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Tropical Ecology 54(3): 275-289, 2013 ISSN 0564-3295

© International Society for Tropical Ecology


www.tropecol.com

Tree species diversity and structure of a Nigerian strict nature reserve


*
VCTOR AJIBOLA JIMOH ADEKUNLE , ADEWOLE OLASIYAN OLAGOKE &
SHADRACH OLUFEMI AKINDELE

Federal University of Technology, Department of Forestry and Wood Technology, Akure, Nigeria

Abstract: This study was carried out to examine the efficacy of Strict Nature Reserves
(SNR) as a means of biodiversity conservation. This was achieved by assessing the status of the
forest in terms of tree species diversity, abundance and growth yield. Data were collected from
four plots of one hectare size each, randomly located in the forest. In each plot, all tree species
(dbh ≥ 10 cm) were identified and their height, diameter at breast height (dbh) and diameters at
the top, middle and base were measured. There were 387 stems ha-1, belonging to 94 tropical
hardwood species, 80 genera and 30 families in the SNR. The most abundant species and family
were Celtis zenkeri of Ulmaceae family (41 stems ha-1) and Sterculiaceae (10 species),
respectively. The Shannon-Wiener index (3.75) and evenness (0.82) and other diversity indices
were very high, indicating that the forest is a potential biodiversity hotspot. The indices
compared favourably with several protected areas that are biodiversity hotspots. In addition,
the total basal area and volume of 674.33 m2 and 4,646.93 m3 and the vertical and horizontal
structure confirm that the SNR is a mature natural forest. This study provides a baseline for
the management of protected areas in developing countries and it shows the potential of in situ
method in nature conservation.

Resumen: Se llevó a cabo este estudio para investigar la eficacia de las Reservas Naturales
Estrictas (SNR, siglas en inglés) como medio de conservación de la biodiversidad. Se evaluó el
estado del bosque en términos de la diversidad, la abundancia y el rendimiento en el crecimiento
las especies arbóreas. Se obtuvieron datos en cuatro parcelas, cada una de una hectárea,
localizadas al azar en el bosque. En cada parcela todas las especies arbóreas (DAP ≥ 10 cm)
fueron identificadas y se midieron sus alturas, diámetros a la altura del pecho (DAP) y
diámetros en la parte superior, intermedia y basal. En la SNR se registraron 387 tallos ha-1
pertenecientes a 94 especies tropicales de maderas duras, 80 géneros and 30 familias. La
especie y la familia más abundantes fueron Celtis zenkeri de la familia Ulmaceae (41 tallos ha-1)
y Sterculiaceae (10 especies), respectivamente. El índice de Shannon-Wiener (3.75), la
equitabilidad (0.82) y otros índices de diversidad fueron muy altos, lo que indica que este bosque
es potencialmente un hotspot de biodiversidad. Los índices se compararon favorablemente con
los de varias áreas protegidas que son hotspots de biodiversidad. Además, el área basal total
(674.33 m2) y el volumen total (4,646.93 m3), así como las estructuras vertical y horizontal,
confirman que la SNR es un bosque natural maduro. Este estudio proporciona una base para la
gestión de áreas protegidas en los países en desarrollo y muestra el potencial del método in situ
para la conservación de la naturaleza.

Resumo: Este estudo foi realizado para examinar a eficácia das reservas naturais estritas
(SNR) como um meio de conservação da biodiversidade. Isto foi conseguido avaliando o estado
da floresta em termos de abundância de espécies arbóreas, diversidade e crescimento. Os dados
foram coletados a partir de quatro parcelas de um hectare cada, aleatoriamente localizadas na
floresta. Em cada parcela, todas as espécies de árvores (DAP ≥ 10 cm) foram identificadas e sua
altura, diâmetro à altura do peito (DAP) e diâmetros no topo, meio e base foram medidos.

*Corresponding Author; e-mail: adekunlevaj@rediffmail.com


276 TREE SPECIES DIVERSITY AND STRUCTURE

Contabilizaram-se 387 caules ha-1, pertencentes a 94 espécies lenhosas tropicais, 80 géneros e


30 famílias na SNR. As espécies mais abundantes e famílias foram a Celtis zenkeri da família
Ulmaceae (41 espécimes ha-1) e Sterculiaceae (10 espécies), respectivamente. O índice de
Shannon-Wiener (3,75) e similaridade (0,82) bem como outros índices de diversidade eram
muito elevados, o que indica que a floresta é um ponto quente de biodiversidade potencial. Os
índices comparam-se favoravelmente com outras áreas protegidas diversas que são também
ponto quente de biodiversidade. Além disso, a área basal total, de 674,33 m2 e o volume 4.646,93
m3 bem como a estrutura vertical e horizontal confirmam que a SNR é uma floresta natural
madura. Este estudo fornece uma base para a gestão de áreas protegidas nos países em
desenvolvimento e evidencia o potencial do método in situ na conservação da natureza.

