An Assessment of Canopy Stratification Ratio for Tectona grandis L.

f in Oluwa Forest Reserve

Omobolanle Temitope FALEYE1*, Lucas Aderemi AKOMOLEDE2, Olalekan Kehinde AJAYI2, Opeyemi
Philips, AKINSULIRE2

1*
Department of Forest Resources Management, University of Ibadan, Ibadan, Nigeria
2
Forestry Research Institute of Nigeria, Jericho Hill, Ibadan, Nigeria
3
Department of Botany, Obafemi Awolowo University, Ile-Ife, Nigeria

ABSTRACT: This study research was set out to investigate determined crown ratio models of for Tectona
grandis plantation in Oluwa Forest Reserve in Ondo State, of Nigeria using easily measured variables of
slenderness coefficient and merchantable height. Using Tthree non-linear regression functions was employed to
describe the tree crown ratio. in the experimental site, Tthe results showed that the basal area and tree
densitystem volume per hectare were far to be higher than the values reported for suggested for a well-stocked
most tropical rainforests in Nigeria. Few trees species were found in the higher girth and height classes.
Similarly, there were fewer tree species in the higher diameter and height class than there were in the lower
class while, there were also more trees were recorded in the co-dominant and intermediate classes than the
dominant and suppressed layer. Meanwhile, the lack of emergence in the plantation reflected the past
disturbance of the ecosystem. The higher mean basal area, stem volume, and mean tree height per hectare
recorded in the study compared with other tropical rainforest in other parts of the country are also very
significant. The mean basal area obtained in the study area was greater than 90m 2 /ha suggested for a well-
stocked forest plantation and a robust tropical forest plantation in Nigeria. Similarly, most of the tree growth
variables were significantly different under different canopy layer in the study area. However, based on the
models evaluation, the three functions investigated for tree crown ratio modeling - Chapman Richard, Logistic
and Exponential function gave constant and accurate results in all the canopy layers as indicated by going by
their indices. Chapman Richard and Exponential function gave a consistent trend and a good fit for canopy layer
crown ratio models. It Is Therefore, we recommended that strict measures should be put in place in order to
avert any level of illegality that may likely disrupts the delicate equilibrium of the ecosystem balance of the
plantation to maintaining its resilence and biodiversity., as it is also highly recommended that such results as
revealed in this study should be sustained in the region and by extension, encouraged in other parts of the
country.

Author for correspondence: amostemitope85@yahoo.com

Keywords: Crown ratio, Linear regression, Plantation, Tectona grandis, Oluwa Forest Reserve.

1
INTRODUCTION
A forest is a large area dominated by dense growth of trees and underbrush. Within the forest, complicated
layers of vegetation are common from ground cover of mosses and low flowering plants through middle layers
of bushes and secondary trees (Adekunle et al.2006). Tree crown is one complement of net primary production
and its dimensions reflect general tree health. Crown ratio is a measure of the ratio of live crown length to tree
height (ref.). It is often used as an important predictor variable for tree growth equation. It indicates tree vigour
(Hasenauer et al.1996) and wood quality (Kershaw et al. 1990), wind firmness (Navaratil 1997), stand density
(Clutter et al. 1983) and is a feature of interest in management of many non-timber resources including wildlife
habitat and recreation (Mcgaughey 1997) is a and useful parameter in forest health assessment. Dense and large
crowns are associated with potential growth rates. Sparse and small crowns can prove responsive to unfavorable
site conditions (competition, moisture, diseases) (Kozlowski et al. 1991).

Crown ratio is a useful indicator of tree vigor (Hasenauer et al.1996), wood quality (Kershaw et al. 1990), wind
firmness (Navaratil 1997), stand density (Clutter et al. 1983) and is a feature of interest in management of many
non-timber resources including wildlife habitat and recreation (Mcgaughey 1997). Crown dimensions can be
important components of forest growth and yield models, and are used in many tree and crown level growth–
modeling systems (Cole et al. 1994). For instance, tree crown parameters can be considered when simple
competition indices are not able to adequately predict recovery from competition when a competitor is removed
(e.g. by thinning) (Vanclay, 1994). Tree crown parameters have been used as predictor variables in diameter
and height growth equations (Monserud et al. 1996). Stand crown parameters have also been used to distinguish
different stages of stand development. (Soars et al.1999).

However, within a forest stand are different forms of canopy layers. These canopy layers provides protection
from strong winds and storms, while also intercepting sunlight and precipitation, leading to a relatively sparse
vegetated understory layer (Lowman, et al.1993). The canopy of a rainforest is typically about 10m thick, and
intercepts around 95% of sunlight. The canopy is below the emergent layer, a sparse layer of very tall trees,
typically one or two per hectare. With an abundance of water and a near ideal temperature in rainforests, light
and nutrients are two factors that limit tree growth from the understory to the canopy (ref).

