Forest Science and Technology

ISSN: 2158-0103 (Print) 2158-0715 (Online) Journal homepage: https://www.tandfonline.com/loi/tfst20

Allometric models and aboveground biomass of

Lumnitzera racemosa Willd. forest in Rawa Aopa
Watumohai National Park, Southeast Sulawesi,
Indonesia

Analuddin Kangkuso, Jamili Jamili, Andi Septiana, Rasas Raya, Idin Sahidin,
Usman Rianse, Saban Rahim, Alfirman Alfirman, Sahadev Sharma & Kazuo
Nadaoka

To cite this article: Analuddin Kangkuso, Jamili Jamili, Andi Septiana, Rasas Raya, Idin Sahidin,
Usman Rianse, Saban Rahim, Alfirman Alfirman, Sahadev Sharma & Kazuo Nadaoka (2016)
Allometric models and aboveground biomass of Lumnitzera racemosa Willd. forest in Rawa
Aopa Watumohai National Park, Southeast Sulawesi, Indonesia, Forest Science and Technology,
12:1, 43-50, DOI: 10.1080/21580103.2015.1034191

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Forest Science and Technology
Vol. 12, No. 1, March 2016, 43
50

Allometric models and aboveground biomass of Lumnitzera racemosa Willd. forest in Rawa
Aopa Watumohai National Park, Southeast Sulawesi, Indonesia
Analuddin Kangkuso a*, Jamili Jamilia, Andi Septianaa, Rasas Rayaa, Idin Sahidinb, Usman Riansec, Saban Rahimd,
Alfirman Alfirmand, Sahadev Sharmae and Kazuo Nadaokae
a
Faculty of Mathematics and Natural Sciences, Halu Oleo University, Kendari, Indonesia; bFaculty of Pharmacy, Halu Oleo University,
Kendari, Indonesia; cFaculty of Agriculture, Halu Oleo University, Kendari, Indonesia; dPostgraduate Program, Halu Oleo University,
Kendari, Indonesia; eGraduate School of Information Science and Engineering, Tokyo Institute of Technology, Tokyo, Japan

(Received 9 August 2014; Accepted 17 March 2015)

Information on aboveground biomass of mangrove forest at Rawa Aopa Watumohai National (RAWN) Park is
fundamental for the management of mangroves (Lumnitzera racemosa Willd.) in the park. Allometric relationships of
partial or whole L. racemosa trees were examined using independent variables D30, DBH, and DB or quadratics D302H,
DBH2H, and DB2H from different individuals of L. racemosa trees. Aboveground biomass was estimated by allometric
equations and tree census data. Results showed that the best fit for allometric models of stem biomass (WS) and leaf
biomass (WL) used the independent variable of DBH2H, while the best fit for branch biomass (WB) used quadratic DB2H.
The quadratic DBH2H is the most reliable parameter for estimating aboveground biomass of L. racemosa trees.
Aboveground biomass of L. racemosa varied among stands, ranging from 26.23 to 191.01 ton ha¡1. Biomass density of L.
racemosa forest varied from 4.07 to 29.80 ton ha¡1 m¡1. The high aboveground biomass of L. racemosa indicates its high
blue carbon stock, which maintains the productivity of nearby coastal areas. Therefore, the results of this study can help
the sustainable management of mangroves in the RAWN Park and surrounding areas.
Keywords: allometric model; aboveground biomass; Lumnitzera racemosa; RAWN Park; Indonesia