Key words: Biodiversity indices, forest structure hotspots, in-situ conservation, stand
volume.

Introduction only on paper. They have either been converted to


farmland of arable and cash crops or other land
Strict Nature Reserve (SNR) is prominent uses, such as large scale afforestation pro-
among the methods for in situ conservation of grammes. In addition, the lucrative nature of
biodiversity in Nigeria in particular and the world timber trade attracted many Nigerians into the
in general. Other common methods in Nigeria business, causing continuous timber harvesting in
include Biosphere Reserves, Game Reserves, both the constituted forest and the free areas.
Regeneration Plots, Permanent Sample Plots, and According to Adekunle et al. (2010), a total of
Sacred Groves. SNRs are created to protect 111,377 timber stems, belonging to 62 different
representative samples of natural ecosystems for indigenous hardwood species of tropical rainforest
preservation of biodiversity and ecological ecosystem, distributed among 16 families, were
processes, scientific study, environmental moni- exploited from Ondo State forest ecosystem
toring, education and the maintenance of genetic between 2003 and 2005.
resources in a dynamic and evolutionary state Other factors responsible for the drastic
(Isichei 1995). There are six SNRs in Nigeria, one reduction of the rich floristic resources of Nigeria
biosphere reserve, 160 constituted forest reserves are illegal activities in the forest, declining
and six National Wildlife Parks. Constituted forest manpower and capacity in Forestry Department,
reserves covered a total land area of about 1,075 inadequate forest patrol, stoppage of the payment
km2. This is 10 % of the 25 % of forestland of annual royalty (formerly 5 % of total income)
earmarked for conservation during the reservation from what accrued from logging activities to rural
phase in the history of forestry development in communities, outdated forestry laws and regu-
Nigeria. These forest reserves are scattered in the lations. Similarly, Maliyat & Datt (2010) reported
36 States of the Federation, and they are locally that the expansion of biotic activities is also
controlled by their respective State Governments responsible for overexploitation of natural resour-
through the Department of Forestry and Wildlife, ces, and this has subsequently disturbed the
but all the SNRs and the biosphere reserve were delicate equilibrium that exists between living
established and managed by the Forestry Research organisms and their environment. So, both the
Institute of Nigeria. All other woodlands, apart reserve where logging is allowed with government
from the constituted areas are regarded as free regulations and the free areas that are not under
areas. strict regulations have suffered devastation in
Lafrankie et al. (2006) noted that the tropical Nigeria. It appears, therefore, that intact old-
rainforests are vulnerable to deforestation and growth forests, where biodiversity is conserved for
degradation. In Nigeria, population growth has led posterity are restricted only to the SNRs,
to an astronomical increase in anthropogenic biosphere reserve and the sacred groves and
activities, excessive logging and over exploitation. community forests. These now serve as refuges for
As a result, most of these forest reserves now exist animals driven by human disturbances from other
ADEKUNLE, OLAGOKE & AKINDELE 277