A canopy layer is the horizontal and vertical distribution of tree crowns in a forest stand. Vertical canopy
arrangement is often simplified by dividing canopy cover into height layers. Horizontal canopy is commonly

2
quantified as the vertically-projected percentage of cover of plant canopies, and the abundance and size of
canopy gaps. Additional attributes used to describe tree crown include the number of vertical canopy layers,
heights of the vertical layers, and the proportions of cover contributed by different species groups (Fiala 2003).

Forest canopy is strongly influenced by stand density due to changing competitive interactions among the
individual trees and in turn directly influences stem-wood volume production. Stand density also influenced the
amount and distribution of leaf area in the forest stands. It is a measure of the stocking of a stand of trees based
on the number of trees per unit area and diameter at breast height of the tree of average basal area. It may also
be defined as the degree of crowding within stocked areas, using various growing space ratios based on crown
length or diameter, tree height or diameter, and spacing. Stand density index is usually well correlated with
stand volume and growth, and several variable-density yield tables have been created using it (ref).

The most commonly cited stand development model is Oliver’s (1981) four-stage model, which comprises
stand-initiation, stem-exclusion, understory-reinitiation, and old-growth phases, all of which include canopy
cover criteria. Franklin et al. (2002) proposed an alternative stand development model for natural stands. Their
model highlights eight commonly encountered development stages: disturbance and legacy creation, cohort
establishment, canopy closure, biomass accumulation/competitive exclusion, maturation, vertical
diversification, horizontal diversification, and pioneer cohort loss, with canopy attributes described for each of
these stages. Quantifying canopy structure attributes across forest stands of different ages can aid in evaluating
these stand development models. The development of understory plant communities is usually related to
changes in the over-story (Franklin et al. 2002, Henderson 1981, Naesset et al 2002, Oliver 1981, Stewart
1988, Zamora 1981). According to Connell et al. 1977 “tolerance” model of succession, shade-tolerant species
are generally present in all stages of succession, but invade the understory and increase in abundance across the
gradient of development stages.

Meanwhile, Johns (1992) It has been noted that very little of the world’s tropical rainforest area can be
considered to be are under effective management (Johns, 1992). In Nigeria, these ecosystems are poorly
managed with intense anthropogenic activities on-going. including Nigeria forest. Clear-cut logging, tree
planting, and short stand rotation lengths have greatly reduced the structural variability of forests (Garman et al.
1992; Hansen et al. 1991, 1995; Smith et al. 1996). Moreover, it is generally understood that Ttree canopy
changes as forests develop with age (Bond et al. 2002, Franklin et al. 2002, Oliver 1981, Van Pelt et al.1996),
but these changes have been rarely been quantified. The lack of data and the difficulty of accurately measuring
the height to the live crown base which is more pronounced in species with asymmetric crowns may justify the
relatively little research done on modeling crown parameters (Burkhart et al. 1992).
3
To justify this research, Tthough much is known about how forests change in the number and size and identity
of their stems (e.g., Oliver and Larson, 1990), littlescant information is available on the forest canopy (Aber,
1979; Brown and Parker, 1994) and almost nothing is known of the development of canopy layers in a forest
stand (Parker, 1995). Since, This is regrettable since: numerous functional characteristics of forests are linked
to developmental stages (e.g., Waring and Schlesinger, 1985). Therefore, it is paramount to assess the tree
crown ratio of dominant species in the forest reserve using relevant models which are generally lacking in
Nigeria. Thus, lack of information on the development of the outer canopy precludes the prediction of some
stand functional characteristics from remote information. The shape of the outer canopy as well as the different
levels below is important for several reasons. This portion of the forest is the interface of atmospheric
interactions. The extent and shape and disposition of this surface have implications for the distribution of
illuminated foliage, the penetration of heat, and the extent of turbulent mixing, among other properties.
Furthermore, the shape of the outer canopy necessarily constrains some aspects of the internal structure
included below that surface. Consequently, changes in the canopy reflect the development of the forest.

Tree crown research contributes to several key forest ecosystem attributes: biodiversity, productivity, forest
management, forest env1ironment, and wildlife. Hence crown ratio is used as an input variable to estimate
growth and mortality of individual trees and also to display changes in the appearances of stands over time for
habitat suitability and visual changes (Laar et al. 1997; Avery and Burkhart 2001). The crown displays the
leaves to allow the capture of radiant energy for photosynthesis. The size of a tree crown has a marked effect on
and is strongly correlated with the growth of the tree and its various parts (Temesgen et al. 2005). Thus,
measurement of a tree crown is often used to assist in the quantification of tree growth (Kozlowski et al. 1991).

