Biodivers Conserv (2008) 17:2735–2755

DOI 10.1007/s10531-007-9279-y

ORIGINAL PAPER

Diversity and biovolume of tree species in natural forest

ecosystem in the bitumen-producing area of ondo state,
Nigeria: a baseline study

V. A. J. Adekunle Æ A. O. Olagoke

Received: 16 May 2007 / Accepted: 23 October 2007 / Published online: 17 November 2007
Ó Springer Science+Business Media B.V. 2007

Abstract Tree species diversity, biovolume and forest stand structure were investigated
in natural forest ecosystem located around some selected communities in the bitumen-
producing area of Ondo state, Nigeria. Two forest reserves and four free areas distributed
in 4 Local Government Areas (LGA) of the state were selected for data collection, out of
the total 6 LGAs in the bitumen belt of the state. The two forest reserves are Oluwa FR at
Legge in Odigbo LGA and Eba Island FR in Ese-Odo LGA while the four free areas are
located close to each of Omotoso community (Odigbo LGA), Ode-Aye community
(Okitipupa), Igbo-Egunrin community (Ilaje) and Igbotako community (Okitipupa). Eight
plots of equal size (20 9 20 m) were located in each of the selected location, using
systematic line transect sampling design. In each plot, all living trees with dbh C 10 cm
were identified with their botanical names and their dbhs were also measured. The results
of the study reveal that there were ninety nine (99) tropical hardwood timber species
(range: 21 to 48 species per selected forest). These species were distributed among twenty
nine (29) families. While Funtumia elastica has the highest population distribution across
the selected communities’ forest, Euphorbiaceae was the dominant family in the entire
area. Although there was a moderate variation in the biodiversity indices among the
selected communities’ forest, the Shannon-Weiner diversity index of H1 = 4.02 and spe-
cies evenness of E = 0.88 were obtained for the entire study area. Tree density summing up
to 2,740 trees/6 ha varied moderately, with a range of 361–609 tree/ha, among the com-
munities. Though most of the trees encountered belonged to the lowest diameter size class,
the mean basal area and biovolume were 26.69 m2/ha and 262.36 m3/ha respectively.
Recommendations guiding the decision on the allocation of the communities’ forest to the
bitumen exploratory industries are made and the need for good forest management of
the prospective area of bitumen exploration of Ondo state is emphasized. This is to prevent
the imminent loss of biological diversity that would eventually accompany the exploration.

V. A. J. Adekunle (&)
A. O. Olagoke

Department of Forestry and Wood Technology, Federal University of Technology, Akure,
Ondo State, Nigeria
e-mail: vajadekunle@yahoo.com
A. O. Olagoke
e-mail: woleola@justice.com

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Keywords Tree Diversity
Biovolume
Forest stand Structure
Bitumen exploration

Abundance
Dominance
Biodiversity indices
Forest management

Introduction

Evidence has shown that the tropical rainforest ecosystem is among the most complex and
species-rich single ecosystem of the world (Bhat et al. 2000 and Gillespie et al. 2004) and
there is a wide variation in the composition and abundance of species between various
tropical forests (Parthasarathy 2001). According to Cannon (1998), the diversity of trees is
fundamental to total rainforest biodiversity. In fact, the canopy system and favourable
environmental conditions of tropical rainforests further increases the diversity. Also, there
has been indication that 70–90% of living flora and fauna depend on trees for survival in
the rainforest ecosystem (Tilman and Lehman 2001). Forest biodiversity provides a wide
array of goods and services, from timber and some non-timber forest resources to playing
an important role in mitigating climate change as carbon sinks. It also has important
economic, social and cultural roles in the lives of many indigenous and local communities
that even our very opportunity to exist as humans depends on the very capacity of the
forests to support living things (CBD-UNEP 2005). However, the high species diversity
and the notable number of goods and services obtainable from this ecosystem are partly
responsible for the pressure to which it has been subjected for centuries, and which is
presently on the increase (Onyekwelu et al. 2005).
The earth is undergoing rapid environmental changes because of human actions
(Houghton et al. 1996). Humans have greatly impaired the rates of supply of the major
nutrients and thus constrain the productivity, composition, and diversity of terrestrial
ecosystems (Tilman and Lehman 2001). Forest degradation is now becoming increasingly
popular in all major blocks of the remaining tropical forests as indicated by Verissimo
et al. (1995), Coomes et al. (2000) and Kammesheidt (2002). Forest biodiversity is being
lost due to rapid deforestation, fragmentation, and degradation of all forest types. NEST
(1991) reported that over 30,000 ha of forest and natural vegetations are being lost
annually in Nigeria and for this reason, several individuals, national and international
organizations have shown a great concern in biological diversity conservation (Adekunle
2005).
According to Dudley and Stolton (2002), mineral exploration has played a substantial
role in forest destruction in many parts of the world, Nigeria inclusive and this has led to a
number of problems. Mineral prospectors have caused serious damage to the forests in the
process of their search and processing of these minerals (Olagbende 1997), as these often
times require vegetal clearance and digging of trenches which sever the roots of trees and
so make them liable to windthrow. Younger Granite province around Jos, Plateau State,
Nigeria where mining of tin and columbite have led to the destruction of landscape and
vegetation cover greatly exemplified the deleterious impact of mineral exploration on the
environment. Similar damage is done to almost all mining districts, including at Ewekoro,
Ado-Ekiti, Ise-Ekiti, Sagamu, Ukpilla, Jakura, Ashaka, Enugu, Abakiliki, etc, all in Nigeria
(Adeyemo and Jegede 2002). Likewise, forest growth and value have been tampered with
during petroleum exploration and exploitation activities in the Niger-Delta area of Nigeria
as a result of oil spillage; this has resulted in a number of community clashes with the
prospector in the area (Olagbende 1997). Allowing mineral prospecting and exploitation in
the forest reserves is, however, tantamount to the removal of appropriate ecological and

