International Research Journal of Plant Science (ISSN: 2141-5447) Vol. 4(8) pp. 214-221, September, 2013
DOI: http:/dx.doi.org/10.14303/irjps.2013.047
Available online http://www.interesjournals.org/IRJPS
Copyright © 2013 International Research Journals
Full Length Research Paper
The Spatial Trends in the Structural Characteristics of
Mangrove Forest at the Rawa Aopa Watumohai National
Park, Southeast Sulawesi, Indonesia
Kangkuso Analuddin*1/2, Jamili1, Rasas Raya3, Andi Septiana1 and Saban Rahim2
1
Department of Biology, Faculty of Mathemathics and Natural Sciences, Halu Oleo University Kendari, Southeast
Sulawesi, Indonesia
2
The Museum and Wallacea Research Center, Halu Oleo University Kendari, Southeast Sulawesi, Indonesia
3
Department of Mathemathics, Faculty of Mathemathics and Natural Sciences, Halu Oleo University Kendari, Southeast
Sulawesi, Indonesia
*Corresponding authors e-mail: zanzarafli@gmail.com;udin_zanza@yahoo.com
Abstract
Mangrove ecosystem at the Rawa Aopa Watumohai (RAW) National Park is one of among the most
protected ecosystems in the Southeast Sulawesi Region of Indonesia. The present study was aimed to
elucidate the structural characteristics of mangrove forest at RAW National Park, knowledge of which
fundamental for conservation and management that of National Park. The growth parameters of
mangrove species including stem diameter at breast height DBH, tree height H and crown diameter
CD were measured in the 50 quadrats of 100 m2 wide each, which were placed near the seashore, midle
area and near the land of mangrove forest. The Rhizophora apiculata and Rhizophora mucronata were
the most dominant mangrove near the seashore and midle sites, while Ceriops tagal and Lumnitzera
Littorea were the most dominant mangrove near the land site. The spatial structural of mangroves at
the Rawa Aopa Watumohai National Park seems to realize the zonation because many similarity index
values of paired stands were estimated as zero. The skewness of DBH for R. apiculata and R.
mucronata was found nearly zero, which indicated normal distribution, while the skewness of DBH for
Ceriops tagal and Lumnitzera littorea was possitive, which indicated L-shaped distribution.
Keywords: Mangrove forest, size distribution, Rawa Aopa Watumohai National Park, Indonesia
INTRODUCTION
Mangroves are one of among the most important
ecosystems, because they play very important roles for
the carbon storage in the subtropical and tropical coastal
areas of the world. The structural development and
growth of mangrove forest are influenced by the
hydrological processes, such as tidal activity, waver, river
discharge, and freshwater inputs, which have a
particularly strong influence on the chemical and physical
conditions in mangrove ecosystem (Sherman et al.
2003). Edaphic factors, such as water logging, nutrient
availability and salinity have great impact on the
mangrove growth and productivity. However, the growth
parameters of mangrove trees such as tree height, stem
diemeter, height at the lowest living leaves and crown
structure are playing important function during their life
time. Therefore, maintenances the tree size structural of
mangrove trees are important for their self-regulation,
which are essential for ensuring how the mangrove trees
could function.
Many scientits realize the important role of crown
structure of plants for their functioning and stand
productivity (e.g.- Pearcy and Yang 1996; Osada and
Takeda 2003; Weiskittel et al. 2007). Crown structure
influences on competition and survival abilities of plant in
the community (e.g.-Kuulivainen1992). The leaves
arrangement within the crown influences many
aspects of whole-plant function including photosynthesis,
transpiration and energy balance (Pearcy and Yang
Analuddin et al. 215
1996). As trees including mangroves grow larger, they
have to produce sound crown to make their life function.
For this reason, study on tree crown structure provides
critical information to assess a variety of ecological
functioning mechanisms of forests including mangroves.
Many researchers have examined the crown of trees in
relations to structure and shape, while others have
concentrated upon modeling of the impact of shape on
physiological activity of tree crown. Measurement of a
tree crown is often used to assist in the quantification of
tree growth (Kozlowski et al. 1991). The surface area of
forest trees is a useful measurement for the study of
rainfall interception, light transmission through forest
canopies, forest litter accumulation, soil moisture loss,
and transpiration rates (Husch et al. 2003).