No single model can be expected to be best for all purposes. It is therefore important to consider forest crown
under different canopy layers to provide useful alternative model for management decisions. Many authors have
based crown ratio equation on logistic functions (Hasenauer et al. 1996; Temesgen et al. 2005) or the
exponential function (Holdaway 1986; Dyer and Burhart 1987) or chapman richard function (Soares et al. 2007;
Adesoye and Oluwadare 2008). Most of these models were formulated for either even aged single species
stands, or multi-species stands comprising of different ages or mixed stand with two or more species. These
statistical functions have rarely been tested on even aged stand consisting of different canopy layers. The
purpose of this study provides information is to enhance improving the understanding of canopy at different
strata across different ages of forest stands using inventory data. Specifically, horizontal and vertical canopy
were will be compared under different layers, quantified and assessed the impact on the growth level at different
ages were assessed. Hence, this study focus on developed ing crown ratio models, stand density and growth

4
characteristics for of Tectona grandis stands under different canopy layers in Oluwa forest reserve in Ondo
State, Nigeria which are generally lacking.

Thus, the main objective of this research was to develop crown ratio models for Tectona grandis stands in
Oluwa Forest Reserve under different canopy layers. This work was also aimed at assessing tree growth
characteristics of Tectona grandis in Oluwa Forest Reserve; determining stand density index for Tectona
grandis stands in Oluwa Forest Reserve and developing crown ratio models for Tectoma grandis stands under
different canopy layers

MATERIALS AND METHODS

Study Area
The study was carried out conducted in Oluwa forest reserve located in the Ssouthern part of Ondo State,
Nigeria (Figs. 1 & 2). Four stands of different ages which covers a total area of 2000 ha were used for the study.
The forest is under the properly managedment of by the Ddepartment of Fforestry of the Ondo State
Government of Nigeria.

The study area is Oluwa Forest Reserve in Odigbo Local Government of Ondo State. Oluwa Forest Reserve is
about 828km2 in area. It is located along the Lagos-Benin expressway and is being the largest reserve in the
State. Most of the reserve lies north of the road but some (perhaps about one-sixth of the total area) lies south of
the road. Its eastern boundary lies very close to the Ondo road, a major road that goes to both Ondo and Akure
town. The area is part of the Wwestern plains of Nigeria which lies approximately between latitudes 6035/ and
7020/N, and longitudes 3045/ and 4032/ E with much of it lying approximately between 300 and 600 metre above
the sea level (Iloeje, 1981). Most rivers and streams draining this area rise from the Ssouthern part of the study
area. Notable among the rivers are Oni, Oluwa, Ominla and Owena.

The study area is under the influence of Koppens humid tropical rain forest climate. Mean annual rainfall ranges
from 1,200mm to 1,450mm and temperatures are high throughout the year with a mean of about 27 ◦C with
annual range of 3◦C (Nigeria Meteorological Observation, 2007). The natural vegetation of the area is tropical
rainforest characterized with multiple canopies and lianas. Some of the most commonly found trees in the area
include; Melicia excelsa, Afzelia bipindensis, Antiaris africana, Brachystegia nigerica, Lophira alata, Lovoa
trichiliodes, Terminalia ivorensis, T. superba, and Triplochiton scleroxylon. However, the natural vegetation of
the area, with the exception of the areas devoted to forest reserve has now been reduced to secondary re-growth
forest thickets and fallow re-growth in various stages of development or replaced by perennial and annual crops
(Osunade, 1991). These perennial crops include cocoa, kola and citrus. Most of the rural settlements in the

5
study area came into existence between 1920 and 1950 and by 1970, human colonization of the area had been
completed or tending towards completion (Adejuwon, 1971). There are, for example, many bush meat sellers
along the expressway, while Oluwa used to be contiguous with Omo Forest Reserve, there is now a dense
human population along the western boundary.

Fig.1: Administrative Map of Ondo State Showing the Study Area

Source: Ministry of Land and Housing, Akure, Ondo State, Nigeria

6
Data Collection (Sampling procedure)
Four stands of different ages were used for the study. Semi-systematic sampling method were employed in
which five (5)Ssamplinge plots of 25m x 25m (0.06 ha ) were selected randomly and laid within each stand
giving a total of twenty (20) sample plots and as total enumeration is too cumbersome and time consuming. S
systematic sampling techniques were used to assess structures of Tectonia grandis were employed for plot
laying stands within the sampling plots. Thus within each stand, five sample plots of 25m x 25m (0.0625 ha)
were laid and used for the study. These summed up to a total of twenty (20) sample plots.

Measurement of Tree Variables

Within each sample plot, tree growth variables such as total height of trees, merchantable height, and crown
length crown diameter were all measured. Similarly, Ddiameter at breast height (DBH) were measured at 1.3m
above the ground level, diameters at the base, middle, and top of the tree were also measured. In addition, trees
within each plot were classified into four canopy layers namely; dominant, co-dominant, intermediate, and
suppressed.

Data Analysis
The data collected from tree measurement were processed into suitable form for statistical analysis. Data
collected and processed includes; basal area estimation, crown projection area estimation at different strata,

7
slenderness coefficient, crown ratio of individual tree, stand volume per plot, mean crown ratio and stand
density index of trees at different layers.

