this axis are the number of grains per fruit (0.66). This axis can be defined as the
axis of grain growth. This axis is used to determine which trees have low and
high production. The third axis expresses 12.80% of the total variability. The
mass of the grains is the only parameter that contributes strongly to the forma-
tion of this axis (0.84). Axis 3 can be defined as the axis of seed performance.
The first two components explain most of the variability revealed by the
quantitative variables studied. The two principal axes express 38.34% of the total
variation.

3.6. Phylogenetic Relationships between Trees

The study of phylogenetic relationships between trees was carried out by means
of ascendant hierarchical classification (AHC). The principle of AHC is to group

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A. O. Komenan et al.

together individuals that have a sufficient degree of similarity in the same set.
The dendrogram resulting from the ascendant hierarchical classification (AHC)
obtained by Ward’s method allowed the identification of two large sets (A and
B) with 25 and 69 trees respectively. These sets are composed of trees from both
populations. A truncation at resemblance level 50 gives a finer classification with
three groups (I, II and III). Group I is identified with set A and groups II and III
are included in set B (Figure 1). Group I contains 25 trees with 16 from Affery
and 9 from Biankouma. Group II is composed of 23 trees including 20 trees
from Affery and 3 trees from Biankouma. Group III was the largest, composed
of 46 trees including 23 trees of each population (Table 6).

Table 5. Eigenvalues, and percentage of variation expressed by the first three axes from the
08 variables.

Variables Axis 1 Axis 2 Axis 3

Dm 0.27 −0.55 −0.12

HtF −0.56 0.06 0.31

largF 0.70 0.25 0.10

LgP 0.42 0.51 −0.22

larP 0.64 −0.23 0.20

DiF 0.62 −0.12 0.05

Mgr 0.08 0.28 0.84

Ngr −0.01 0.66 −0.28

Eigenvalues 1.89 1.25 1.02

% Expressed variance 23.56 15.59 12.80

% Cumulative variance 23.56 39.16 51.95

NB: The underlined values indicate the variables that contribute the most to the formation of the axes.

Table 6. Composition of the three groups (I, II and III) from the Ascending Hierarchical Classification (AHC).

Groups Staff Trees

A1; A26; A33; A40; A42; A48; A51; A54; A66; A68; A82; A84; A86; A91; A103; A110; BK25; BK27; BK29; BK41;
I 25
BK44; BK52; BK58; BK59; BK71

A5; A8; A12; A16; A22; A27; A34; A35; A36; A41; A43; A45; A50; A78; A99; A106; A108; A109; A111; A112;
II 23
BK19; BK39; BK45

A6; A10; A11; A13; A17; A28; A32; A37; A46; A65; A70; A74; A77; A79; A80; A83; A85; A93; A96; A101; A104;
III 46 A105; A107; BK4; BK5; BK7; BK10; BK11; BK12; BK13; BK18; BK34; BK40; BK46; BK47; BK48; BK49; BK56;
BK57; BK61; BK62; BK66; BK68; BK70; BK72; BK74

Values in bold indicate the most representative trees in each group.

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 A. O. Komenan et al.

Figure 1. Dendrogram of the ascendant hierarchical classification (AHC) of trees according to the Ward method; Group I (25 trees),
Group II (23 trees) and Group III (46 trees).

The differentiation between the three phenotypic groups resulting from

AHC was refined by the analysis of variance to a classification criterion
(ANOVA) of all the variables analysed. The results indicate significant differ-
ences (P < 0.05) for all variables except petiole length (LgP) and number of seeds
per fruit (Ngr) (Table 7). Among the six variables that revealed a differentiation
between the three groups, three (Dm, Hfr and Mgr) allowed a complete distinc-
tion. The three (3) other variables (largF, largP and HtF) allowed a partial diffe-
rentiation.
Group I is characterised by trees with low average values for trunk diameter
(34.65 cm) and fruit height (53.27 mm). Individuals in this group showed inter-
mediate mean values for the height of the first branch (6.09 m), leaf width
(widthF) and petiole width (0.21 mm). For this group, the seed mass is relatively
high (7.14 g).
Group II had the highest mean values for trunk diameter (60.47 cm), height of
first branch (6.09 m) and seed mass (8.17 g). No low mean values of the variables
were recorded in this group. However, relatively high values were recorded for
leaf width (4.51 mm), petiole width (0.20 mm) and fruit height (67.69 mm).
In Group III, leaf width (5.55 mm), petiole width (0.24 mm) and fruit height
(76.90 mm) were the variables with the highest average values. On the other
hand, seed mass recorded a low average value (4.49 g). For the individuals in this
group, the height of the first branch is relatively high (4.56 m).

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A. O. Komenan et al.

Table 7. Mean values and standard deviations of the morphological variables analysed in the 3 groups of phenotypic diversity
derived from AHC and analyses of variance (ANOVA).