Introduction measurements (Khan et al. 2005), the overall productivity

Mangroves are woody plant communities inhabiting the
intertidal zones of tropical and subtropical coasts. Man-
grove forest is a highly productive ecosystem that exports
organic matter to support a variety of organisms and
enhances phytoplankton production (Rivera-Monroy et al.
1998). Mangroves maintain the ecological equilibrium in
coastal waters and preserve rich biological resources and
species diversity. The carbon sequestrated in vegetated
coastal ecosystems has been termed “blue carbon.”
Because of their high productivity, these ecosystems play
an important role in the global carbon cycle, and therefore
occupy a position of increasing importance in climate
change debates and mitigation programs. However, due to
increasing human population, global mangrove cover is
being destroyed, adversely affecting mangrove ecosystem
function and productivity. Therefore, estimation of
remaining mangrove biomass is needed for the develop-
ment of realistic carbon budgets in coastal ecosystems.
The UN REDDC program requires developing countries
to produce robust estimates of forest biomass accompa-
nied by an appropriate measure of uncertainty.
The carbon fixation capacity of mangrove forests is
higher than that of terrestrial forests (Clough 1998). For
this reason, mangroves have been considered an important
carbon sink in coastal ecosystems (Ong 1993). Although
many studies have been conducted on biomass
of mangrove forests remains poorly understood. Conse-
quently, biomass estimation is a key tool for studying the
production structure and functions of forest ecosystems.
Many studies have been focused on allometry and bio-
mass estimation as a first step in the evaluation of carbon
sinks in different plant ecosystems. Estimation of man-
grove biomass in the field is difficult due to extreme con-
ditions such as mud, dangerous animals and insects, and
inundation at high tide. Therefore, many ecologists have
developed allometric models for quantifying the whole or
partial biomass of mangroves from measurable tree
dimensions (Clough et al. 1997; Komiyama et al. 2005;
Dahdouh-Guebas et al. 2006; Deshar et al. 2012). Many
studies using allometric models have quantified above-
ground biomass of mangrove forest worldwide (Christen-
sen 1978; Tamai et al. 1986; Day et al. 1987; Lee 1989,
1990; Day et al. 1996; Ross et al. 2001; Sherman et al.
2003; Soares and Schaeffer-Novelli 2005; Khan et al.
2009), but few data are available for aboveground bio-
mass of mangrove forest from Indonesian sites
(Komiyama et al. 1987; Kusmana et al. 1992; Sukardjo
and Yamada 1992), and there are no aboveground bio-
mass data available for Lumnitzera racemosa Willd.
forest.
Mangrove forest at the Rawa Aopa Watumohai
National (RAWN) Park is one of the most protected

*Corresponding author. Email: zanzarafli@gmail.com

ISSN 2158-0103 print/ISSN 2158-0715 online

Ó 2015 Korean Forest Society
http://dx.doi.org/10.1080/21580103.2015.1034191
http://www.tandfonline.com
44 A. Kangkuso et al.
forests in the Southeast Sulawesi region of Indonesia.
Spatial trends in the structural characters of this mangrove
forest show clear zonation of different stands, with L.
racemosa growing near the land site (Analuddin et al.
2013). Nevertheless, it is important to establish allometric
models for estimating the aboveground biomass of man-
groves, which can improve the understanding and knowl-
edge of biomass stock and future productivity of
mangroves. These data are fundamental for the conserva-
tion and management of mangrove forests in the RAWN
Park, which support the habitat of the endangered
endemic buffalo, anoa (Bubalus sp.), and also play an
important role as a blue carbon source for the coastal zone
of this region. Therefore, the aims of this study were to:
(1) develop an allometric model for estimating whole or
partial aboveground biomass of L. racemosa; (2) elucidate
Figure 1. Maps of Indonesia (above) and of southeast Sulawesi (below). Small box is study site of Rawa Aopa Watumohai National
Park (RAWN Park 2011).
the trends in aboveground biomass, biomass density, and
biomass allocation in L. racemosa forest; and (3) estimate
the stock of aboveground biomass of mangroves in the
RAWN Park.
Materials and methods
Site description
The present study was carried out in the mangrove
forest of the RAWN Park (Figure 1), which is located
in the eastern part of Kendari city (04o33’12.100 S
122o0.3’20.400 E), Southeast Sulawesi, Indonesia (RAWN
Park 2011). It is an excellent location for studying man-
grove biomass because mangroves at this site are continu-
ously watered by the Lanowulu River. The L. racemosa
forest is located near the land site of the RAWN Park