The knowledge concerning tree crown structure is
important in the sense that trees use this tree component
as a source of absorption of carbon dioxide (Hussein
2001). Taking into consideration the crown as a primary
element in the vegetation development, several scientific
studies relating to tree crown have been undertaken to
determine tree growth through models for tree crown
profiles (Gadow 1999; Gill et al. 2000). Much interest has
been paid to analyze the tree height-crown shape
supporting mechanism (e.g. Givnish 1986), growth
strategies and allocation pattern to crown dimension (e.g.
King 1990; Kohyama and Hotta 1990), but little
information is known about tree growth parameters
maintenace for mangrove forests (e.g. Analuddin et al.
2009c). In the former study, Analuddin et al. (2009a)
found the changes in the tree size inequality of
monospecific mangrove Kandelia obovata stand as the
stands grew. Meanwhile, the study on spatial trends in
the structural characteristics and growth parameters
development of mangrove is scarse.
Mangrove forest at the Rawa Aopa Watumohai
National Park is one of among the most fascinating and
protected forests in the Southeast Sulawesi Region of
Indonesia. However, little scientific information is known
concerning this mangrove forest. An investigation on the
trends in the structural characteristics including tree
growth parameters performance and changes in the
compositional of mangroves from the seashore to the
landward sites are important, knowledge of which can
improve the understanding of the present condition of
mangroves. These information are ensential for
conservation of the mangroves at the Rawa Aopa
Watumohai National Park. Therefore, the objectives of
this study were: (1) to elucidate the structural
characteristics of mangrove trees grown along the
Lanowulu River; and (2) to know the self- maintenances
of mangrove forest in the Rawa Aopa Watumohai
National Park. The data were analyzed to elucidate the
structural characteristics of mangrove forest by estmating
on the importance value index, similarity index and
exponent value of self-thinning line of mangroves stands.
In addition, we assesed the size inequality of mangroves
dominant by analyses on the frequency distribution and
coefficient of variance of the mangrove trees growth
parameters.
MATERIALS AND METHODS
Study site
The present study was carried out at the mangrove forest
of Rawa Aopa Watumohai National Park, which is located
at the
at eastern part of Kendari city, Southeast
Sulawesi, Indonesia. The mangrove condition provides
an excellent site for studying mangrove forest
conservation, because this is the only pure mangrove
condition till now, while many sorrounded mangrove have
been degraded due to marine aquculture development.
The mangrove forest is one of most important vegetation
here, because this mangrove forest becomes habitat for
various organisms, including crocodile, and endemic
animals of birds and Bubalus sp, which is ane of among
the most protected animals in Indonesia. The growth and
development of mangroves in this sites were continuosly
supported by Lanowulu River, because it flows inside the
mangrove vegetation. The Lanowulu river becomes
habitat for various marine organisms, and provides an
easy way to access the mangrove forest.
Data Analyses
The community structure of mangrove forest was
quantitatively analyzed, such as species density and
dominance as well as their importance value index. In
addition, the spatial trend of mangrove forest was
elucidate by analyzing the similarity index of mangrove
among stands using Bray and Curtis Formula. On the
other hand, size inequality of the mangrove tree growth
parameters, such as coefficient of variation and
skewness frequency distribution of tree heigh H and stem
diameter at breast heigh DBH, were evaluated for each
dominant mangrove species. The coefficients of the
curvilinear and nonlinear equations for the relationships
among growth parameters in each species, were
obtained by software (KaleidaGraph ver. 4,0, Synergy
Software).