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environmental conditions that are required for optimal agricultural productivity (ENS
2003).
Biodiversity assessment has been recognized by international policy processes such as
the Convention on Biological Diversity, as inevitable tool guiding biodiversity conserva-
tion (Margules and Pressey 2000; Phillips et al. 2003; Royal Society 2003). According to
Slik et al. (2003), floristic analyses are very useful for identifying spatial patterns in plant
diversity and composition. Quantitative floristic inventories have been used in recent years
to characterize forest vegetation throughout the tropics, though many of the investigators
were interested in documenting the structure and floristic composition of forest commu-
nities (Smith and Killeen 1995). Adekunle (2005) reported that tree species composition,
size and age distributions and their horizontal and vertical order were found to be the key
values to determine diversity in a forest stand.
The massive vegetal clearance expected to accompany the commencement of bitumen
exploration and dearth of recent information on the diversity and biovolume of tree species
in this area brought the impetus for the floristic, structural and quantitative description of
the selected communities’ forest. This paper, aimed at characterizing and comparing this
communities’ forest, reports the patterns of diversity and biovolume of tree species in the
bitumen-producing area of Ondo state, Nigeria and recommend appropriate measure to
ensure biodiversity conservation during mineral exploration. So this study provides
baseline information prior to exploration of bitumen in these communities’ forest.

Methodology

The study area

This study was conducted in forests around each of six bitumen-producing communities in
Ondo State within the tropical lowland rainforest ecological zone of southwest Nigeria.
Two forest reserves and four free areas distributed in 4 Local Government Areas (LGA)
were selected for data collection, out of the total 6 LGAs in the bitumen belt of the state.
The two forest reserves are Oluwa FR at Legge in Odigbo LGA and Eba Island FR in
Ese-Odo LGA while the four free areas are located close to each of Omotoso community
(Odigbo LGA), Ode-Aye community (Okitipupa), Igbo-Egunrin community (Ilaje) and
Igbotako community (Okitipupa) (Table 1).

Table 1 Communities, type of forest assessed and their location coordinates

Community name and the type of forest assessed. Location mid-point coordinates

Latitude Longitude Easting Northing

Atijere (Eba Island forest reserve) 60 251N 40 311E 667739 709447

0 1 11
Igbo-Egunrin (Free area forest) 6 27 30 N 40 371E 678789 713167
Igbotako (Free area forest) 60 341 3011N 40 381 E 680588 726995
Legge (Oluwa forest reserve) 60 531 3011N 40 441 E 691524 762051
Ode-Aye (Free area forest) 60 351 3011N 40 451E 693485 728881
Omotoso (Free area forest) 60 441 3011N 40 391E 682370 745431

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N
Nigeria
W E

S Map of Africa

Ondo State

Map of Nigeria

1#
2#
4# #3
6 #5
# LEGEND
1. LEGGE (OLUWA FOREST RESERVE)
2. OMOTOSO FOREST
3. ODE-AYE FOREST
4. IGBOTAKO FOREST
100000 0 100000 Meters 5. IGBO-EGUNRIN FOREST
6. ATIJERE (EBA FOREST RESERVE)
Scale: 1:2000000
OR 1CM TO 20 KILOMETERS

Fig. 1 Map of Ondo State showing the location of the forests with maps of Africa and Nigeria in inset

The climate and site conditions

Characteristic of all the study sites is two distinct seasons (rainy and dry seasons), with
frequent rainfall that normally starts in March and ends in November. The annual rainfall
ranges from 1,700 to 2,200 mm. The dry season is experienced from December to
February. Mean annual temperature falls between 26°C and 28°C while the average daily
humidity is 80%.
The soils are predominantly ferruginous tropical soils and are typical of the variety
found in the intensively weathered areas of basement complex formations in the rainforest
zone of Southwest in Nigeria (Onyekwelu et al. 2005).

Data collection

Systematic sampling design (systematic line transect) was employed for the laying of plots
(Fig. 1). Two transects of 1000 m in length with a distance of at least 500 m between them
were laid in each of the study sites. Sample plot of 20 X 20 m in size was laid in alternate
along each transect at 250 m interval and thus summing up to 4 sample plots per 1000 m
transect and a total of 8 sample plots per study site. All living trees with dbh C 10 cm were
identified and their dbh were measured in every plot. All computed values were extrap-
olated for a hectare.

Tree species identification

The botanical name of every living tree that was encountered in each sample plot was
recorded for each of the study sites. In cases tree’s botanical name was not known

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immediately, such a tree was identified by its commercial or local name. Such commercial
or local names were translated to correct botanical names using Gbile (1984) and Keay
(1989). Trees that could not be identified were tagged ‘unknown’. Specimens of such
unknown trees were collected and preserved for the identification in the herbarium. Each
tree was recorded individually in the field forms and possible effort was made not to omit
any eligible stem in a sample plot.

Computational and statistical procedures

Basal area calculation

The basal area of all trees in the sample plots in the six study sites was calculated using the
formula:
pD2
BA ¼ ð1Þ
4
Where BA = Basal area (m2), D = Diameter at breast height (cm) and p = Pie (3.142).
The total basal area for each of the sample plots was obtained by the sum of the BA of
all trees in the plot while mean BA for the plot (BAp ) was obtained by dividing the total BA
by the number of sample plots (i.e. 8 plots).
Basal area per hectare was obtained by multiplying mean basal per plot with the number
of 20 9 20 m plots in a hectare (25).
BAha ¼ BAP  25 ð2Þ
Where BAha = basal area per hectare.

