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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.
Key words: Biodiversity indices, forest structure hotspots, in-situ conservation, stand
volume.
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 )
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
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 5. Status of tree species in the SNR according to diameter (cm) and height classes.
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.
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
management, and its acquisition is very time References
consuming and expensive because it is normally
obtained from field surveys (Tonolli et al. 2011). Adekunle, V. A. J. 2006. Conservation of tree species
Therefore, for volume estimation, the 145.22 m3 ha-1 diversity in tropical rainforest ecosystem of south-
for this study is less than 391 m3 ha-1 reported by west Nigeria. Journal of Tropical Forest Science 18:
Wittmann et al. (2008) and the 406 - 416 m3 ha-1 91-101.
reported by Tonolli et al. (2011) for multilayer Adekunle, V. A. J., A.O. Olagoke & L.F. Ogundare. 2010.
forest areas in Italian Alps. The difference may be Rate of timber production in a tropical rainforest
due to the different methods adopted for volume ecosystem of southwest Nigeria and its implications
computation. For instance, form factor was used by on sustainable forest management. Journal of
Wittmann et al. (2008), Tonolli et al. (2011) Forestry Research 21:225-230.
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