Table 1. List of SNRs in Nigeria, their sizes and format, of the present status of Nigerian’s nature
locations. reserves for international audience. This is not the
case with the countries listed as mega-diverse
Name Area State Ecological countries. As a result, recognising the worth of
(ha) zone Nigerian protected areas as potential biodiversity
Akure 32 Ondo Rainforest hotspots is in jeopardy and it could hinder the
Bam Ngelzarma 142 Yobe Savannah country from benefiting from the REDD projects.
Bonu 145 Bauchi Savannah
Börner & Wurner (2008) and Dewi et al. (2009)
Lekki 78 Lagos Mangrove
Milliken Hill 49 Enugu Rainforest emphasized the need for credible temporal base-
Omo (Biosphere 460 Ogun Rainforest lines in developing projections for forest-cover
Reserve) change. This is fundamental to REDD project
Ribako 170 Kaduna Savannah development (CCBA 2008) and for biodiversity
Urhonigbe 64 Edo Rainforest hotspots identification.
Myers (2003) reported that a great number of
forests and free areas, and home for so many endemic species are found in relatively few areas.
endangered indigenous plant species. He noted further that certain of the 25 already
In situ conservation methods are veritable identified terrestrial hotspots have been assessed
tools for the preservation of genetic resources as the hottest hotspots and they are still receiving
currently decreasing at an alarming rate. To more conservation efforts. But the hotspot strategy
promote nature conservation worldwide, Myers et should be made to include other areas and lo-
al. (2000) brought about the concept of hotspot cations for conservation efforts because most
that considered the regions with exceptional con- species-rich habitats are usually prone to
centration and assemblage of endemic species that overexploitation and abuse. In view of the above,
are experiencing a high rate of habitat loss. They this study was carried out in a SNR within Akure
noted that conservation and protection efforts Forest Reserve to document its present tree
should be prioritised in these areas. Twenty five diversity and structure. The results were com-
such biodiversity hotspots were identified world- pared with other forest ecosystems to assess its
wide (Myers et al. 2000). At present, there is no worth as a potential biodiversity hotspot.
recognised biodiversity hotspot in Nigeria. The
country is not also among the 16 mega-diverse
countries of the world. This does not preclude that
there are potential biodiversity hotspots in the
country. The country is very rich in forest
resources, with eight important tropical ecological
zones, broadly categorised into three (the tropical
rainforest ecosystem, the mangrove in the
southern part and the savannah in the northern
part). The biosphere reserve and the SNRs in the
country could be potential biodiversity hotspots,
which could qualify the country to be among the
mega-diverse countries of the world if adequate
biodiversity assessments are carried out and
documented. Fig. 1. Map of Nigeria showing Ondo State.
To benefit from the initiative on Reducing
Deforestation and Degradation of Tropical Forests Materials and methods
(REDD), some countries have prepared proposals
that include nature reservation, protection and Study area
monitoring. Some of the nature reserves are
Rimba Raya Biodiversity Reserve in Indonesia, This study was carried out in SNR 1 located
Maashan Nature Reserve in China, Maya Bio- within Akure Forest Reserve in Ondo State,
sphere Reserve in Guatemala and the Juma Southwest, Nigeria (Fig. 1). It lies within 4º 30’ and
Reserve in Brazil (CCBA 2011). There is scarcity of 6º East and 5º 45’ and 8º 15’ North. The climate of
baseline data, ecological information and docu- the area is humid sub-tropical, indicating that it is
mented evidence, either in printed or electronic basically within the tropical rainforest ecological
278 TREE SPECIES DIVERSITY AND STRUCTURE

zone. It is dominated by broadleaved hardwood following Kent & Coker (1992), Magurran (2004)
trees that form dense, layered stands. The mean and Lu et al. (2010):
annual temperature is about 26 ºC (minimum 19 (i) Shannon-Wiener diversity index:
ºC and maximum 34 ºC) and the rainy season lasts s
for 9 months annually, between March and
November (about 2500 mm with bimodal rainfall
H' = −
∑i =1
p i ln( p i )

pattern) while the dry seasons usually last for 3


months, between December and February. where, H' is the Shannon-Wiener diversity
This portion of the reserve (Akure Forest index; S is the total number of species in the
Reserve) was carved out and designated as a Strict community; pi is the proportion of S made up of the
Nature Reserve (SNR) in 1954 by the Forestry ith species; Ln is natural logarithm.
Research Institute of Nigeria (FRIN). Plantations (ii) Pielou’s species evenness index:
of indigenous and exotic tree species were s

established at the boundaries and buffer zone of H' ∑ p i ln( p i )


the reserve. The SNR is bounded on one side by a E H = = i = 1
H max ln(S)
river. This made accessibility to the forest very
difficult. There are several rural communities
around the reserve. They are involved in its mana- (iii) Margale f’s index of species richness (M):
gement and protection. All their anthropogenic (S–1)
activities, namely, organized taungya farming, M=
LnN
collection of falling twigs and branches as firewood
and gathering of other non-timber forest products (iv) Simpson concentration index:
(NTFPs), were restricted to the buffer zone. These ni
activities were carried out with adequate moni- λ = ∑ ( )^2
Ni
toring and effective patrol by forest guards. The (v) Number 1 of Hill diversity index (N1) = eH
names, sizes and locations of all SNRs in Nigeria 1
is presented in Table 1. (vi) Number 2 of Hill diversity index (N2) = λ