(Analuddin et al. 2013), and shows different growth stages
and stand performance. This mangrove is monospecific
dominant on the border of the land site, and the tree can-
opy is closed due to crowded stands.
Tree sampling and allometric establishment
Forest survey and tree sampling were carried out from
February to June 2014 using a quadrat method within nine
100 m2 stands. The stem and height of all individual trees
were measured. Stem diameter at breast height (DBH)
was measured using diameter tape, while tree height (H)
was measured by measuring pole meter. An allometric
model was established by selecting eight different sized
L. racemosa trees for estimation of stem mass, branch
mass, leaf mass, and aboveground mass. All selected trees
were harvested, and divided into main stem, branch/twig,
as well as leaf components. The main stem was cut into
1 m length intervals from the bottom to the top. The fresh
weight of all tree components was measured at the sample
plot. Main stem samples of c. 5 cm discs were taken from
each 1 m length, together with branches/twigs and leaf
samples, and brought to the Biology Laboratory at Halu
Oleo University to be oven dried at 80  C. Total dry mass
of L. racemosa parts (stem, branch/twig and leaf) were
estimated as dry/fresh biomass ratios (Brown et al. 1997).
An allometric model was established using independent
variables of stem diameter at 30 cm from the ground
(D30), at breast height (DBH), and branch diameter (DB)
at minimum position or quadratic those of parts and tree
height (D302H, DBH2H and DB2H). In addition, the age
of the stands was estimated by ring analysis from a sample
tree taken from each stand on the basis of mean DBH.
Data analysis
Allometric models of stem weight (WS), branch weight
(WB), and leaf weight (WL) of L. racemosa trees were
established as WS D aDBH2Hb, WB D aDB2Hb, and
WL D aDBH2Hb, respectively; where a and b are the two
constant values. Total WS of L. racemosa trees in each
stand was estimated by summation method using the fol-
lowing equation:
X
n (Table 1).
WS D WS1 C WS2 C ::::WSn
iD1
where WS is stem weight of trees, while WS1, WS2, … ..
WSn are stem weight of individual trees.
Total WB of L. racemosa trees in each stand was esti-
mated by summation method using the following
equation:
X
n ship of DBH to WL were estimated as 1.30 £ 10¡2 and
WB D WB1 C WB2 C ::::WBn
iD1
where WB is branch weight of trees, while WB1, WB2,
… ..WBn are branch weight of individual trees.
Forest Science and Technology 45
Total WL of L. racemosa trees in each stand was esti-
mated by summation method using the following equa-
tion:
X
n
WL D WL1 C WL2 C ::::WLn
iD1
where WL is leaf weight of trees, while WL1, WL2, … ..
WLn are leaf weight of individual trees.
Total aboveground biomass (W) of L. racemosa in
each stand was estimated by summation method using the
following equation:
X X X
WD WS C WB C WL
P P
where
P WS is total stem biomass, while WB and
WL are total branch biomass and leaf biomass,
respectively.
Results and discussion
Allometric model of partial and whole aboveground
biomass
Table 1 represents the coefficients and R2 values for allo-
metric models of partial and whole aboveground mass of
L. racemosa trees. The best fit for allometric models of
stem biomass (WS) and leaf biomass (WL) used the inde-
pendent variable DBH2H. The constant values of a and b
for the allometric model of WS were estimated as 5.82 £
10¡2 and 0.9434 (R2 D 0.993), respectively; while the val-
ues of a and b for the allometric model of WL were esti-
mated as 3.92 £ 10¡3and 1.025 (R2 D 0.934),
respectively. On the other hand, the best fit for the allome-
tric model of branch biomass (WB) used the independent
variable DB2H with constant values of a and b estimated
as 8.46 £ 10¡5 and 2.25 (R2 D 0.997), respectively. These
results suggested that the independent variable DBH2H is
the most applicable for the allometric model of stem bio-
mass in L. racemosa trees, while independent variables
DB or quadratic DB2H are the most precise for estimating
branches/twigs biomass of L. racemosa trees, although
the independent variable DBH2H could be also applied
Conversely, the allometric model of leaf biomass
(WL) is more proportional using the independent varia-
bles D30 and DBH rather than D302H or DBH2H, because
the R2 values for the relationship of D30 to WL, and DBH
to WL were high (R2 D 0.955 and R2 D 0.960, respec-
tively). The constant values of a and b for the relationship
of D30 to WL were estimated as 9.85 £ 10¡3 and 2.48,
respectively, while the values of a and b for the relation-
2.44, respectively. On the other hand, the R2 values for
the relationships of D302H to WL and of DBH2H to WL
were 0.927 and 0.934, respectively. The constant values
of a and b for the relationship of D302H to WL were esti-
mated as 3.13 £ 10¡3 and 1.034, respectively, while the
46 A. Kangkuso et al.