Volume calculation

The volume of individual trees was estimated using the equation developed for trees
volume estimation in the lowland rainforests by FORMECU (1999). This equation is
expressed as follows:
V ¼ e8:433þ2:331LnðDÞ ð3Þ
Where V = Volume of tree (m3) and D = dbh (cm).
Total plot volume was obtained by adding the volume of individual trees encountered in
the plots. Mean volume for sample plots was calculated by dividing the total plot by the
number of sampling plots (8).
Volume per hectare was obtained by multiplying mean volume per plot V P with the
number of 20 9 20 m plots in a hectare (25).
Vha ¼ V P  25 ð4Þ

Tree species classification and diversity indices

All trees were assigned to families and number of species in each family was obtained for
tree species diversity classification. Frequency of occurrence was obtained for species

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abundance/richness. This was repeated for all plants encountered in the sample plots for the
six sites. The following biodiversity indices were used to obtain tree species richness and
evenness within the forest in each community. They were also used as indices for com-
paring biodiversity among the communities’ forest.
I. Species relative density (RD) was obtained using the formula given by Oduwaiye
et al. (2002):
hn i
i
RD ¼  100 ð5Þ
N
Where RD = relative density, ni = number of individuals of species i and N = total
number of individuals in the entire population.
II. Relative dominance was obtained using the formula given by Brashears et al. (2004):
P
ð BAi  100Þ
RDo ¼ P ð6Þ
BAn
Where RDo = relative dominance, BAi = basal area of all individual trees belonging to a
particular species i and BAn = stand basal area.
III. Community diversity was obtained using a mathematical formula that takes into
account the species richness and abundance of each species in the ecological community.
The equation for the Shannon-Wiener diversity index given by Price (1997) that was used
is:
X
S
H1 ¼ pi Lnpi ð7Þ
i¼1

H1 is the Shannon diversity index, S is the total number of species in the community, pi is
the proportion of a species to the total number of plants in the community and Ln is the
natural logarithm.
IV. Species evenness (E) in each community was determined using Shannon’s equi-
tability (EH) as stated by Kent and Coker (1992):
H1
E¼ ð8Þ
LnðSÞ
S is the total number of species in each community.
V. Sorensen’s species similarity index (SI) of Nath et al. (2005) was used to compare
diversity across the different communities selected for the study.


2C
SI ¼  100 ð9Þ
aþbþcþdþeþf
C is the total number of species in six communities (i.e. aggregate of all species
encountered in the entire study area); while a, b, c, d, e and f are number of species at
communities 1, 2, 3, 4, 5 and 6, respectively.

Test of significance

One-way analysis of variance (ANOVA) was used for test for significant difference
between tree growth variables obtained in each study site. These variables are trees per

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hectare, tree species and family richness, mean dbh of trees, basal area and volume. The
analysis was performed using SPSS for windows 13.0 (SPSS 2003).

Results

Species diversity, evenness and similarity indices in the bitumen-producing area of

Ondo state

The total number of tree species (C10 cm dbh) encountered when data from the selected
communities’ forest were pooled was 99 species in 82 genera and 29 families. Number of
species (species richness) encountered varied significantly (Table 2). The overall Shannon-
Wiener diversity index was H1 = 4.02 and species evenness was E = 0.88. Shannon
diversity index and equitability varied moderately. Oluwa FR at Legge (3.42) had the
highest species diversity index. This is followed by free area forests at Igbotako (3.26),
Omotoso (3.21) and Ode-Aye (3.16), Eba Island FR at Atijere (2.76) and the least, Igbo-
Egunrin free area forest (2.46). Species evenness result were similar in all the selected
communities’ forest, expect for Igbo-Egunrin which had the least value of 0.77. The
highest E value of 0.91 was recorded for Atijere. Sorensen’s species similarity index (SI)
obtained for the entire communities’ forest was 97. Of the 82 genera, Ficus with 4 species
had the highest number of species. Other genera with more than one species include Cola
(3 species), Sterculia (3), Alstonia (2), Dialium (2), Diospyros (2), Uapaca (2), Antho-
cleista (2), Guarea (2), Trichilia (2), Albizia (2), Pterocarpus (2) and Celtis (2).

Family composition

A total of 29 families were encountered (Table 5). The families, Euphorbiaceae and
Moraceae with 10 species (10%) each dominated the forest canopy, followed by Meliaceae
and Sterculiaceae (9 species each). Apocynaceae, Caesalpinoidae, Mimosoidae, Papilio-
noidae and Rubiaceae had 5 species each, Anacardiaceae, Annonaceae, Guttiferae,
Sapotaceae and Ulmaceae had 3 species each, Bombacaceae, Ebenaceae, Loganiaceae,

Table 2 Summary of the tree growth variables obtained for some selected forests in the bitumen-producing
communities of Ondo State
Variables Communities All sites

Omotoso Legge Ode-Aye Igbo-Egunrin Atijere Igbotako

Number of trees/ha 361 609 380 423 541 426 2740

Basal area/ha (m2) 12.13 54.36 16.73 19.19 28.66 28.30 26.46
Volume/ha (m3) 117.47 572.90 164.93 172.36 259.30 287.19 262.36
Number of families 21 22 19 19 15 17 29
Number of species 35 48 36 25 21 39 99
Mean Dbh (cm) 16.58 27.16 18.84 20.92 22.82 23.33 21.60
Dominant Dbh (cm) 130 130 132.5 99.6 88.1 120 132.5
Shannon diversity index 3.21 3.42 3.16 2.46 2.76 3.26 4.02
Species evenness 0.90 0.88 0.88 0.77 0.91 0.89 0.88

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Myristicaceae, Sapindaceae and Verbenaceae had 2 species each while Agavaceae,

Bignonaceae, Burseraceae, Combretaceae, Dichapetalaceae, Ochnaceae, Palmae, Rutaceae
and Simaroubaceae were only represented by 1 species each.
The result of the relative dominance indicated that Apocynaceae (21%) was the dom-
inant family. Other co-dominant families are Euphorbiaceae (15.11%), Sterculiaceae
(12.02%), Papilionoidae (5.84%), Ochnaceae (5.76%) and Annonaceae (5.53%). Palmae
(0.02%) was the least of all the families encountered (Table 5).

Forest stand density, basal area and biovolume

An average of 457 stems/ha was obtained in the entire study area. Oluwa FR at Legge
(609) had the highest tree density, followed Eba Island at Atijere (541), Igbotako forest
(426), Igbo-Egunrin forest (423) and Ode-Aye forest (380) while the least was recorded in
Omotoso forest (361). There was no significant difference (P B 0.05) in the number of tree
encountered per hectare (tree density) among the selected communities (Table 3). In
contrast, the tree basal area differed significantly (P B 0.05) among the communities’
forest. Except for Oluwa FR, which had the highest BA/ha (54.36 m2/ha), the values
obtained for all other forests were similar. The mean basal area was 26.69 m2/ha while the
least was obtained for Omotoso forest (12.13 m2/ha). The volume of trees followed a
similar pattern with basal area. It varied significantly (P B 0.05) among the forests selected
for this study, with a mean of 262.36 m3/ha (Table 2). There was no significant difference
(P B 0.05) in the volume of trees in the forests located at Igbotako, Atijere, Igbo-Egunrin,
Ode-Aye and Omotoso communities while significant difference was discovered to exist
between Oluwa FR at Legge when compared with others, as this forest had the highest
volume/ha (572.90 m3/ha).