Group I Group II Group III

Variables F P
(N = 25) (N = 23) (N = 46)

Trunk diameter (Dm) cm 34.65 ± 9.53c 60.47 ± 6.29a 42.17 ± 9.44b 51.92 <0.001***

Height of first branch (HtF) m 6.09 ± 7.31ab 7.34 ± 3.08a 4.56 ± 2.23b 3.32 0.041*

Sheet width (largF) mm 4.59 ± 1.09ab 4.51 ± 0.95b 5.55 ± 2.11a 4.34 0.016*

Petiole length (LgP) mm 1.36 ± 0.23a 1.30 ± 0.19a 1.43 ± 0.43a 1.20 0.306ns

Width of the petiole (larP) mm 0.21 ± 0.04ab 0.20 ± 0.03b 0.24 ± 0.07a 4.01 0.020 *

Fruit height (HFr) mm 53.27 ± 15.25c 67.69 ± 6.26b 76.90 ± 9.10a 40.51 <0.001***

Seed mass (Mgr) g 7.14 ± 2.57b 8.17 ± 2.99a 4.49 ± 2.24c 7.28 <0.001***

Number of seeds per fruit (Ngr) 2.05 ± 0.89a 1.81 ± 0.66a 2.23 ± 0.98a 1.63 0.202ns

For each parameter, the means with the same letters are statistically identical to the threshold P<0.05, according to the ppds test; ns: not significant; ** and
*** indicate highly significant and very highly significant respectively.

3.7. Study of the Distribution of Trees

The analysis of the distribution of Garcinia kola Heckel trees in Affery is shown
in Figure 2, which shows the geographical distribution (A1) of the trees and a
distribution curve according to the Average Nearest Neighbor model. The curve
(A2) indicates an aggregate distribution of trees. According to Table 8, the ob-
served distance between trees in this zone is 84.28 m while the expected distance
is 333.91 m; the ratio of the nearest neighbour is 0.25 with a z-score value of
−14.79. Thus the observed distance less than the expected distance the z-score
less than 0 indicates that the distribution is aggregated. The same is true for Bi-
ankouma (Figure 3 and Table 8) because the observed distance (157.46 m) is
less than the expected distance (320.35 m) and the z-score (−8.42) is less than 0.
For these two zones, there is less than a 1% probability that these models are the
result of chance.

Table 8. Average distances and nearest neighbor values (ANN) by site.

Site DO (m) DE (m) ANN z-score p Area (m2)

Biankouma 157.46 320.35 0.49 −8.42 0.01 30,788,879.98

Affery 84.28 333.91 0.25 −14.79 0.01 47,720,769.21

DO: Distance observed; DE: Expected distance; P: probability; ANN: Average Nearest Neighbor.

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 A. O. Komenan et al.

Figure 2. Cartography and curve of the distribution of Garcinia kola trees in Affery.

Figure 3. Cartography and distribution curve of Garcinia kola trees in Biankouma.

4. Discussion
This study was initiated as part of a program for the sustainable management of
non-timber forest products other than timber. More specifically, it is a pro-
gramme aimed at the domestication of the Garcinia kola (Heckel) species, which
is widely exploited in Côte d’Ivoire for its therapeutic values. This study aimed
to assess the morphological diversity of Garcinia kola H. (Clusiaceae) in two
preferential agro-ecological areas for tree growth in Côte d’Ivoire.
All the characteristics studied revealed a significant difference between the in-
dividuals sampled at the two sites combined. There is great heterogeneity be-
tween Garcinia kola H trees. These results are similar to those of Bationo [16] on
Sclerocarya birrea in Burkina Faso. Similarly, in South Africa and Namibia, [11]
[12]) on Sclerocarya birrea, subsp caffra yielded similar results. Only the length
of the petiole (LgP) and the number of grains per fruit (Ngr) do not differ sig-
nificantly.
The study of intra-population variability revealed high variability for fruit

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A. O. Komenan et al.