Table 1. Coefficients and R2 values for allometric equations between independent variables and dependent variables of partial or whole
aboveground biomass in Lumnitzera racemosa trees.

Coefficient values

Independent variables Dependent variables a b R2 values

D30 Stem biomass WS 0.146 2.24 0.982

DBH 0.179 2.23 0.983
D302H 0.0498 0.9421 0.989
DBH2H
DB 0.0582 0.9434 0.993
DB2H 0.0444 2.0222 0.964
0.17869 1.1167 0.983
D30 Branch/twig biomass WB 0.00561 3.146 0.88
DBH 0.00802 3.095 0.86
D302H 0.00126 1.321 0.91
DBH2H
DB 0.00121 1.31 0.885
DB2H 0.00082 2.935 0.993
0.0000846 2.25 0.997
D30 Leaf biomass WL 0.00985 2,48 0.955
DBH 0.013 2.44 0.96
D302H 0.00313 1.034 0.927
DBH2H
DB 0.00392 1.025 0.934
DB2H 0.00269 2.23 0.78
0.00476 1.704 0.77
D30 Aboveground biomass W 0.146 2.4 0.98
DBH 0.184 2.384 0.98
D302H 0.0464 1.0086 0.99
DB
H2H 0.056 1.0065 0.99

values of a and b for the relationship of DBH2H to WL
were estimated as 3.92 £ 1032 and 1.035, respectively.
However, the R2 values for the relationships of DB to WL
and of DB2H to WL were lower at 0.78 and 0.77, respec-
tively. The constant values of a and b for the relationship
of DB to WL were estimated as 2.69 £ 10¡3 and 2.23,
respectively, while the values of a and b for the relation-
ship of DB2H to WL were estimated as 4.76 £ 10¡3 and
1.704, respectively. Nevertheless, when the parameters of
DB and DB2H were used for leaf biomass estimation, the
allometric relationship for leaf biomass was less site-
dependent for each, but it was well fitted by DBH2H as
the independent variable for leaf biomass. Therefore, the
quadratic DBH2H is the most reliable parameter for
estimating aboveground biomass of L. racemosa trees.
Many previous studies have developed allometric
models for estimation of aboveground biomass for man-
groves by using independent variable DBH (Table 2),
which applied well for various mangrove species includ-
ing Avicennia germinans and Laguncularia racemosa
(Fromard et al. 1998a), A. marina (Comley and McGuin-
ness 2005), and Rhizophora apiculata (Ong et al. 2004).
The independent variable DBH applied well for estimat-
ing aboveground biomass of Bruguiera gymnorrhiza, Cer-
iops australis, and Xylocarpus granatum (Clough and
Scott 1989). Therefore, independent variable DBH or qua-
dratic DBH2H are the best parameters for allometric mod-
els estimating the whole or partial aboveground biomass
of various mangrove species. However, the findings of
this study indicate that there is considerable variation in

Table 2. Allometric equations for aboveground biomass for various mangrove species based on DBH (cm).