Basal Area Estimation

The diameter at breast height (DBH) will be computed using this equation…………… (1)
Where,
BA = Basal Area (m2)

= 3.142

D = Diameter at breast height

Tree Slenderness coefficient computed as

TSC = ………………..... (2)

Where, TSC=Tree slenderness coefficient

THT=Tree total height (m)
D=Diameter at breast height (m)

Crown projection area estimation

CPA = ………………… (3)

Where CD = Crown diameter

Stand volume per plot

Newton’s formula ………............. (4)

Where
V = represent the stand volume
Ab = represent the cross sectional area at the base
8
Am = represent the cross sectional area at the middle
At = represent the cross sectional area at the top respectively

h = height

Stand density index estimation

Stand density index can be estimated using this formula;
The maximum density line is expressed by the equation:
log10N = -1.605 (log10D) + k……………………… (5)
Where N = number of trees per acre,
D = dbh of the tree of average basal area,
k = a constant varying with the species
When the quadratic mean diameter equals 10 inches (250 mm), the log of N equals the Stand Density Index.

In equation form:
log10SDI = -1.605(1) + k…………………………… (6)
Which means that: k = log10SDI + 1.605
Substituting the value of k above into the reference-curve formula gives the equation:
log10N = log10SDI + 1.605 - 1.605 log10D
This equation can be rewritten as:
log10SDI = log10N + 1.605 log10D - 1.605…………………………………… (7)

The above equation is an expression for computing the stand density index from the number of trees per acre
and the diameter of the tree of average basal area.

Estimates of stand density will express the degree to which the growing space available for tree growth is
utilized. Thus, stand density is a function of three elements:

1. Number of trees - which is readily determined by counting.

2. Tree size - which involves a number of factors, e.g.:-

o Stem - characterized by diameter, height and taper

o Crown - characterized by spread and depth

o Root - characterized by spread and depth (both difficult to measure).

9
 3. Spatial distribution on the ground - which is not readily determined. Generally, a square or triangular
spacing is assumed.

In even-aged stands, crown closure may be proportional to basal area/ha. This relationship has led to the
development of indices between estimates of crown closure obtained from aerial photographs and basal area.
The value of crown closure as a variable depends on how well variation in stand volume is correlated with it.

The functional form of this relationship is:

Log10 N = k - 1.605(Log10 QMD) ……………………………………. (8)

Where, N =number of trees per hectare,
K = a species specific constant,
QMD = quadratic mean diameter

Crown ratio model

The tree crown ratio models that will be employ for this study are; Logistic function, Chapman-Richard and
Exponential models since a crown ratio value ranges from 0 (i.e. no crown, dead or defoliated) to 1 (i.e. crown
extends over the entire tree bole). The tree crown ratio model formulated to express crown ratio as a function of
tree size (e.g. basal area, merchantable height)
The original forms of the models are:

-1
CR = …………………. (9)

CR = ………………………. (10)

CR = bo + (11)

Where,

10
CR is estimated crown ratio, βx is linear equation with parameter β and independent variable x (which includes
individual tree characteristics such as merchantable height-MHT, Diameter at breast height, crown length and
total height) and e is Naperian constant.
The multiple linear regression equation for the independent variables is given as follows:
Bx = βo +bx1 + cx2 + dx3 …………………………… (12)
Where,

is intercept,

b to d is set of parameters,
x1 to x3 is the independent variables.

CR = ]-1 ……………………. (13)

RESULTS AND DISCUSSION

A total of 1042 individual trees were recorded and identified from the sampling plot covering an area of 2000ha
in the study area. A total of 95 trees sampled were recorded in the dominant canopy layer which covered about
9.1% of the total population of the forest reserve plantation. About 267 of the individual trees species were
recorded in the co-dominant class which accounted for about 25.6% and 403 trees were recorded for in the
intermediate height class which accounted with 38.7% of the total population. while Tthe suppressed class
accounted for about 277 of the individual trees species which contributed were found to be 26.6 % to the total
population (appendix 6).

The trees stand density in this study had higher values compared to reported values of some authors who
worked in same eco region numbers of individual trees per hectare obtained in the forest plantation were higher
compared to those reported by (Adekunle et.al (2004);, Ojo (2004), ;Adekunle and Olagoke (2008) for other
tropical rainforests of Nigeria. This is an indicationg that the protection of the forest is properly managed.
plantation is very effective. The mean basal area of the stands in the per hectare obtained for the forest
plantation (92.64785m2/ha) compared well with values() that reported in Onigambari forest reserve (Akinyemi
et. al 2002) and in Nigeria and also higher than the reported research at Omo forest reserve by (Adekunle, 2007)
in Nigeria. It was also discovered that the value of Tthe mean basal area of the stands were higher compared to
those reported by (Temesgen et al 2005) a research conducted on multispecies and multilayered stands of
11
southeastern British Columbia. The mean stem volume values obtained for the forest plantation
(7184.4573m2/ha) were far higher than those reported () by some authors who worked in tropical rainforest
ecosystem in Nigeria (Adekunle et al (;2004;), Alder and Abayomi, (1994), ;Adekunle and Olagoke (2008) for
tropical rainforest ecosystem in Nigeria.