Data collection Forest structure analyses


The entire forest was divided into rectangular Basal area
plots of 100 x 100 m (1 ha) in size and four of the The basal area of all trees in the sample plots
1-ha plots were selected using the simple random were calculated using the formula:
sampling technique. Within each of the selected BA = (л D2)/4
four sample plots, measurement and identification where, BA = Basal area (m2), D = Diameter at
were limited to all woody plants with diameter at breast height (cm) and л = pie (3.142). The total
breast height (dbh) of ≥ 10 cm. With this minimum BA for each plot was obtained by adding all trees
diameter at breast height, most of the classes of BA in the plot.
woody plants were captured. The tree data Volume
collected in each sample plot for further analysis The volume of each tree was calculated in
were dbh, diameters over bark at the base, middle every plot using the Newton’s formula (Hush et al.
and merchantable top of all trees in the plots, and 2003):
height of all trees in each plot using Spiegel V = (h/6) (Ab + 4 Am + At).
relaskop. where, V = Tree volume (m3), Ab, Am and At =
The number and scientific names of all the tree tree cross-sectional area at the base, middle and
species encountered in each field plot were recor- top of merchantable height, respectively (m2) and h
ded. When it was difficult to identify the species in tree height (m). Plot volumes were also obtained
the field, the common/local name were recorded, by adding the volumes of all the trees in the plot.
and plant specimens were collected for iden- The basal area and volume ha-1 were obtained
tification at the Forestry Research Institute of by dividing the sum of basal area and volume,
Nigeria, Ibadan, Nigeria herbarium. respectively, for the four plots (one ha plot) by 4.
Values for the entire stand were obtained by
Data analyses
multiplying the value of one ha with the forest size
Tree species diversity (32 ha).
The following indices were also employed (i) Relative density (%) of each species was
ADEKUNLE, OLAGOKE & AKINDELE 279

computed following the equation of Brashears addition, about 22 % of the species had densities of
et al. (2004): less than 5 individuals ha-1 (Fig. 2). Sterculiaceae
⎛ ni ⎞ had the highest number of species (10 species),
RD = ⎜⎜ ⎟⎟ × 100
⎝N⎠ followed by Meliaceae and Moraceae with eight
species each (Table 3). Mimosoideae had seven
where, RD is the relative density of the species while Apocynaceae and Rubiaceae had five
species; ni is the number of individuals of species i species each. About 36 % of the families were
and N is the total number of all individual trees. represented by only one species in the forest and
(ii) Relative dominance (%) of each species was Sterculiaceae had the highest number of
estimated using the following equation: individuals (127 stems ha-1, about 33 % of the total
((Σ 100 )
Bai ×× 100)
∑ Ba
stand density). Very high Shannon-Weiner index
i
RDo = (3.74) and equitability index, using Pielou’s
∑ Ba n
Σ Ban evenness index of 0.82 were obtained (Table 4).
The results of the other biodiversity indices were
where, RDo is the relative dominance of the
64.72 for Margalef’s index of species richness, 0.07
species; Bai is the basal area of all individual trees
for Simpson concentration index, 42.09 for number
belonging to a particular species i; Ban is the basal
1 of Hill’s diversity index and 14.29 for number 2
area of the stand.
of Hill’s diversity index.
(iii) Importance Value Index (IVI): The sum of the
RD and RDo divided by 2 (RD x RDo/2) gave
the importance value index for each species
(Brashears et al. 2004; Yang et al. 2008). This
was used to express the share of each species
in the tree community (Rajkumar & Partha-
sarathy 2008).
(iv) Family Importance Value (FIV): This was used
to estimate a family’s share in the forest
community. It is defined as the sum of its
relative dominance (RDo) and density (RD) E. angolense

divided by 2.
Based on the forest structural analysis of
Proctor et al. (1983) & Newbery (1991), the size
class distributions were classified under four
distinct categories, namely, smaller (10 - 20 cm Fig. 2. Density curve of 15 species with > 5 indivi-
dbh); medium (21 - 50 cm dbh); large (51- 100 cm duals ha-1 in the SNR.
dbh) and largest (> 100 cm dbh). The stems were
further classified into 10 diameter and six height Forest structure
classes to show the graphical pattern of tree
Celtis zenkeri Engl. (Ulmaceae) had the
population distribution and vertical stratification
highest number of occurrence (41 stems ha-1) and a
respectively.
relative density of 10.9 (Table 2). So, it could be
To examine the relationship among the growth
regarded as the most abundant species in the
variables, Pearson Correlation Coefficient was
forest. This was closely followed by Mansonia
adopted.
altissima A. Chev. (Sterculiaceae) with 36 stems ha-1,
and a relative density of 9.30. The third abundant
Results
species was Pterygota macrocarpa K Schum. with
33 stems ha-1 and a relative density of 6.97.
Tree species diversity and abundance
Cleistopholis patens (Benth.) Engl. & Diels
The total number of trees (≥ 10 cm dbh) in the (Annonaceae) had the highest mean dbh (99.20
forest was 387 stems ha-1. These were dispro- cm) while the least dbh (10.20 cm) was recorded
portionately distributed among 94 species, 80 for Enantia chlorantha Oliv. in the same family.
genera and 30 families (Table 2). About 47 % of the The highest (25 m) and lowest (8.64 m) mean
species were represented by only a single indivi- heights were recorded for Khaya grandifoliola
dual ha-1 and 78 % with less than 5 stems. In C.DC and Sterculia tragacantha Lindl., respec-
280 TREE SPECIES DIVERSITY AND STRUCTURE
ADEKUNLE, OLAGOKE & AKINDELE 281
282 TREE SPECIES DIVERSITY AND STRUCTURE
ADEKUNLE, OLAGOKE & AKINDELE 283

Table 3. Families important index value for the study area.