Population structure of the communities’ forest vegetation

The population structure of the forest vegetation in each of the selected communities is
represented in Fig. 3a–f.
The structure of all the selected communities was typical inverted J-shaped curve. Basal
area distribution in various diameter size class was similar in four communities while Igbo-
Egunrin and Atijere, which had no [100 cm dbh tree, were different from others.

Stem diameter distribution and density

Tree density consistently deceased with increasing stem diameter size from 10 cm to
100 cm (Fig. 3). The lowest size class (10–20 cm) captured 57.65% of the total forest
stand density and about 8% were above the recommended dbh value of 48 cm for trees to
be merchantable. Mean dbh of trees in the selected communities differed significantly
(P B 0.05). There were no significant difference in mean dbh obtained at Omotoso forest,
Ode-aye forest, Igbo-Egunrin forest, Eba Island FR at Atijere and Igbotako forest and that
of Eba Island forest, Igbotako forest and Oluwa FR at Legge (P B 0.05) (Table 3). Large
dimensioned trees whose dbh are 100 cm were not encountered in Eba island FR at Atijere
and Igbo-Egunrin forest. Dominant dbh had greatest value in Ode-Aye forest (132.5 cm
dbh), followed by Omotoso forest and Oluwa FR at Legge (130 cm dbh each), Igbotako

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Table 3 One-way ANOVA test of significance in some growth variables among the selected communities’
forest
Variables Source of variation Sum of squares df Mean square F Sig.

Basal area Between groups 9069.75 5 1813.95 4.84 0.002

Within groups 14241.16 38 374.767
Total 23310.91 43
Volume Between groups 1085584 5 217116.9 3.822 0.007
Within groups 2158626 38 56805.96
Total 3244211 43
No. of trees/ha Between groups 344161.9 5 68832.39 1.688 0.161
Within groups 1549531 38 40777.14
Total 1893693 43
Mean tree dbh Between groups 503.646 5 100.729 2.880 .027
Within groups 1329.046 38 34.975
Total 1832.692 43

forest (120), Igbo-Egunrin forest (99.6) and the least (88.1) was recorded in Eba Island at
Atijere.

Distinction in species composition and abundance among the selected communities

There was a considerable variation in the population density of the most abundant tree
species among the selected communities (Table 3). Funtumia elastica was the most-
abundant in Omotoso and Igbotako with 34 and 66 trees/ha respectively. Ricinodendron
heudelotii (75), Spondias mombin (66), Millettia thonningii (141) and Lophira alata (94)
had the highest population in Oluwa FR at Legge, Ode-Aye forest, Igbo-Egunrin forest and
Eba Island FR at Atijere respectively. The abundant species were well represented,
occasional or rare in all the selected communities. Funtumia elastica (most abundant) was
followed by Dialium guineensis (31), Macaranga barteri (28), Alstonia boonei, Aningeria
robusta and Pycnanthus angolensis (each with 25 trees/ha), and Gmelina arborea (22);
these together formed 52.63% of the stand density in Omotoso forest. In Oluwa FR at
Legge, Ricinodendron heudelotii (most abundant), Celtis zenkeri (66), Cleistopholis patens
(35), Cola millenii (34), Funtumia elastica (28), Mitragyna ciliata and Staudtia stipitata
(each with 25 trees/ha) accounted for 47.29% of the forest stand density. Spondias mombin
(most abundant) which was followed by Albizia ferruginea (41), Sterculia tragacantha
(25), Funtumia elastica (22) and Pycnanthus angolensis (19) formed 45.53% of Ode-Aye
forest stand density. Millettia thonningii, which was the most abundant, formed with
Funtumia elastica (56), Anthocleista vogelli, Cleistopholis patens and Pycnanthus
angolensis (each with 31 trees/ha) 68.56% of the forest stand density in Igbo-Egunrin
forest. While Lophira alata (most abundant) coupled with Uapaca heudelotii (56),
Millettia thonningii (44), Malacantha alnifolia (39), Baphia nitida and Diospyros mes-
piliformis (38 trees/ha each), Canthium subcordatum and Hannoa klaineana (31 trees/ha
each), Piptadeniastrum africana and Rothmania hispida (25 trees/ ha each) captured
67.47% of the forest stand density in Eba Island FR at Atijere, Funtumia elastica which
was the most abundant, in addition with Anthostemma aubryanum (38), Trema orientalis

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Table 4 Mean separation for growth variables with DMRT

Community Variable mean

Basal area Volume Mean tree dbh No. of trees/ha

Omotoso 12.13a 117.47a 16.58a 361a

a a a
Ode-Aye 16.73 164.93 18.84 380a
a a a
Igbo-Egunrin 19.19 172.36 20.92 423a
a a b
Igbotako 28.30 259.30 23.33 426a
a a b
Atijere 28.66 288.34 22.82 541a
b b b
Legge 54.36 572.90 27.16 609a
Mean with the same alphabets in columns are not significant (P B 0.05)

(25), Pausinystalia talbotii and Pycnanthus angolensis (with 22 trees/ha each) accounted
for 40.61% of Igbotako forest stand density (Table 4).