RESULTS AND DISCUSSION
RESULTS
Spatial trend in mangrove structure
Table 1 shows the structure of mangrove in each stand at
the Rawa Aopa Watumohai National Park. There was
216 Int. Res. J. Plant Sci.
Table 1. Mangrove forest structure at the study site of Rawa Aopa Watumohai National Park,
Southeast Sulawesi, Indonesia
Location
Stands
Nama Jenis
Near the
land site
1
Lumnitzera littorea
Ceriops tagal
Rhizopora apiculata
Ceriops tagal
Lumnitzera littorea
Lumnitzera littorea
Ceriops tagal
Rhizopora apiculata
Rhizopora stylosa
Xilocarpus granatum
Bruguiera gymnorhiza
Rhizopora apiculata
Rhizopora stylosa
Bruguiera gymnorhiza
Rhizopora mucronata
Ceriops tagal
Ceriops decandra
Xilocarpus granatum
Rhizopora apiculata
Rhizopora stylosa
Bruguiera gymnorhiza
Xilocarpus granatum
Avicennia alba
Rhizopora mucronata
Bruguiera gymnorhiza
Rhizopora apiculata
Xilocarpus granatum
Bruguiera gymnorhiza
2
3
4
5
6
7
Midle site
8
9
Near the
seashore
10
existed nine major mangrove species, which showed
various spatial structure. The Lumnitzera littorea was the
higher importance value index (IVI) or the most dominant
mangrove species at the stands 1 and 3, while Ceriops
tagal was the highest IVI or the most dominant mangrove
at the stand 2. On the other hand, the Rhizophora
apiculata was the most dominant mangrove species at
the stands 4, 5, 8 and stand 10. Furthermore, the
Rhizophora mucronata was most dominant mangrove at
the stands 6 and 9. However, the Ceriops tagal was the
most dominant mangrove at the stand 7.
Mangrove structure according to the site condition
showed different trend from the seashore to the landward
sites. As shown in Table 1 that the Ceriops tagal and
Lumnitzera littorea were the two dominant mangroves
species near the landward site, while the Rhizophora
apiculata and Rhizophora mucronata were dominant in
the middle and near the seashore of Rawa Aopa
Density
(Ind/Ha)
2400
1300
100
2720
580
1000
1220
940
120
220
120
560
40
180
1600
1420
80
540
780
40
280
120
180
860
40
900
80
120
Basal area
2
(m /Ha)
1002
568
115
675
129
1673
492
4437
912
903
365
5978
276
1510
5755
1843
77
1215
5995
344
1525
246
682
3613
235
5688
141
605
Important value
index (%)
172,60
105,42
21,98
216,40
83,60
172,31
127,69
169,91
36,64
57,93
35,52
198,79
18,68
82,53
300,00
173,86
15,46
110,68
179,51
15,86
75,09
29,54
46,02
230,81
23,18
215,67
36,74
47,59
Watumohai National Park. On the other hand, Rhizopora
stylosa and Bruguiera gymnorrhiza were distributed
sparse, while Avicenia alba was just found near the
seashore only.
Table 2 descibes the similarity index among mangrove
stands along the Lanowulu river. Many similarity index
(SI) among stands were estimated as zero, which means
that mangrove composition for many mangrove stands
were completely different. There was six paired stands
only having SI values more than 70%, while it was nearly
30% paired stands having SI values less than 10%
meaning low degree similarity of mangrove among
stands at the study site. These trends seem to realize the
spatial zonation of mangrove species at the Rawa Aopa
Watumahi National Park, Southeast sulawesi indonesia.