Correlation among tree growth variables for individual dominant trees

The karl Pearson’s product correlation co efficient for the growth variables are presented in Tables 1-4. The
correlation values obtained for individual dominant trees variedy between -0.012 and 0.681. Most of the trees
growth variables are have significantly positive correlated ion with one another. The highest r-value was
between mht and tht (r = 0.681); this was followed by that between < dbh and tht (r =0.618) < and dbh and mth
(r = 0.534). Small Negative r-value were obtained between dbh and cl (r = -.012); mht and sc (r = -0.305); dbh
and cr (r = -0.300); tht and cr (r =-0.260). Meanwhile Tthe correlation values obtained for individual co-
dominant trees variedy between -0.85 and 0780. Most of the tree growth variables are have significantly
positive correlated ion with one another. The highest r-value was between mht and tht (r = 0.780) <; this was
followed by that between cl and cr (r = 0.776) < and dbh and mth (r = 0.656). Negative Small r- value were
obtained between mht and cl <, tht and cr with (r = -.085); <dbh and cr (r = -0.150); < tht and sc (r = -0.217).

The correlation values recorded for individual trees in the intermediate layer vary between - 0.34 and 0.919.
Most of the tree growth variables have significant positive correlation with one another. The highest r-value was
between cl and cr (r = 0.919); this was followed by that between dbh and sc (r =0.566); tht and sc (r = 0.519)
and dbh and mht (r = 0.508) Small r- value were obtained between dbh and cr, (-0.34); tht and cr (-0.61); cr and
sc, mht and cl with (r = 0.28). Furthermore, the values obtained for suppressed trees vary between -0.235 and
0.765. Most of the tree growth variables have significant positive correlation with one another. The highest r-
value was between cl and cr (r = 0.765); this was followed by that between dbh and sc (r =0.538); mht and sc (r
= 0.403) and dbh and cl (r = 0.380). Small r- value were obtained between tht and cr, (-0.235); cl and sc (-
0.244); cr and sc, (r = -0.255).

Generally, most of the tree growth variables were significantly and positively correlated with one another,
which impliedy that an increase in the one, tend to be associated with an increase in the other variable(s). For
instance, tree basal area correlated well with diameter at breast height since diameter at breast height was input
variable in the computation of the tree basal area. However, tree slenderness coefficient and the tree crown
12
ratio, crown length and slenderness coefficient had a weak correlation (0.022, and 0.025) respectively. There
were Nnegative correlation were obtained between crown ratio and diameter at breast height, crown length and
diameter at breast height, crown ratio and total height. These negative correlation trends are expected which
indicated that trees with small diameter, tall and slender had lower crown ratio values.

Table 1: correlation matrix for individual dominant tree growth variables

DBH MHT THT CL SC CR
DBH 1
MHT .534** 1
THT .618** .681** 1
CL -.012 -.559** .224** 1
SC -.847** -.305** -.272** .099 1
CR -.300** -.880** -.260** .878** .223** 1
Correlation is significant at 0.05 level (2- tail)
Total height (tht), merchantable height (mht), diameter breast height (dbh), slender coefficient (sc), crown
length (cl) and crown ratio (cr)

Table 2: correlation matrix for individual co-dominant tree growth variable

DBH MHT THT CL SC CR
DBH 1
MHT .567** 1
THT .656** .780** 1
CL .278** -.085 .542** 1
SC -.723** -.270** -.212** .022 1
CR -.150** -.660** -.085 .776** .211** 1
Correlation is significant at 0.05level (2- tail)
Total height (tht), merchantable height (mht), diameter breast height (dbh), slender coefficient (sc), crown
length (cl) and crown ratio (cr)

13
 Table 3: correlation matrix for individual intermediate tree growth variables
DBH MHT THT CL SC
CR
DBH 1
MHT .508** 1
THT .280** .296** 1
CL .119** .028 .180** 1
SC -.566** -.173** .519** .028 1
CR -.034 -.297** -.061 .919** .025
1
Correlation is significant at 0.05level (2- tail)
Total height (tht), merchantable height (mht), diameter breast height (dbh), slender coefficient (sc), crown
length (cl) and crown ratio (cr)

Table 4: correlation matrix for individual suppressed tree growth variables

DBH MHT THT CL SC
CR
DBH 1
MHT .186** 1
THT .393** -.698** 1
CL .380** .538** .286** 1
SC .639** .403** .160 -.244** 1
CR .345** .376** -.235** .765** -.255** 1
correlation is significant at 0.05level (2- tail)
14
 total height (tht), merchantable height (mht), diameter breast height (dbh), slender
coefficient (sc),crown length (cl) and crown ratio (cr)

Basal area and stem volume estimation

Information on tree basal area and stem volume per hectare for the forest plantation is presented in appendix 5.
The forest had an average basal area of 92.64785m2/ha while the average trees stem volume recorded
7184.4573m2/ha.