Discussion

The richness of tree species in forests located in the bitumen-producing area of Ondo state
is noteworthy. For tree species diversity in the area as a whole, high value of species
richness (99), distributed among 89 genera and 29 families were present in this study area.
The breakdown of species richness in each of the forests selected for this study revealed
that the number of species present are 21, 25, 35, 36 and 39 for Eba Island FR at Atijere,
Igbo-Egunrin, Omotoso, Ode-Aye and Igbotako free area forests in that order. These
values were lower in comparison with the values reported by Lowe (1997), Onyekwelu
et al. (2005) and Adekunle (2006) for other rainforests in Nigeria. This could indicate the
extent to which man has previously and presently degraded the forest vegetation in these
communities, since species diversity of a given forest area besides climate and geographic
location, would largely depend on the extent of human interaction in the past and present
(Parthasarathy 2001). In contrast, species richness of 48 obtained for Oluwa FR at Legge is
higher than that of 45 reported for other forests by these workers. Eba Island Forest
reserve, because of its environmental conditions—a riparian forest (surrounded by
water)—that could only permit the survival of few species adapted to such conditions,
possessed the least number of species. Other places with lower species richness were in the
free area where logging, farming and other form of encroachment were not controlled.
Although, forest reserves are under the control of the state government and thus logging
and entrance are protected under the state forest laws, the fate of this ecosystem when the
exploration of bitumen commences is uncertain as the exploration would also be done with
the permission from the same government that owns the forest reserves (Table 5).
Parthasarathy (2001), Guo et al. (2003), Onyekwelu et al. (2005) and Adekunle (2006)
have in the recent time considered the use of Shannon diversity index for characterizing
community diversity in the tropics. The overall Shannon diversity index obtained for the
entire bitumen–producing area (4.02) is higher than the general limit of 1.5–3.5 reported by
Kent and Coker (1992), but compares well with 3.898 reported by Parthasarathy (2001).
Though all the values fall within the general limit, there is still a considerable variation in
this index among the selected communities’ forest. The overall Shannon’s equitability of
0.86 obtained in this study is higher than 0.66 reported by Onyekwelu et al. (2005) for

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 Table 5 Diversity and relative dominance of tree families encountered in selected communities’ forest in the bitumen-producing area of Ondo state
S/N Family Omotoso Legge Ode-Aye Igbo-Egunrin Atijere Igbotako

No. of RDO No. of RDO No. of RDO No. of RDO No. of RDO No. of RDO
Species Species Species Species Species Species

1 Agavaceae – – 1 1.8865 – – – – – – 1 0.3043

2 Anacardiaceae 1 1.6641 1 1.7537 2 6.3411 1 3.7927 – – 1 0.1715
3 Annonaceae 1 1.0797 1 13.306 1 9.1135 1 5.1238 1 0.4403 2 4.1156
4 Apocynaceae 4 48.5993 3 19.33 3 9.9886 1 13.9954 1 0.6401 3 33.5457
5 Bignonaceae – – – – 1 0.9977 1 0.7385 – – – –
6 Bombacaceae – – 1 0.2307 – – – – – – 1 4.0232
Biodivers Conserv (2008) 17:2735–2755

7 Burseraceae – – – – 1 1.6074 1 1.9659 – – – –

8 Caesalpinoidae 3 5.2856 2 2.0574 2 0.749 1 1.6527 – – – –
9 Combretaceae 1 0.3214 1 2.167 – – – – – – 1 0.2319
10 Dichapetalaceae – – 1 0.1788 – – – – – – – –
11 Ebenaceae 1 2.1405 2 1.2286 – – – – 1 1.4477 – –
12 Euphorbiaceae 3 8.8668 2 18.3106 4 6.4248 1 0.4884 3 40.6421 8 15.9521
13 Guttiferae 2 1.458 – – – – 2 2.7086 1 1.2117 1 2.8294
14 Loganiaceae 1 1.1275 – – 2 3.7271 1 4.0428 1 4.7435 – –
15 Meliaceae 2 1.2511 5 5.6527 1 0.159 3 1.7564 1 1.1586 5 1.6007
16 Mimosoidae 1 0.839 1 1.0062 3 7.6568 2 5.4996 2 8.7952 2 0.5702
17 Moraceae 3 1.5215 5 5.4242 5 5.2047 2 4.0903 1 1.6734 3 5.8794
18 Myristicaceae 1 3.8078 2 3.6885 1 3.2118 1 6.2918 – – 1 5.325
19 Ochnaceae 1 0.2234 1 0.1586 – – 1 19.5841 1 14.3522 1 0.267
20 Palmae – – – – 1 0.1494 – – – – – –
21 Papilionoidae 1 0.2061 4 1.3434 2 1.2517 1 21.7428 3 10.4944 – –
22 Rubiaceae 1 6.1651 4 4.6309 1 0.1907 2 3.9047 2 6.5865 2 1.8502
23 Rutaceae 1 2.6927 1 0.5764 – – 1 0.1329 – – 1 4.3173
2745

123
 Table 5 continued
2746

S/N Family Omotoso Legge Ode-Aye Igbo-Egunrin Atijere Igbotako

123
No. of RDO No. of RDO No. of RDO No. of RDO No. of RDO No. of RDO
Species Species Species Species Species Species

24 Sapindaceae 1 0.7204 1 0.2178 1 0.8004 – – – – – –

25 Sapotaceae 2 3.91 1 1.0448 – – – – 1 4.3954 – –
26 Simaroubaceae – – – – – – – – 1 2.2604 – –
27 Sterculiaceae 3 3.3331 7 12.0296 2 40.3437 – – – – 4 16.4071
28 Ulmaceae – – 1 3.7678 1 1.2463 1 1.1695 1 1.1586 3 2.6091
29 Verbenaceae 1 4.787 – – 1 0.1494 1 1.3181 – – – –
Biodivers Conserv (2008) 17:2735–2755
Biodivers Conserv (2008) 17:2735–2755 2747