Families BA ha-1 Vol ha-1 RD RDo FIV


Agavaceae 0.03 0.14 0.71 0.15 0.43
Anacardiaceae 0.19 1.63 0.19 0.91 0.55
Annonaceae 0.15 0.90 1.81 0.71 1.26
Apocynaceae 1.06 5.90 8.14 5.01 6.57
Aquifoliaceae 0.04 0.22 0.32 0.17 0.25
Bombacaceae 0.26 1.55 0.78 1.23 1.00
Boraginaceae 1.41 10.30 1.94 6.68 4.31
Caesalpiniodeae 0.47 2.80 2.84 2.21 2.53
Capparaceae 0.26 1.58 0.26 1.24 0.75
Combretaceae 0.18 1.01 0.52 0.85 0.68
Ebenaceae 0.13 0.75 0.78 0.63 0.70
Euphorbiaceae 0.89 4.74 1.68 4.20 2.94
Guttiferae 0.03 0.20 0.26 0.12 0.19
Irvingiaceae 1.56 10.59 5.62 7.38 6.50
Loganiaceae 0.20 1.40 0.58 0.94 0.76
Meliaceae 1.80 11.46 6.20 8.56 7.38
Mimosoideae 1.31 9.32 3.62 6.24 4.93
Moraceae 0.74 4.67 4.33 3.49 3.91
Myristicaceae 0.47 3.25 2.06 2.21 2.14
Ochinaceae 0.05 0.16 0.26 0.22 0.24
Olacaceae 0.03 0.22 0.52 0.16 0.34
Pandaceae 0.06 0.29 0.84 0.27 0.55
Papilionioideae 0.45 2.90 2.84 2.15 2.50
Rubiaceae 0.51 2.89 1.81 2.42 2.11
Rutaceae 0.02 0.13 0.45 0.11 0.28
Sapindaceae 0.28 1.77 2.33 1.35 1.84
Sapotaceae 0.86 5.33 3.88 4.09 3.98
Simaroubaceae 0.14 0.93 1.03 0.68 0.86
Sterculiaceae 6.92 48.82 32.69 32.84 32.76
Ulmaceae 2.06 9.34 10.79 9.78 10.28
Total 22.54 145.22

tively. The highest basal area and volume per Ulmaceae with a basal area and volume of 2.06 m2
hectare (2.10 m2 and 15.68 m3, respectively) were and 9.34 m3, respectively. The families with the
contributed by Mansonia altissima. This was highest mean dhb (52.6 cm) and height (24.8 m)
followed by Triplochiton scleroxylon K Schum. with were Anacardiaceae and Olacaceae, respectively.
basal area and volume of 1.55 m2 and 11.95 m3, Sterculiaceae also had the highest of 32.76 (i.e.
respectively. The least basal area and volume (0.01 31.65 % of the total FIV), followed by Ulmaceae
m2 and 0.06 m3, respectively) were recorded for with FIV of 10.28 (about 9.93 % of the total FIV).
Ceiba pentandra (L.) Gaertn. However, Mansonia Guttiferaceae had the lowest FIV of 0.19 (0.18 % of
altissima had the highest species importance with total FIV). The mean dbh and height for the forest
an IVI of 9.11 %. This was closely followed by were 22.68 ± 0.36 cm and 13.55 ± 0.14, respectively
Celtis zenkeri with IVI of 8.23. This species (Table 4). The total number of trees in the entire
contributed about 8.24 % to the total IVI in the forest was 12,384 with a total basal area and
forest. volume of 674.33 m2 and 4,646.93 m3, respectively.
Sterculiaceae had the highest total basal area The dominant dbh (Triplochiton scleroxylon) and
(6.92 m2) and volume (48.82 m3), followed by height (Celtis zenkeri) were 131.5 cm and 29.5 m,
284 TREE SPECIES DIVERSITY AND STRUCTURE

Table 4. Arboreal Biodiversity indices and growth


variables of the SNR.