Height (m) distribution of tree per plot into canopy layers in the study area
The distribution into height classes corresponding to each of the strata (canopy layers) in the study area are is
shown in table 5. Four strata existed in the all the plots sampled in the plantation. Trees belonging to the
dominant height class (measured> 25m and above) accounted for about 9.1% of the individuals sampled in the
plantation. About 25.6% of the forest plantation were found in belongs to the co-dominant layer (height range
of 20 m-24.9 m). The intermediate layer (height class of 14m-19.9m) accounted for about 38.7% of in the forest
plantation area sampled within the forestrespectively. The suppressed layer, with trees height class ≤ 13.9m,
accounted for about 26.6% of the individuals sampled in the plantation. There was a decline major drop in the
number of trees attaining reaching above 25m height. Also there were Ffewer numbers of individuals in the
dominant height class was noted in the area than there were in the suppressed and co-dominant height classes.
The intermediate class recorded the highest number of individuals in the overall canopy layer.

Table 5: height (m) distribution of tree per hectare into canopy layers in the study area

Canopy layer Height class (m) Number of trees per hectare

Dominant ≥ 24 95
Co-dominant 16-23 267
Intermediate 10m-15 403
Suppressed ≤ 10m 278
Total 1042

Crown ratio model

15
Three non-linear regression models (Logistics, Richards, and Exponential) were fitted into the data in the study
area. All the tree growth variables apart from the crown ratio (the dependent variable), were tried during model
fitting processes. The selected version of the Logistics, Richards and Exponential models, their parameters
estimation and fit statistics for the canopy layers are represented in table 6, 7, 8, and 9 respectively.
Merchantable height and slenderness coefficient were found to consistently predict crown ratio in all the
functions.

The R2 values for the three functions were fairly high under the dominant, and co-dominant and intermediate
layer with low values of standard errors of estimates (SEE). The suppressed layer which showed gave a lower
fit to the data set in the functions however produced significant result for the estimated parameters in all the
functions.

However, there are significant differencet among growth variables under different canopy layers was reflected
for the models used, hence they three models are fitted for to the data set on the layers basis. The merchantable
height and slenderness coefficient gave better fit to the data set and were found to be important in defining the
tree crown ratio for Tectona grandis stand in Oluwa forest reserve. The suitability of the other tree growth
variables wereas investigated. They were Hhowever, they failed to explain the tree crown ratio in the entire
canopy layer, and were therefore not included in the model presentation result. The R 2 values for the three
functions were consistently high for under dominant, co-dominant, and intermediate layers with low standard
error of estimate (SEE). The suppressed layers, with thewhich gave much lowestr fit to the data set in all the
functions, were however produced significant result infor all the estimated parameters forin all the functions.
The R2 value obtained in this study were generally higher than in comparison with those reported fromby the
previous research infor less diverse ecosystem (Temesgen et.al 2005;and Adesoye and Oluwadare 2008).
Hence, the model best fitted in this study than previous ones.with lower SEE value. This indicates better fit of
the three functions to the data set than those fitted by previous workers.

Table 6: Tree crown ratio models selected with parameter estimate and fit statistic for dominant layers in
Oluwa forest reserve
Function parameter estimate SE
Logistic a0 1.213 0.144

a1 -0.107 0.007

R2 = 0.657; SEE = 0.2 a2 0.000 0.001

Richards a0 0.085 0.217

16
 a1 0.174 0.011

R2 = 0.704; SEE = 0.173 a2 -0.004 0.001

Exponential a0 0.188 0.052

a1 -0.133 0.028

R2 = 0.711; SEE = 0.173 a2 0.001 0.001

Table7: Tree crown ratio models selected with parameter estimate and fit statistic for co-dominant
layers in Oluwa forest reserve
Function parameter estimate SE
Logistic a0 0.274 0.094

a1 -0.110 0.006

R2 = 0.643; SEE = 0.2 a2 0.003 0.001

Richards a0 0.585 0.024

a1 0.273 0.025

R2 = 0.763; SEE = 0.2 a2 0.022 0.003

Exponential a0 -0.400 0.027

a1 0.032 0.003

R2 = 0.738; SEE = 0.2 a2 0.001 0.000

Table 8: Tree crown ratio models selected with parameter estimate and fit statistic for intermediate layer in Oluwa
forest reserve
Function parameter estimate SE
Logistic a0 0.245 0.032

a1 0.025 0.002

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 R2 = 0.647; SEE =0.158 a2 0.001 0.000
Richards a0 -0.614 0.059

a1 0.173 0.071

R2 = 0.747; SEE = 0.167 a2 -0.015 0.003

Exponential a0 -0.066 0.071

a1 -0.046 0.010

R2 = 0.720; SEE = 0.164 a2 -5.743 3.822

Table 9: Tree crown ratio models selected with parameter estimate and fit statistic for suppressed layer in Oluwa
forest reserve
Function parameter estimate SE
Logistic a0 -1.256 0.086

a1 0.024 0.008

R2 = 0.410; SEE = 0.152 a2 0.0001 0.001

Richards a0 -0.555 0.074

a1 14.161 1.494

R2 = 0.430; SEE = 0.158 a2 0.001 141.224

Exponential a0 -0.470 0.016

a1 -32.278 6.873

R2 = 0.427; SEE = 0.158 a2 0.232 49387.189

Mean comparison of the growth variables for the canopy layers

The mean slenderness coefficient had direct link withdecreases with decreasing crown ratio at different stages
of the canopy layer (table 10 -13). Dominant class had the highest value and suppressed individual class

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recorded the lowest value, this indicates that trees that are tall and slender had lower crown ratio values.
However Tthe average diameter at breast height showedfollowed the same trend with as the dominant class
havingrecorded the highest value (27.19) and the suppressed class had the lowest(10.4772).