Queen Forest an inviolate biosphere reserve in Ondo state. The closeness in species
evenness obtained for each of the selected communities’ forest, however revealed the
similarity in the pattern of tree species distribution in the area (Table 6).
A total of 29 families of trees encountered in this area shows that forests in the bitumen
belt are very rich in tree species, which are of immense value to rural livelihood. Isichei
(1995), Were (2001), Akinyemi et al. (2002), Onyekwelu et al. (2005) and Adekunle
(2006) are few among research workers that have reported on the dominance of members
of the Sterculiaceae, Moraceae, Ulmaceae, Meliaceae, Euphorbiaceae families in the
Nigerian tropical forests. The data presented here is in conformity with their unanimous
assertion; the dominant families in the study area are Euphorbiaceae, Moraceae, Meliaceae
and Sterculiaceae. The number of trees (tree density) of 457 trees/ha obtained in the study
area indicates that the area has a dense vegetation and also reflects the recovery status of
preciously disturbed parts by man as a result of his daily indispensable activities in the
forest. Legge (Oluwa Forest) and Atijere (Eba forest), which are forest reserves and
reservations under protection (gazette) greatly demonstrate higher values of tree density
609 and 541 trees/ha respectively. The lesser value in other communities’ forest can be
attributed to the significant disturbances done to the forest vegetation there, as consequent
to human activities, since they were not protected by any law (free areas) (Table 7).
The mean tree basal area of 26.69 m2/ha obtained in this study favourably compares
with the 25.5 m2/ha also obtained for tropical humid forest in Rio Xingu, Brazil (Par-
thasarathy 2001) and higher than 22.68 m2/ha recorded in a similar forest in Onigambari,
Nigeria (Akinyemi et al. 2002). This value is also higher than the recommended 25 m2/ha
for a fully stocked forest (Alder and Abayomi 1994). The variation in the tree basal area
among the selected communities’ forest is pronounced. Except for Omotoso free area
forest that has a value of 12.13 m2/ha and Ode-Aye forest with a value of 16.73, other
communities’ forest are higher than the 16.84 m2/ha reported by Adekunle et al. (2002) for
Omo forest reserve in the tropical rainforest ecosystem of southwest Nigeria.
Although, the mean tree volume obtained in this study could be seen as relatively high
(262.36 m3/ha), but value is relatively lower that for some other forest reserves in the same
ecosystem in Nigeria. This, in support of the diameter distribution result, gives the indi-
cation that most of the trees are not yet mature for sawn-timber harvest.
Nath et al. (2005) asserted that the inverted J-curve, where the abundance decreases
with increasing diameter, is an indication of good regeneration of the constituent species.
The conformity of the population structure of trees in all the selected communities’ forest
with this reverse J-shaped structure as shown in Fig. 2(a–f) clearly reflects the potential of
these communities’ forest to regenerate over a space of time. Larger population of the trees
falls in the lowest diameter size class; the amount of merchantable trees with dbh C48 cm
dbh is very meager. Non-availability of large dimensioned trees with dbh that is above
100 cm in Eba Island FR at Atijere and Igbo-Egunrin forest could be due to the fact that
these forests are very close to the river or as a result of previous human activities in the
area.
The presence of Funtumia elastica in all the forests, Albizia ferruginea, Alstonia boonei,
Baphia nitida, Celtis zenkeri, Cleistopholis patens, Lophira alata, Macaranga barteri,
Pycnanthus angolensis, Ricinodendron heudelotii and Sterculia tragacantha to more than
half of the selected communities’ forest reveal a moderate floristic overlap in their forest
vegetation. In contrast, each of Aningeria robusta, Anthostemma aubryanum, Cola
millenii, Gmelina arborea, Hannoa klaineana, Piptadeniastrum Africana, Trema orientalis
and Uapaca heudelotii is exclusively found in a community. The occurrence of Gmelina
arborea can be attributed to the dispersal of its seeds from the existing plantation in

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2748 Biodivers Conserv (2008) 17:2735–2755

Table 6 Tree species diversity, population density, Shannon index and species evenness in selected
communities’ forest in the bitumen–producing area of Ondo state
Families–species Omotoso Legge Ode-Aye Igbo-Egunrin Atijere Igbotako piLNpi

Agavaceae
Dracaena mannii – 12 – – – 7 -0.034
Anacardiaceae
Lannea welwitschii 7 9 – – – – -0.030
Mangifera indica – – 6 6 – – -0.024
Spondias mombin* – – 66 – – 3 –0.093
Annonaceae
Cleistopholis patens* 3 35 12 31 – 13 -0.116
Enantia chlorantha – – – – 7 – -0.015
Xylopia aethiopica – – – – – 4 -0.010
Apocynaceae
Alstonia boonei* 25 17 6 – – 13 -0.085
Alstonia congensis 6 – – – – – -0.013
Funtumia elastica* 34 28 22 56 13 66 -0.202
Holarrhena floribunda – 4 – – – – -0.010
Rauvolfia vomitoria 6 – 3 – – – -0.019
Bignonaceae
Newbouldia laevis – – 6 10 – – -0.030
Bombacaceae
Bombax buonopozense – – – – – 3 -0.007
Ceiba pentandra – 3 – – – – -0.007
Burseraceae
Canarium schweinfurthii – – 3 8 – – -0.022
Caesalpinoidae
Amphimas pterocarpoides 3 – 3 16 – – -0.039
Daniellia ogea 3 – – – – – -0.007
Dialium dinklagei – 7 – – – – -0.015
Dialium guineense* 31 – 9 – – – -0.062
Distemonanthus – 3 – – – – -0.007
benthamianus
Combretaceae
Terminalia superba 3 12 – – – 3 -0.033
Dichapetalaceae
Dichapetalum – 3 – – – – -0.007
madagascariense
Ebenaceae
Diospyros dendo – 9 – – – – -0.019
Diospyros mespiliformis* 6 17 – – 38 – -0.085
Euphorbiaceae
Alchornea cordifolia – – 3 – – – -0.007
Anthostemma aubryanum* – – – – – 38 -0.059
Drypetes spp – – – – – 12 -0.024
Macaranga barteri* 28 3 9 7 – 6 -0.076

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Biodivers Conserv (2008) 17:2735–2755 2749

Table 6 continued
Families–species Omotoso Legge Ode-Aye Igbo-Egunrin Atijere Igbotako piLNpi