However, the similarity index of mangroves between near
the land and other sites was found less than 50%
meaning low degree similarity, while SI of mangrove
Analuddin et al. 217
Table 2. Similarity index among mangrove stands at the study site of Rawa Aopa Watumohai National Park,
Southeast Sulawesi, Indonesia
S
T
A
N
D
S
SI/DI
1
2
3
4
5
6
7
8
9
10
1
2
63,01
3
92,67
70,43
4
7,33
7,33
0
STANDS
5
7,33
7,33
0
62,86
6
0
0
0
0
0
7
35,14
35,14
35,14
19,31
36,89
0
8
7,33
7,33
0,00
71,77
90,15
0
9,85
36,99
7,33
92,67
92,67
100
64,86
92,67
100
29,57
92,67
92,67
100
64,86
92,67
100
100
100
100
64,86
100
100
37,14
100
80,69
28,23
100
100
63,11
9,85
92,27
100
100
23,06
90,15
100
92,27
92,67
92,67
100
31,12
17,87
100
87,75
40,16
9
0
0
0
0
7,73
76,94
0
7,73
10
7,33
7,33
0,00
68,88
82,13
0,00
12,25
59,84
7,73
92,27
Similarity index SI (upper) and disimilarity index DI (below)
Table 3. Size distribution of four dominant mangrove species along the Lanowulu River, Rawa Aopa
Watumohai National Park
Mangrove species
Growth
parameters
DBH
Rhizophora apiculata
H (m)
CD (m)
CPA (m2)
DBH
H
CD
CPA
DBH
H (m)
CD
CPA
DBH
H (m)
CD
CPA
Rhizophora mucronata
Lumnitzera racemosa
Ceriops tagal
Maximum
Mean
CV (%)
Skewness
52,55
60,00
35,00
132,73
36,62
45,00
30,00
113,10
22,61
25,00
10,00
9,62
31,85
15,00
11,00
23,76
26,72
34,45
16,49
34,03
20,91
25,40
16,55
23,17
8,98
8,51
3,08
2,89
10,81
6,76
4,16
4,72
45,07
39,53
44,28
90,21
34,62
29,28
35,31
116,94
51,74
58,37
49,21
81,58
66,42
35,40
48,87
112,69
-0,03
-0,41
0,16
1,60
0,14
-0,21
0,13
1,74
1,32
1,53
0,89
1,31
1,19
1,32
1,19
2,43
Note: Stem diameter at breast height DBH, tree height H, crown diameter CD and crown projection area CPA.
between the midle and near the seashore sites was
estimated nearly 70% meaning high similarity degree.
Figure 1 represents the relationsip between mean stem
diameter DBH and tree density ρ of mangrove from ten
samples stands. The ρ decreased as increasing in DBH.
The relationship between mean DBH and ρ was well
approximated
by
D BH = Kρ
thinning for this mangrove stands was lower as compared
with the slope of monospecific Kandelia obovata stands
(Analuddin et al. 2009), and Bruguiera gymnorrhiza
stands (Rashila et al. 2012)
Size distribution of four dominant mangrove
−α
equation,
where
coefficients K and α were calculated as 1,68 cm2 m-2α
and 1.24, respectively (R2 = 0.88). the slope of self-
Size structures of four dominant mangrove, such as stem
diameter DBH, tree height H and crown area (Table 3)
were varied among species. The Rhizophora apiculata
218 Int. Res. J. Plant Sci.
Mean stem diameter DBH (cm)
100
10
1000
10
4
-1
Tree density ρ (Ha )
Figure 1. Relationship between mean stem diameter and tree density of mangrove
stands at the Rawa Aopa Watumohai National Park using
where coefficients K and
(R2 = 0.88).
D BH = Kρ − α equation,
α were calculated as 1,68 cm2 m-2α and 1.24, respectively
was the larger and the taller mangrove as compared to
other dominant mangrove species, while the Lumnitzera
littorea was the smaller and the shorter mangrove
species. These trends realized that the R. apiculata was
grown larger in the at the present study site as compared
with other mangrove species. On the other hand, size
inequality or coefficient of variation (CV) of DBH for
Rhizophora mucronata and Ceriops tagal was lower than
other mangrove species, indicating that these two
mangrove species might grow in the same rate, which
resulted in small variation of mangrove tree size.
The size of crown diameter (CD) and crown projection
area (CPA) of four dominant mangrove (Table 3) were
varied among mangrove species. The size of CD and
CPA for Lumnitzera littorea and Ceriops tagal were
smaller as compared the CD and CPA of R. apiculata
and R. mucronata. This means that CD and CPA of R.
apiculata and R. mucronata grow larger, while CD and
CPA of L. littorea and C. tagal tended to be small.