diameter (DiF), fruit height (Hfr), leaf width (largF), fruit mass (maF), grain
mass (Mgr) and number of grains per fruit (Ngr). The high coefficient of varia-
tion at Affery for fruit mass (maF) and grain mass (Mgr) indicates greater varia-
bility between fruits in the Affery area. However, in Biankouman, these two
characters have larger average masses, which suggests larger fruits and seeds
than in Affery. This could be explained by the fact that the trees of Garcinia kola
Heckel in Biankouma are found in a much more preserved forest environment,
in a forest that is difficult to access. The number of seeds per fruit (Ngr) on the
scale of all samples does not show any significant difference. However, this cha-
racteristic makes it possible to make the difference on a smaller scale, in a study
area. Some seeds are larger, egg-shaped and elongated; this is believed to be due
either to the influence of the microclimate on seed formation or to the existence
of several Garcinia species. Thus, the mass of seeds and the number of seeds, the
mass of fruits and the number of seeds per fruit would make it possible to gather
as many divergent individuals as possible within the natural populations of G.
kola. These parameters were highlighted by Nafan [17] in Vitellaria paradoxa
(Shea) as the most variable. The same was true for Detarium microcarpum [18].
The study of inter-population variability reveals a significant difference be-
tween most traits, except Dm, HtF, largF and maF. Thus, these four characteris-
tics do not allow the study of diversity between populations. These results are
similar in J. curcas [19]. Indeed, the results of these authors show that the varia-
bility of morphological descriptors is generally greater at the level of individuals
or between individuals in the same population than between populations.
According to the PCA, the highest variabilities in this study are recorded for
grain mass (Mgr), number of grains per fruit (Ngr), leaf width (largF), petiole
width (largP) and trunk diameter (Dm). Concerning fruits, similar results were
obtained in Santalum austrocaledonicum (Santalaceae) [20] and V. paradoxa
[17]. These authors have shown that the size and shape of the fruits make it
possible to identify different phenotypes of these trees. One other author ob-
tained similar results in Andansonia digitata [18]. Moreover, since fruiting is an
important step in the development cycle, it remains influenced by various factors
that would be responsible for variability. These factors can be of environmental,
nutritional or even anthropogenic origin. It is also assumed that the months of
July and August mark the beginning of the fruiting period of Garcinia kola.
However, the rainfall in August is relatively low and could induce a variation in
water absorption and nutrients depending on the location of the trees. Similar
results on baobab (Andansonia digitata) have shown that the width of the leaves
is part of the morphological descriptors that discriminate trees according to their
origin [18].
The ascending hierarchical classification (AHC) revealed three groups of
phenotypic diversity. Furthermore, the independent distribution of trees of dif-
ferent origins (Affery and Biankouma) in the three groups suggests that genetic
diversity was not related to geographical origin. This result would indicate a
phyletic link between the different trees. Six traits were the most relevant to dif-

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 A. O. Komenan et al.

ferentiate the AHC derived groups. These were, according to their discrimina-
tory power, trunk diameter, fruit height, seed mass, leaf width, petiole width and
height of the first branch. Thus, these characteristics would make it possible to
characterise Garcinia kola in a large-scale study.
The study of the distribution of trees within the two populations (Affery and
Biankouma) revealed an aggregate distribution of trees. The average distance
observed between trees is smaller at Affery (84.28 m) than at Biankouma (157.46
m). These results are similar to those obtained in Strombosia sheffleria [21]. The
aggregate spatial distributions of some tree species can be interpreted as reflect-
ing variations in environmental characteristics [22] [23]. These species will ag-
gregate in areas where environmental conditions are favourable for their devel-
opment [24]. On the other hand, the mode of dispersion may also explain the
aggregation. The limitation of dispersion also results in an aggregate geographi-
cal distribution [25] often observable for tropical tree species [26]. The aggregate
distribution of Garcinia kola H. would be due either to sarcochores, i.e. totally or
partially fleshy diasporas, or ballochores, i.e. expelled by the plant itself during
the spread. Indeed these types of diasporas, which cannot ensure long-distance
dispersal, can give species an aggregated spatial structure made by rodents [21].

5. Conclusion
This work has enabled us to gather information on the level and structure of the
morphological diversity of Garcinia kola Heckel in two agro-ecological zones in
Côte d’Ivoire. The geographical distribution of the trees of the two populations
revealed an aggregated structure. The evaluation of the morphological diversity
of G. kola made it possible to highlight the most discriminating descriptors. In-
deed, at the fruit level, it is the height of the fruit and the mass of the grains that
are the most discriminating. At the foliar organ and tree level, it was the width of
the leaves and trunk respectively that revealed greater variability in the trees. The
ascending hierarchical classification (AHC) has given details on the approxima-
tion of trees by structuring them into three groups of phenotypic diversity. The
differentiation of these groups is based on six most discriminating parameters.
These can be classified according to their order of discriminating power. These
are trunk diameter, fruit height, seed mass, leaf width, petiole width and height
of the first branch. In this way, the representatives of the three groups can be
used to set up a wood yard to bring together most of the diversity of G. kola in
Côte d’Ivoire. But since morphological markers are subject to variations related
to the tree growing environment, we intend to extend this study to molecular
analysis. Microsatellite primers have been selected to conduct this step. The
combined results will make it possible to propose a strategy for the sustainable
conservation of the resource.

Acknowledgements
The authors thank the village populations and the landowners of the study areas

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A. O. Komenan et al.

without which no data would be obtained. Thank you also to the research college
of Genetics Laboratory of Nangui Abrogoua University.

Conflicts of Interest
The authors declare no conflicts of interest regarding the publication of this pa-
per.

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