Biodiversity indices Tree growth variables


Indices Values Variables Values
No. of Individual ha-1 387 Mean Dbh (cm) 22.68 ± 0.36
(Ni)
No. of Species (NS) 95 Dominant Dbh 131.5
(cm)
No. of Families (NF) 31 Mean height 13.55 ± 0.14
(m)
No. of Genera (NG) 80 Dominant 29.5
height (m)
Shannon-Weiner 3.74 Total Basal 22.54
Index (H’) Area/ha (m2) Fig. 3. Diameter class distribution frequency of tree
Pielou’s Evenness 0.82 Total Volume/ha 145.22 species in the SNR.
Index (E) (m3)
Margalef’s Index of 64.72
Species Richness
(M)
Simpson 0.07
Concentration Index
(λ)
Number 1 of Hill 42.09
Diversity Indices
(N1)
Number 2 of Hill 14.29
Diversity Indices
(N2)

respectively. Evidently, there were very wide


Fig. 4. Number of individuals ha-1 in each of the
ranges between the maximum and minimum dbh
height classes (Figures on bars indicate the number
(121.5 cm and height 27.7 m).
of plant families].
As typical of mature natural forest, the
number of stems was inversely proportional to
lies was recorded in the height class of 11-15 m,
diameter sizes, which was also evidenced from the
followed by the height class of 6 - 10 m (111 stems
reverse J pattern of the dbh class frequency
ha-1) of 70 species and 28 families (Fig. 4). The
distribution (Fig. 3), indication of a healthy
lowest proportion of stems (17 stem ha-1 in 23
recruitment of the individuals in the SNR. As the
species and 14 families) was recorded in the height
diameter increased, there was a decrease in the
class of ≥ 25 m. The majority of the trees (64 %)
number of stems (Table 5). The lowest diameter
class (10 - 20 cm) had the highest number of stems, occupied the middle stratum where they form a
species and family. This class had 222 stems ha-1, close canopy (height between 11 - 20 m). This was
about 57 % of the total stems ha-1 and 29 families. followed by 32 % at the lower stratum (height ≤ 10
This class also contributed the highest basal area m) and about 4 % occupied the upper stratum (26 -
ha-1 of 3.87 m2 followed by the second lowest dbh 30 m). Trees that could be referred to as emergent
class (21 - 30 cm) with 81 stems ha-1, three species (height up to 40 m) were not encountered in the
in one family and the highest volume of 29.62 m3. sample plots (Fig. 4).
Only one individual was encountered in the
diameter class of 91 - 100 cm and only two Relationship among variables
individuals each were recorded for dbh classes 71 - There was generally a positive and significant
80, 81 - 90 and > 100 cm (Table 5). linear relationship among the tree growth
For height distribution, the highest proportion variables (Table 6). The r-values ranged between
of stems (135 stems ha-1) of 60 species and 28 fami- 0.34 and 0.93. The highest correlation coefficient was
ADEKUNLE, OLAGOKE & AKINDELE 285

Table 5. Status of tree species in the SNR according to diameter (cm) and height classes.

DBH or Height NS NF Ni BA ha-1 (m2) Vol ha-1 (m3)


Dbh class (cm) 10 - 20 85 30 222 4.61 21.98
21 - 30 3 1 81 3.61 29.62
31 - 40 5 2 37 4.52 24.44
41 - 50 1 1 19 2.89 19.45
51 - 60 1 1 12 1.97 14.33
61 -70 1 1 9 1.78 12.61
71- 80 1 1 2 0.73 4.99
81 - 90 1 1 2 0.79 5.93
91 - 100 1 1 1 0.36 2.52
> 100 2 1 2 1.29 9.36
Total 387 22.54 145.22
Height class (m) <5m 22 12 11 0.48 3.2
6 - 10 70 28 111 6.57 46.59
11 - 15 60 28 135 8.07 46.18
16 - 20 53 27 87 3.86 25.47
21 - 25 27 18 26 2.18 14.35
26 - 30 23 14 17 1.39 9.44
Total 387 22.54 145.22

NS = number of species; NF = number of families; Ni = number of individuals ha-1; BA = Basal area; and
Vol = Volume.