Table 10: Mean comparison of the growth variables for the dominant canopy layer
Growth variables Mean Minimum Maximum S.D

BA 0.0648 0.0072 0.1768 0.03934

DBH 27.19 9.55 47.43 0.7300
MHT 14.43 6.600 21.200 3.3188
SC 97.5788 533.477 209.424 31.4212
CR 0.4118 0.1724 0.7203 0.1044

Table 11: Mean comparison of the growth variables for the co-dominant canopy layers
Growth variables Mean Minimum Maximum S.D

BA 0.049 0.0016 0.203 0.03264

DBH 23.68 4.45 56.93 8.01
MHT 12.6011 4.5000 19.8000 3.08575
SC 94.0571 49.0870 330.3371 33.4477
CR 0.3796 0.4163 0.7000 0.0000

Table 12: Mean comparison of the growth variables for the intermediate canopy layers
Growth variables Mean Minimum Maximum S.D

BA 0.05726 0.0016 0.5216 0.06149

DBH 19.081 4.46 52.2 7.9973
MHT 9.7489 3.8000 17.4000 2.4765
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 SC 92.6185 29.6933 583.8767 39.9820
CR 0.3572 0.0416 0.8239 0.0054

Table 13: Mean comparison of the growth variables for the suppressed canopy layers
Growth variables Mean Minimum Maximum S.D

BA 0.159 0.000 0.159 0.67

DBH 10.4772 0.0255 0.923 0.008384
MHT 4.8789 1.7000 14.9000 0.2099
SC 90.4005 6.2832 202.7972 36.3824
CR 0.3736 0.980 0.7799 0.0109

Relationship between residual and estimated Crown ratio

The evaluation of the residual plots (Fig. 1-5), indicating its error analysis showedrevealed that error variance
wasis constant across the predicted crown ratio. Chapman-Richard logistics and Exponential functions were
observed to have constant error variance.

In the graphical relationship between the residuals and estimated crown ratio obtained with the three functions,
all the residual values are in the positive and negative region which implied that crown ratio values were
consistently predicted. Exponential and Chapman-Richard functions judging from their error analysis appeared
best in the three functions because constant error variance distributed well both in the positive and negative
region of the x-axis (i.e. the estimated crown ratio values) which . This is desirable for a good model and. tThis
trend was similar to the findings of Soares and Tome (2001) and; Adesoye and Oluwadare (2008). Based on the
evaluation of the error analysis, Chapman-Richard and Exponential functions are recommended for predicting
crown ratio in the stand because reflected they are more precise option for predicting variables considered in
this study.in their predictive ability.

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Model fitting and evaluation
Model fitting and evaluation are important parts of model building. Fitting of crown ratio models were based on
the total data set. A number of different models were examined for predicting crown ratio using logistic,
exponential and Chapman Richard functions. In this study coefficient of determination (R 2) and standard error
of estimate (SEE) were computed in order to evaluate the fitted models. In addition, residual plots were carried
out to check the error assumption. The level of significance of the parameter estimates was also testedobserved.
The selected versions of the models are presented in (appendix 1 -4). One unique independent variable that
features in all the models wasis tree slenderness coefficient. This proves that tree slenderness coefficient is one
of the factors that contributeding to the size of tree crown ratio. Some The pattern was observed in the studies
conducted have shown similar pattern by (Hanus et al. (2000); Hasenauer and Monserud (1996); Hann, (1997).
Merchantable height was noted to be another important variable used in modeling in this study. Adesoye and
Oluwadare (2008); Marshall et al. (2003) also found merchantable height as an important variable for modeling
crown. The other variables failed to adequately explain crown ratio variation and were therefore not included in
the models. The SEE differences were small since crown ratio wasis constrained to the interval of 0 and 1. This
was also notediced in the studywork carried out by Temesgen et al. (2005) and Adesoye and Oluwadare (2008).
Generally, the models reflectedconsistently gave good fit of to the Tectona grandis plantation data in Oluwa
Forest Reserve. The evaluation of the residual plots (Fig. 1-5) indicatingrevealed the error analysis reflected that
error variance wasis constant across the predicted crown ratio. Chapman-Richard and Exponential functions
were observed to have constant error variance. In the graphical relationship between the residuals and estimated
crown ratio obtained fromwith the logistic, all the residual values are in both negative and positive region but
are farther from one another which implied that crown ratio values were slightly consistent. Exponential and
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Chapman-Richard functions judging from their error analysis appeared constant error variance distributed both
in the positive and negative region of the x-axis (i.e. the estimated crown ratio values), which . This is desirable
for a good model. This trend was similar to the findings of Soares and Tome (2001); Adesoye and Oluwadare
(2008). Based on the evaluation of the error analysis, Chapman-Richard and Exponential functions are
recommended for predicting crown ratio in the stands. Although the R2 values for suppressed canopy layer were
lower compared to other strata, they are more precise in their predictive ability.