The size distribution of stem diameter at the breast
height DBH (Fig. 2) and H (Fig. 3) of four dominant
mangrove species showed different trends. Histogram
frequency distributuion of DBH for R. apiculata and R.
mucronata showed normal distribution (Figure 2), while
distribution DBH for Lumnitzera littorea and Ceriop tagal
showed L-shaped distribution. In addition, as shown in
Figure 3 that histogram frequency distributuion of H for R.
apiculata and R. mucronata seems to be normal
distrbution, while it was L-shaped distribution for L.
littorea and C. tagal. These trends stand for that the
mangrove species at the present study grow at the
different ways.
DISCUSSION
The present study clearly showed the spatial trends in the
structural characteristics of mangrove forest at the Rawa
Aopa Watumohai National Park. As shown in Table 1 that
there was different mangrove dominant from the
seashore to the near the land sites. The Ceriops tagal
and Lumnitzera littorea were dominant near the land site,
while the Rhizophora apiculata and Rhizophora
mucronata were dominant in the midle and near the
Analuddin et al. 219
80
80
Rhizophora mucronata
70
70
60
60
50
50
Frequency
Frequency
Rhizophora apiculata
40
30
40
30
20
20
10
10
0
0
0
10
20
30
40
50
60
10
20
30
Interval DBH (cm)
40
50
60
Interval DBH (cm)
80
80
Ceriops tagal
70
Lumnitzera littorea
70
60
Frequency
Frequency
60
50
40
30
40
30
20
20
10
10
0
50
0
10
20
30
40
Interval DBH (cm)
50
60
0
10
20
30
40
50
60
Interval DBH (cm)
Figure 2. Histogram frequency distribution of stem diameter four dominant mangrove species at the Rawa
Aopa Watumohai National Park
seashore sites. However, some mangrove species such
as Rhizopora stylosa, Bruguiera gymnorrhiza and
Avicenia alba were dstributed sparse, which showed not
clear trend. These differences in the spatially might be
due to the different in the environmental condition from
near the land to the seashore sites. The sediment
condition of near the land slightly hard, which is preffered
habitat by the mangroves of C. tagal and L. Littorea. On
the other hand, the sediment condition in the midle as
well as seashore site was mudy, which became habitat
for R. apiculata and R. mucronata. However, the new
sedimantetion area at the seashore site was the most
suitable habitat for Avicennia alba, which was just found
in the seashore site of the present study.
The spatial distribution of mangrove vegetation at the
Rawa Aopa Watumohai National Park seems to realize
the zonation, because the mangrove dominant was
different from the seashore to the near the land sites,
while many paired stands showed zero similarity index
(SI). It was just six paired stands was only having SI
values more than 70%, while it was nearly 30% paired
stands having SI values less than 10% meaning low
degree similarity of among mangrove stands at the study
site. The similarity index of mangrove among site was
found less than 50% between near the land and other
sites, which meansg low degree similarity between
mangrove near the land to the other sites, while SI
between mangrove in the midle and near the seashore
sites was estimated nearly 70% meaning high similarity
degree of mangrove for both sites.
The self-thinning process was occurred at the
mangrove forest of RAW National Park, although the
slope of self-thinning line of 1.24 was lower as compared
with the slope of self-thinning at the crowded mangrove
stands in Okinawa Island, Japan (Analuddin et al.
2009; Rashila et al. 2012), which the shelf thinning
220 Int. Res. J. Plant Sci.
120
120
Rhizophora mucronata
100
100
80
80
Frequency
Frequency
Rhizophora apiculata
60
40
40
20
0
60
20
0
10
20
30
40
50
60
00
70
10
20
30
40
50
60
70
Interval tree height (m)
Interval tree height (m)
120
Ceriops tagal
120
Lumnitzera littorea
100
100
80
Frequency
Frequency
80
60
40
40
20
20
0
60
0
0
10
20
30
40
50
60
Interval tree height (m)
0
10
20
30
40
50
60
Interval height (m)
Figure 3. Histogram frequency distribution of tree height four dominant mangrove species at the Rawa
Aopa Watumohai National Park
process of mangrove at these sites was studied on the
longterm of period time. The slope of self-thinning line
(Figure 1) at the present study was very close to -4/3
power law of self-thinning (Enquist et al. 1998). However,
the self-thinning process of many crowded stands follows
the -3/2 power law of self-thinning when the self-thinning
process is studied for the long time priode by following
the trajectory of mean mass and density of crowded
stands. Analuddin et al. (2009) showed that the slope of
self-thinning for crowded Kandelia obovata stands was
very close to -3/2 power law of self-thinning of Yoda’s
Lawa. In addition, Rashila et al. (2012) was also found
the slope of self-thinning for the Bruguierra gymnorrhiza
stands was close to Yoda’s law. However, whatever the
self-thinning slope of 3/2 (Yoda et al., 1963) or 4/3
(Enquist et al. 1998), the self-thinning process of an
organisms follows a self-thinning rule (Dewar and
Magnani, 1999).