obtained between the logarithm transformed basal The floristic diversity of this forest is very close
area and volume (0.93). Very weak correlation was to other tropical rainforests in some Asian countries
observed between tree height and most of the with biodiversity hot spots. Lu et al. (2010)
other growth variables. The r-value between the obtained a total of 428 stems ha-1 in 95 species and
height and basal area and also between height 38 families in tropical rainforest of Xishuangbana,
with volume was 0.34. China while Rajkumar & Parthasarathy (2008) got
a total of 105 species that belong to 32 families in
Discussion the evergreen forest of Andaman Giant, India. An
Forests contain the greatest diversity in terms average stand density of 422 stems ha-1 was
of species, genetic material and ecological reported for Borneo rainforest by Small et al.
processes of all ecosystems. Forest habitats play a (2004) and as high as 544 for a primary forest in
central role in the functioning of the biosphere, as Indonesia by Kessler et al. (2005). In a Mexican
they are the origin of many cultivated plants and tropical deciduous forest, 347 stems ha-1 in 148
animals (EU 2008). SNRs are one of the in situ species distributed among 42 families were
methods that are needed to strictly protect rare reported (Duran et al. 2006). Even though the
habitat types and habitats that are being stand density of the SNR is lower than those
disturbed with anthropogenic activities. The reported from other tropical forests, it fell within
results of this study revealed that this SNR is a the range of 245 and 467 stems ha-1 recorded for
repository of many indigenous tropical hardwood tropical forests by Campbell et al. (1992). The
tree species in different families. This is evidenced number of species is also within the range of 62 -
by the 387 stems ha-1 (dbh ≥ 10 cm) that belonged 247 species reported for a mature tropical forest in
to 94 indigenous hardwood species, distributed in southeast Asia (Losose & Leigh 2004), higher than
80 genera and 30 important families in this forest. 92 species of submontane tropical rainforest in
The number of tree species encountered in a Philippines (Hamann et al. 1999) and the 81
sample survey was adopted as a surrogate for the species of the mature lowland dense forest in
actual species richness in this study (Magnussen et Vietnam (Blanc et al. 2000). Tree density can be
al. 2010).
286 TREE SPECIES DIVERSITY AND STRUCTURE

Table 6. Correlation matrix for tree growth variables in the study area.

Dbh/Dab(cm) Height (m) BA (m2) Vol (m3) Ln Ba Ln Vol


Dbh/Dab (cm) 1.00
Height (m) 0.45 1.00
BA (m2) 0.93 0.34 1.00
Vol (m3) 0.62 0.34 0.67 1.00
Ln Ba 0.93 0.49 0.77 0.51 1.00
Ln Vol 0.88 0.63 0.71 0.59 0.93 1.00
Dbh- Diameter at breast height, Dab – Diameter above buttress.

affected by natural calamities, anthropogenic Orissa, India (Salu et al. 2007). This was also
activities and soil properties. The entire Akure similar to the findings of Lu et al. (2010) where
Forest reserve is a productive natural forest 60 % of the species were represented by only one or
subjected to anthropogenic activities before this two individuals ha-1. The density curve (Fig. 2) of
portion was carved out and constituted as SNR in the 15 species whose densities were ≥ 5 flattened
1954. out, similar to those reported for some protected
Biodiversity indices are generated to bring the forests of East and West Ghats and Orissa, India
diversity and abundance of species in different (Parthasarathy 2001; Salu et al. 2007). The very
habitats to similar scale for comparison and the high values of the diversity indices revealed a
higher the value, the greater the species richness forest with very high tree species diversity and
(IIRS 2002 a & b). The estimate of species abundance. The Shannon-Weiner, Pielou’s even-
diversity could come from different sources of ness, Simpson concentration, Margalef index and
which forest surveys, adopted in this present study, numbers 1 and 2 of Hill diversity indices of the
and biodiversity monitoring programmes have SNR compared favourably with the values of Lu et
been reported as major sources (Baffetta et al. al. (2010). The Shannon and Evenness indices of
2007). But, Beck & Kitching (2007) reported that this study are higher than the values of Rajkumar
the observed richness can only be a good & Parthasarathy (2008) and Yang et al. (2008). In
approximation of the true richness when it can be a similar study, Duran et al. (2006) obtained a
demonstrated that the survey is very unlikely to Shannon index that ranged between 2.69 and 3.33,
have missed any forest tree species. The SNR is which indicated a lesser diverse ecosystem than
dominated by the family Sterculiaceae with 10 the SNR. The Shannon index of this SNR is higher
important species, representing 32 % of the total than the mean of 3.34, but within the range of 2.94
stand density and highest FIV of 32.76 (31.65 % of - 3.96 that was obtained by Rao et al. (2011) for
the total FIV). Other important families domi- sacred groves in south eastern ghats, India. Sacred
nating this forest include the Meliaceae, Moraceae groves are traditional in situ conservation methods
and Mimosoideae. This finding corroborated the of biodiversity.
works of Adekunle (2006) and Adekunle et al. The horizontal and vertical structures of the
(2010) that the tropical rainforest ecosystem of forest as revealed by the diameter and height
southwest Nigeria is dominated by these set of tree distribution show a forest whose population
species and families. In a similar study, Meliaceae, structure is expanding, ensuring its stability. The
Euphorbiaceae and Moraceae were reported as the floristic composition is dominated by a suite of
families that dominated the tropical rainforest of understorey species because of the dominance of
Doi Inthanon, Thailand (Kanzaki et al. 2004), small-stemmed trees. Only 12 % of the trees could
some sites in southeast Asia (Kessler et al. 2005), be regarded as mature or big trees (dbh ≥ 40 cm)
Andaman Giant evergreen forest in India (Raj- while 88 % of them were represented by small
kumar & Parthasarathy 2008) and the Xishuang- diameter logs. The proportion of big trees in this
banna forest in southwest China (Lu et al. (2010). forest is more than the 4.5 and 3.5 % reported by
For species rarity (species represented by ≤ 2 Huang et al. (2003) and Lu et al. (2010), respec-
stems ha-1), the value obtained for the SNR (64 %) tively. In addition, the diameter distribution curve,
is lower than the 72 % that was obtained for which exhibited an inverse J-shape, indicated
tropical dry deciduous forest of Boudh District, healthy recruitment potentials. Also, the 64 % of
ADEKUNLE, OLAGOKE & AKINDELE 287