Maesobotrya barteri – – – – – 3 -0.007

Phyllanthus discoideus 13 – 12 – 19 – -0.066
Ricinodendron heudelotii* 6 75 7 – – 3 -0.113
Uapaca guineensis – – – – 13 – -0.025
Uapaca heudelotii – – – – 56 – -0.080
Unknown 1 – – – – – 6 -0.013
Guttiferae
Allanblackia floribunda 9 – – – 19 – -0.047
Garcinia kola – – – 9 – – -0.019
Harungana madagascariensis 3 – – 3 – 19 -0.043
Loganiaceae
Anthocleista djalonensis 9 – 19 – – – -0.047
Anthocleista vogelli* – – 6 31 13 – -0.073
Meliaceae
Carapa procera 3 6 – 3 – 6 -0.033
Ekebergia senegalensis – – 3 – – – -0.007
Entandrophragma angolensis – 9 – – – – -0.019
Guarea cedrata – – – 3 – 6 -0.019
Guarea thompsonii – – – – 6 – -0.013
Khaya ivorensis – 3 – – – 4 -0.015
Trichilia monadelpha – 19 – 9 – 3 -0.051
Trichilia prieureana 12 19 – – – 3 -0.054
Unknown 2 – – – – – 3 -0.007
Mimosoidea
Albizia ferruginea* – 10 41 4 – 10 -0.089
Albizia zygia – – 3 – – – -0.007
Brachystegia eurycoma 9 – 7 – 6 3 -0.043
Pentaclethra macrophylla – – – 16 – – -0.030
Piptadeniastrum africana* – – – – 25 – -0.043
Moraceae
Antiaris africana – – – – – 12 -0.024
Bosqueia angolensis – 3 4 – 6 – -0.025
Ficus capensis – – 3 – – – -0.007
Ficus exasperata 4 – 6 – – 6 -0.030
Ficus mucuso – 3 6 3 – – -0.024
Ficus sur – – 13 – – – -0.025
Melicia excelsa 3 – – – – – -0.007
Mussanga cecropiodes 6 3 – 6 – 6 -0.037
Myrianthus arboreus – 9 – – – – -0.019
Treculia africana – 12 – – – – -0.024
Myristicaceae
Pycnanthus angolensis* 25 9 19 31 – 22 -0.126

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2750 Biodivers Conserv (2008) 17:2735–2755

Table 6 continued
Families–species Omotoso Legge Ode-Aye Igbo-Egunrin Atijere Igbotako piLNpi

Staudtia stipitata* – 25 6 – – – -0.051

Ochnaceae
Lophira alata* 3 7 – 6 94 3 -0.131
Palmae
Elaeis guineensis – – 3 – – – -0.007
Papilionoidae
Baphia nitida* 3 6 6 – 38 – -0.076
Lonchocarpus sericeus – 16 – – – – -0.030
Millettia thonningii* – – 7 141 44 – -0.186
Pterocarpus osun – 3 – – – – -0.007
Pterocarpus soyauxii – 3 – – 13 – -0.030
Rubiaceae
Canthium subcordatum* – 13 3 – 31 – -0.070
Mitragyna ciliata* – 25 – – – 3 -0.047
Nauclea diderrichii 19 17 – – – – -0.057
Pausinystalia talbotii* – – – 3 – 22 -0.043
Rothmannia hispida* – 3 – 9 25 – -0.058
Rutaceae
Zanthoxylum zanthoxyloides 10 9 – 3 – 19 -0.063
Sapindaceae
Blighia sapida 6 6 – – – – -0.024
Lecaniodiscus cupanioides – – 12 – – – -0.024
Sapotaceae
Aningeria robusta* 25 – – – – – -0.043
Chrysophyllum albidum – 3 – – – – -0.007
Malacantha alnifolia* 3 – – – 38 – -0.063
Simaroubaceae
Hannoa klaineana* – – – – 31 – -0.051
Sterculiaceae
Cola gigantea – 3 9 – – 6 -0.033
Cola millenii* – 35 – – – – -0.056
Cola nitida 3 – – – – – -0.007
Mansonia altissima – – – – – 19 -0.034
Pterygota macrocarpa – 6 – – – 9 -0.029
Sterculia oblonga – 4 – – – – -0.010
Sterculia rhinopetala 3 6 – – – 16 -0.043
Sterculia tragacantha* 7 9 6 6 – – -0.047
Triplochiton scleroxylon – 6 – – – – -0.013
Ulmaceae
Celtis integrifolia – – – – – 3 -0.007
Celtis zenkeri* – 66 9 3 6 9 -0.115
Trema orientalis* – – – – – 25 -0.043

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Biodivers Conserv (2008) 17:2735–2755 2751

Table 6 continued
Families–species Omotoso Legge Ode-Aye Igbo-Egunrin Atijere Igbotako piLNpi

Verbenaceae
Gmelina arborea* 22 – – – – – -0.039
Vitex doniana – – 3 – – – -0.007
Total 361 609 380 423 541 426 -4.02
* Most common species in one or more of the selected communities’ forest

Table 7 A comparison of recent forest inventories in the tropical rainforest ecosystem

Locality Source Number Basal Number of Number of H1
of trees/ha Area (m2)/ha families/ha species/ha

Nigeria
Omotoso This study 361 12.13 21 35 3.21
Oluwa FR This study 609 54.36 22 48 3.42
Ode-Aye This study 380 16.73 19 36 3.16
Igbo-Egunrin This study 423 19.19 19 25 2.46
Eba Island FR This study 541 28.66 15 21 2.76
Igbo-Egunrin This study 426 28.3 17 39 3.26
Ala FR Adekunle (2006) 148 n.a 24 54 3.62
Omo FR Adekunle (2006) 115 n.a 21 41 3.34
Shasha FR Adekunle (2006) 159 n.a 20 55 3.66
Queen FR* Onyekwelu et al. 742 85.4 26 51 3.31
(2005)
Onigambari FR Akinyemi et al. 308 22.68 22 50 n.a.
(2002)
Selected other localities
Sengaltheri* (India) Parthasarathy 965 55.34 n.a 82 3.69
(2006)
Cumbre Pilón (Bolivia) Smith and Killeen 647 30.62 37 146 n.a
(1995)
Rio Colorado (Bolivia) Smith and Killeen 588 26.4 31 78 n.a.
(1995)