The size inequality or coefficient of variation (CV) for
stem diameter DBH and tree height H of mangroves at
Rawa Aopa Park showed different trends, i.e. the CV of
DBH and H for Rhizophora mucronata was the lowest,
while the highest CV of DBH and H was found for
Ceriops tagal. The self-thinning reduces the size
variability of terresterial trees (e.g. Hara 1986; Westoby
and Howell 1986) as well as mangroves (Analuddin et al.
2009).
Skewness of DBH and H for mangroves Rhizophora
apiculata and Rhizophora mucronata at present study
was almost negative and very close to zero, while
skewnesses of DBH and H for mangroves Cerops tagal
and Lumnitzera littorea was positive. These trends
realized the diferences of growth and development of
each mangrove species. Kohyama et al. (1990) pointed
out that skewness of tree height and stem diameter
increased with stand age until the stand reached the full
density state and then decreased with age during the
self-thinning process. Many studies testified that
skewness of stem diameter changed as stand grew, and
was correlated with high mortality rate of smaller trees
Analuddin et al. 221
(e.g. Xue and Hagihara 1999; Mcarthy and Weetman
2007), and with increasing stand age (Coomes and Allen
2007). However, other studies found that the skewnesses
of DBH and H was almost constant, though the mortality
of smaller trees was high (e.g. Hara 1985; Knox et al.
1989; Analuddin et al. 2009).
CONCLUSION
The present study clearly showed the spatial trend in the
structure of mangrove forest grown at the Rawa Aopa
Watumohai National Park, Southeast Sulawesi
Indonesia. The mangrove seems to realize the zonation
from the seashore to the landward sites. The mangroves
showed variation in spatial distribution and size structural,
which are useful information for the management and
sustanable conservation of the Rawa Aopa Watumohai
National Park.
ACKNOWLEDGMENT
We thank the Ministry of Education and Culture of
Indonesia Government, and Haluoleo University for the
financial support with grant no. 0019/ES.2/PL/2013. We
also thank Official of Rawa Aopa Watumohai National
Park for co-operation with and support for our research
working in the field.
REFERENCES
Analuddin K, Suwa R, Hagihara A, (2009). The self-thinning process in
mangrove Kandelia obovata stands. J Plant Res 122: 53-59
Analuddin K, Shahadev S, Suwa R, Hagihara A (2009). Crown foliage
dynamics of mangrove Kandelia obovata in Manko Wetland, Okinawa
Island, Japan. J Oceanogr 65: 121-127
Analuddin K (2009). Self-thinning and its concomitant changes in the
growth parameters and productivity of crowded mangrove Kandelia
obovata stands. PhD Disertation. Graduate school of Engineering
and Science, Faculty of Science, Univesity of the Ryukyus Japan.
Anonim (1997). Informasi kawasan Konservasi Sulawesi Tenggara.
Departemen Kehutanan Kantor Wilayah Propinsi Sultra Sub Balai
KSDA
Coomes DA, Allen RB (2007). Mortality and tree-size distributions in
natural mixed-age forests. J. Ecol. 95: 27-40
Dewar RC, Magnani F (1999). Plant energetics and population density.
Nature 398: 572.
Enquist BJ, Brown JH, West GB (1998). Allometric scaling of plant
energetics and population density. Nature 395: 163-165.