the trees occupying the middle stratum (height obtained volume with airborne laser scanning and
class of 11 - 20 m) had the highest number of an analytical formula, namely the Newton’s
species and families. This vertical and horizontal volume estimation formula (Husch et al. 2003),
structure is peculiar to mature natural forests. was used in this study. The variation in these
Mature forests are ecosystems with a recognized values could also be attributed to factors like the
ability to maintain both structure and floristic sampling intensity, inter-location variations, soil
diversity that is stable over time through the properties and the different climatic conditions.
dynamic balance of mortality, recruitment and Banda et al. (2006) and Munishi et al. (2011)
growth of plants (Stephenson & Van Mantgem observed that, generally, different management
2005). This mature phase is composed of mature regimes had significant influence on plant species
trees in various layers and a closed canopy (Saiter composition and richness, and, thereby, on forest
et al. 2011). Eventually, the mature trees die or growth and yield.
are damaged, knocking down the smaller trees
surrounding them and forming gaps and in a short Conclusions and recommendations
time, these gaps are filled with herbs, climbers and
treelets, which can arise from exposed roots and The results of this study revealed the potential
stumps or a bank of seeds and seedlings (Saiter et of an in situ method in nature conservation. The
al. 2011). The small diameter trees could also phytosociological assessment as well as the species
develop into mature trees and replace the old ones diversity and abundance compared favourably
in the future if the present conservation effort is with other similar forest ecosystems, including
maintained. those located in the biodiversity hotspots of the
The average basal area per ha-1 (22.54 m2 ha-1) world. This forest, therefore, is a potential biodi-
of this forest corroborates the range reported for versity hotspot that requires improved conser-
two of the biodiversity hotspots in the tropical vation and management efforts, and intensive
rainforest of the Ghat regions of India, i.e. 10.51- research of all the biodiversity indicators. Species
47.20 m2 ha-1 for Western Ghats (IIRS 2002a) and with low rarity index value should be considered
3.73 - 59.33 m2 ha-1 for North East (IIRS 2002b). as rare. Conservation efforts should be stepped up
The value is also close to the pan-tropical average for such species to prevent them from going into
of 32 m2 ha-1 (Dawkins 1959) and 32.3 m2 ha-1 extinction. By virtue of their narrow range, they
reported by Small et al. (2004) but less than 54.64 are usually vulnerable to extinction. The results of
m2 ha-1 reported by Yang et al. (2008) for broad - this work will serve as baseline data that could be
leaved rainforest in Mid-South Taiwan, the 47.8 helpful in the appraisal of plant resources of the
m2 ha-1 reported by Wittmann et al. (2008) for a tropical rainforest ecosystem for its effective
riparian forest of the Lower Miranda River, management. The continuous involvement of rural
Southern Pantanal, Brazil, the 59.6 m2 ha-1 for communities around the forest should be
natural forests in Chitrepani, Siwalik Region of rewarded, in form of incentives, by the government
Central Nepal by Shrestha et al. (2000) and the agency responsible for the protection and mana-
139.7 m2 ha-1 for a primary tropical forest in gement of the SNR. This is to avoid encroachment
Indonesia (Kessler et al. 2005). The value reported which may occur immediately the communities are
by Kessler et al. (2005) was one of the highest no more satisfied with using the buffer zones
values recorded in tropical forests (Lu et al. 2010). where anthropogenic activities are allowed
Tree stem volume at stand level was reported to be presently.
one of the most important parameters in forest
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computation. For instance, form factor was used by on sustainable forest management. Journal of
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(Received on 11.09.2011 and accepted after revisions, on 08.05.2012)

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