* Undisturbed natural forest; n.a. not available

Omotoso. Likewise, the presence of Elaeis guineensis in Ode-Aye forest is typical of

degraded natural forest (Fig. 3).
The demonstration of high level of tree species diversity in this area gives the impetus
for the placement of an economic value on the forest vegetation and many biological
resources there. Failure to recognize this would be tantamount to failure of these resources
to compete on a level playing field with the forces driving their decline (OECD 2002). The
need for paying cognizance to the practical valuation of these communities’ forest vege-
tation and their constituent biodiversity therefore becomes conspicuous. The inevitability
of this deeply seated upon the premises that the clearance of forest vegetation to pave way

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2752 Biodivers Conserv (2008) 17:2735–2755

500m

20m

20m
1000m

250m

Transect 1 Transect 2

Fig. 2 Plots location using systematic line transect

for bitumen exploration will result in the further loss of habitat, biodiversity and envi-
ronmental values in the area. It must however be pointed out that any prospective bitumen
exploratory industry who wishes to clear forest vegetation must establish a process to
protect and manage the biodiversity in the area, as any habitat destroyed is very difficult to
regenerate. Also, once the vegetation is cleared, some important species may go into
extinction and the value of those species to mankind may be lost forever.
Mining industries have often failed to make contribution to sustainable development
and to protect the environment adequately (Dudley and Stolton 2002). This necessitates the
regulation of their movement into these remote fragile ecosystems and the area of unique
biodiversity as a means of protecting the environment and the people who live there. The
Ondo state government in her recent proposals for industrialization has earmarked this
study area for the location of bitumen exploratory industries owing to its inherent bitumen
deposition; it now becomes imperative to adhere to a number of innovative ideas presented
in the 2002 report of WWF by Dudley and Stolton (2002). This requests the prospective
industries as a first step to respect the IUCN Amman 2000 resolution, which calls on the
industries to stay out of the protected areas and also must help to conserve critical area of
high biodiversity wherever they are sited for exploration.

Conclusion and recommendations

This study investigated the diversity and biovolume of tree species (C10 cm dbh) in some
communities in the bitumen producing area of Ondo state to provide baseline information
prior to commencement of bitumen exploration. Results showed that the forest vegetation
in this area has been under intense human disturbances, but currently displays a sign of
rejuvenation. Evidently, the area is very rich in species that are relied solely upon by the
rural communities close to them. The various communities selected are dominated by

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Biodivers Conserv (2008) 17:2735–2755 2753

5 (a) Omotoso Forest 350 16 (b) Oluwa FR 300

300
Basal area (m )

14
2

4 250

Basal area (m )
2
No. of trees
250 12

No. of trees
3 10 200
200
8 150
2 150
6 100
100 4
1 50
50 2
0 0 0 0

90 0
00

00
0

90 0
00

00

--2

--3

--4

--5

--6

--7

--8

--9
--2

--3

--4

--5

--6

--7

--8

--9

--1

>1
--1

>1

10

20

30

40

50

60

70

80
10

20

30

40

50

60

70

80 Diameter size class (cm) Diameter size class (cm)

6 (d) Igbo-Egunrin Forest 300

5 (c) Ode-Aye Forest 300 5 250

Basal area (m )
2

No. of trees
Basal area (m )
2

4 250 4 200

No. of trees
200 3 150
3
150
2 2 100
100
1 50 1 50
0 0 0 0

0
00

00
0

90 0
00

00

--2

--3

--4

--5

--6

--7

--8

--9
--2

--3

--4

--5

--6

--7

--8

--9

--1

>1
--1

>1

10

20

30

40

50

60

70

80
10

20

30

40

50

60

70

80

90
Diameter size class (cm) Diameter size class (cm)

7 (f) Igbotako Forest 300

9
8
(e) Eba Island FR 300 6 250
Basal area (m )
2

250
Basal area (m )

No. of trees
2

7 200
No. of trees

6 200 4
5 150
150 3
4
3 100 2 100
2 1 50
50
1
0 0 0 0
0

0
00

00
0

0
00

00

--2

--3

--4

--5

--6

--7

--8

--9
--2

--3

--4

--5

--6

--7

--8

--9

--1

>1
--1

>1

10

20

30

40

50

60

70

80
10

20

30

40

50

60

70

80

90
90

Diameter size class (cm) Diameter size class (cm)

Basal Area Adundance

Fig. 3 (a–f) Population structure of tree species in selected communities’ forest in the bitumen-producing
area of Ondo state based on diameter-frequency and basal area

different combination of species. Most of the tree species are distributed in the lowest
diameter size class and the percentage of the merchantable trees ready for sawn-timber
harvest is very meager. The area, besides being distinct in its biodiversity constituents, also
harbours a vast number of genetic resources. Allocation of the area for bitumen exploration
would therefore demands for good management of the area for the conservation and
sustainability of the constituent resources.
The variation in the species composition and abundance among the selected commu-
nities’ forest necessitates the preservation of the vast acreage of the forest in this area, as a
single large reserve. The forest should also be managed to provide other services like
tourism, wildlife sanctuary, watershed management and climatic amelioration. Meanwhile,
the authority in control of these forests should eschew from subsequent conversion of
natural forest to industrial plantation of exotic and indigenous species as the case of Oluwa
forest reserve- one of the richest ecosystem of Nigeria whose greater part had been clear-
felled for plantation species, which defy biodiversity conservation in this area.
In view of the proposed allocation of this area to bitumen exploratory industries, it is
suggested that Legge (Oluwa forest) and Atijere (Eba forest) which are zones of high
protection and high conservation value should be tagged ‘no-go area’ for bitumen
exploration while exploration can only commence in Igbotako with intense biodiversity

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2754 Biodivers Conserv (2008) 17:2735–2755

conservation in view. Also, the industries can proceed into Omotoso, Ode-Aye and Igbo-
Egunrin under good forest management strategies. The prospective industries should also
be made to agree for compensation measures, including habitat creation to maintain
constant natural assets. Therefore, the decision to allow these bitumen exploratory
industries should be based on their readiness to sign a contract for good management of the
biological resources, with little deleterious effect and low environmental risk, should they
be allowed. Another study should be carried out a few years after the commencement of
the exploration such that the results in term of tree species diversity and yield could be
compared with the result of this baseline study. This will help allow for the effect of
bitumen exploration on the habitat to be assessed.

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