CONCLUSIONS AND RECOMMENDATIONS

Live crown ratio models for Tectona grandis stand in Oluwa Forest Reserve were fitted using easily measured
variables of slenderness coefficient and merchantable height. The result of the study providedgave some
explanation on the uses of three non-linear regression functions to describe the tree crown ratio of Tectona
grandis plantation in Oluwa Forest Reserve. Based on thise examined result of the study, the basal area and tree
stem volume per hectare in forest plantation were far higher than the values suggested for a well-stocked
tropical rainforest in Nigeria.

There were fewer tree species in the higher diameter and height classes than there were in the lower classes
respectively. These set of higher diameter classes formed the dominant classes which could be as a result of
their early and fast growth. There were more trees recorded in the co-dominant and intermediate classes than the
dominant and suppressed layer. The lack of tree that could be classified as emergence reflected the past
disturbance of the ecosystem. The higher mean basal area, stem volume, and mean tree height per hectare
recorded in the study compared other tropical rainforest in other part of the country are also very significant.
The mean basal area obtained in the study area is greater than 90m 2 suggested for a well-stocked forest
plantation and a robust tropical forest plantation in Nigeria. These is evident that there are level of commitment
by the stakeholder as it compare well with what is obtainable in other part of west Africa with similar
ecosystem. It is highly recommended that such should be sustained in the region and by extension, encourage in
other part of the country.

Most of the tree growth variables showed were significantly differencet under different canopy layer in the
study area. The forest canopy wereas well diverse and the tree growth variables compared considerably well
with each other. It is recommended on this note that strict measure should be put in place to prevent any level of
illegality that may likely disrupts the delicate equilibrium of the ecosystem. However, based on the models
evaluated,ion the three functions investigated in the study for tree crown ratio modeling indicated, Chapman
Richard, Logistic and Exponential function weregave constant and reflected an accurate result in all the canopy
layers, going by their indices. Notably, Chapman Richard and Exponential function showed gave a consistent
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trend and a good fit for predicting canopy layer crown ratio models in for Tectona grandis plantation of the in
Oluwa forest reserve.

ACKNOWLEDGEMENT
The authors wish to acknowledge Professor P.O Adesoye of the University of Venda, South Africa, for the
assistance rendered regarding design and methodology of this study. We all acknowledge Mr Afolabi gbenga
and Mr Olugade Tayo both in the Department of Forest Resources Management, University of Ibadan, for the
technical assistance rendered. Anonymous reviewers are also greatly appreciated for their constructive
comments.

CONFLICT OF INTEREST
The authors have declared that there is no conflict of interest.

REFERENCES (Have been removed)

APPENDIX
Appendix 1: selected tree crown ratio model for dominant stratum
Function Models R2 SEE
Logistics 0.657 0.2

Richards 0.704 0.173

Exponential 0.711 0.173

Appendix 2: selected tree crown ratio model for co-dominant stratum

Function Models R2 SEE

Logistics 0.643 0.2

Richards 0.763 0.2

Exponential 0.738 0.2

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 #

Appendix 3: selected tree crown ratio model for intermediate stratum

Function Models R2 SEE
Logistics 0.647 0.156

Richards 0.747 0.167

Exponential 0.720 0.164

Appendix 4: selected tree crown ratio model for suppressed stratum

Function Models R2 SEE
Logistics 0.412 0.152

Richards 0.430 0.158

Exponential 0.427 0.158

Appendix 5: Tree basal area (m2) and stem volume per plot in forest reserve
Plot basal area stem volume

1 43.286 3,716.887
2 49.560 4,204.157
3 53.022 4,816.724
4 53.257 4,941.889
5 38.766 3,476.266
6 133.910 10,259.627
7 160.173 10,477.570
8 101.550 3,064.326
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 9 106.043 7892.709
10 151.607 11,081.719
11 132.937 9,215.091
12 136.927 9,368.108
13 78.872 5,544.587
14 112.215 8,478.861
15 147.011 9,598.887
16 82.612 7,182.559
17 103.260 8,616.163
18 64.327 6,460.889
19 76.533 7,214.863
20 87.089 8,077.264
Total 1912.957 143689.146
Mean 92.64785 7184.4573

Appendix 6: Tree canopy and percentages of individual layers

Canopy layer Number of trees Percentage (%)

Dominant 95 9.1

Co-dominant 267 25.6

Intermediate 403 38.7

Suppressed 277 26.6

Total 1042 100

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