Gadow KV (1999). Modeling forest development. Kluwer academic
Publishers. 213 pp
Gill S, Biging G, Murphy E (2000) Modeling conifer tree crown radius
and estimating canopy cover. For Ecol Manage 126: 405-416
Givnish TJ (1986). On the economy of plant form and function.
Cambrigde University Press, Cambrigde. 717 pp
Hara T (1985). A model for mortality in a self-thinning plant population.
Ann Bot 55: 667-674
Hara T (1986). Effects of density and extinction coefficient on size
variability in plant populations. Ann Bot 57: 885-892
Hossain M, Othman S, Bujang JS, Kusnan M (2008). Net primary
production of Bruguierra parfiflora (Wight & Arn.). For Ecol Manage
255: 179-182
Husch B, Beer Th, Kershwa J (2003). Forest mensuration. J. Wiley,
Th
New York. USA 4 Edition. 443 pp
King DA (1990). Allometry of sapling and understory trees of
Panamanian forest. Funct Ecol 4: 27-32.
Knox RG, Peet RK, Christensen NL (1989). Population dynamics in
loblolly pine stands: changes in skewness and size inequality.
Ecology 70: 1153-1166
Kohyama T, Hotta M (1990). Significance of allometry in tropical
saplings. Funct Ecol 4: 515-521
Kohyama T, Hara T, Tadaki Y (1990) Patterns of trunk diameter, tree
height and crown depth in crowded Abies stands. Ann Bot 65: 567574
Kozlowski T, Kramer P, Pallardy S (1991). The physiological ecology of
woody plants. Academic Press, New York, USA. 443 pp
McCarthy JW, Weetman G (2007). Self-thinning dynamics in a balsam
fir (Abies balsamea (L.) Mill.) insect-mediated boreal forest
chronosequence. For Ecol Manage 241: 295-309
Osada N, Takeda H (2003) Branch architecture light interception and
crown development in saplings of a plagiatropically branching tropical
tree, Polyalthia jenkinsii (Anonnaceae). Ann Bot 91: 55-63
Pearcy RW, Yang W (1996) A three-dimensional shoot architecture
model for assessment of light capture and carbon gain by understory
plants. Oecologia 108: 1-12
Rashila D, Sharma S, Kamara M, Wu M, Hoque ATMR, Hagihara A
(2012) Self-thinning exponents for partial organs in overcrowded
mangrove Bruguiera gymnorrhiza stands on Okinawa Island, Japan.
For Ecol Manage 2012; 278:146-154.
Shimano K (1997). Analysis of the relationship between DBH and crown
projection area using a new model. J. Sci. 2: 237-242
Sherman RE, Fahey TJ, Martinez P (2003). Spatial patterns of biomass
and aboveground net primary productivity in a mangrove ecosystem
in the Dominican Republic. Ecosystem 6: 384-398
Smith III TJ (1992). Forest structure. In: Robertson A.I. and Alongi D.M.,
editors. Tropical mangrove ecosystems. Coastal and Estuarine
Studies. Washington (DC): American Geophysical Union. Pp. 101136.
Weiskittel AR, Maguire DA, Monserud RA (2007) Modeling crown
structural responses to competition vegetation control, thinning,
fertilization, and Swiss needle cast in coastal Douglas-fir of the
Pacific Northwest, USA. For Ecol Manage 245: 96-109
Westoby M, Howell J (1986) Influence of population structure on selfthinning of plant populations. J Ecol 74: 343-359
Xue L, Hagihara A (1999) Density effect, self-thinning and size
distribution in Pinus densiflora Sieb. et Zucc. stands. Ecol Res 14:
49-58
Yoda K, Kira T, Ogawa H, Hozumi K (1963). Self-thinning in
overcrowded pure stands under cultivated and natural conditions
(Intraspecific competition among higher plants XI). J Biol, Osaka City
Univ 14: 107-129
How to cite this article: Analuddin K, Jamili, Raya R, Septiana A and
Rahim S (2013). The Spatial Trends in the Structural Characteristics
of Mangrove Forest at the Rawa Aopa Watumohai National Park,
Southeast Sulawesi, Indonesia. Int. Res. J. Plant Sci. 4(8):214-221