Species Allometric model References

Avicennia germinans Wtop D 0.140DBH R D 0.97, n D 45

2.40 2
Fromard et al. (1998)
Avicennia marina Wtop D 0.308DBH2.11 R2 D 0.97, n D 22 Comley and McGuinness (2005)
Laguncularia racemosa Wtop D 0.102DBH2.50 R2 D 0.97, n D 70 Fromard et al. (1998)
Rhizophora apiculata Wtop D 0.235DBH2.42 R2 D 0.98, n D 57 Ong et al. (2004)
Bruguiera gymnorrhiza Wtop D 0.186DBH2.31 R2 D 0.99, n D 17 Clough and Scott (1989)
Ceriops australis Wtop D 0.189DBH2.34 R2 D 0.99, n D 26 Clough and Scott (1989)
Xylocarpus granatum Wtop D 0.0823DBH2.59 R2 D 0.99, n D 15 Clough and Scott (1989)
Lumnitzera racemosa Wtop D 0.184DBH2.384 R2 D 0,98, n D 8 Present study
Wtop D 0.056DBH2H1.0065 R2 D 0.99, n D 8
 Forest Science and Technology 47

Table 3. Biomass in the stem, branch/twig, leaf, aboveground, and biomass density of Lumnitzera racemosa stands.

Stand Density Stem biomass Branch biomass Leaf biomass Aboveground Biomass density
Stands age (yrs) (no. indiv. ha¡1) (ton ha¡1) (ton ha¡1) (ton ha¡1) biomass (ton ha¡1) (ton ha¡1 m¡1)

1 54 2000 121.60 54.31 15.10 191.01 23.34

2 46 3400 123.04 45.88 14.32 179.72 28.54
3 42 8500 135.68 37.70 15.13 188.50 29.80
4 49 2300 88.14 34.24 10,60 132.98 21.56
5 28 8000 30.89 8.26 3.42 42.58 5.80
6 24 11900 19.82 4.32 2.10 26.23 4.07
7 26 4700 52.12 11.86 5.60 69.55 10.90
8 29 4100 65.47 17.34 7.23 90.04 14.31
9 29 2900 48.73 13.22 5.41 67.36 10.73
Mean 36.33 5311 76.16 25.24 8.76 109.77 16.56

the use of independent variables in allometric equations
for estimating the biomass of aboveground components of
mangrove species. The allometric relationships described
in this article may not be appropriate in mixed or open for-
est stands, because this study was carried out under mono-
specific and closed canopy conditions. For estimating the
biomass of trees outside the size range of this investiga-
tion, care should be taken in extrapolating the present
allometric relationships. Further studies to include other
plant species could improve the goodness of fit found in
this research.
Trends in aboveground biomass
Table 3 describes the trends in stem biomass, branches/
twigs biomass, leaf biomass, and aboveground biomass,
as well as biomass density of L. racemosa stands. Stem
biomass of L. racemosa ranged from 19.82 to 135.68 ton
ha¡1 for young and mature stands, respectively, with a
mean of 76.16 ton ha¡1. Branch biomass was estimated as
54.31 ton ha¡1 in mature stands, but as just 4.32 ton ha¡1
in young stands, with a mean of 25.24 ton ha¡1. On the
other hand, leaf biomass ranged from 2.10 ton ha¡1 in
young stands up to 15.13 ton ha¡1 in mature stands, with
a mean of 8.76 ton ha¡1. However, aboveground biomass
of L. racemosa varied among stands, ranging from 26.23
ton ha¡1 in young stands (24
29 years old) up to 191.01
ton ha¡1 in mature stands (42
54 years old), with a mean
of 109.77 ton ha¡1. Biomass studies of mangroves have
been done in many parts of the world with many different
species of mangroves including Bruguiera gymnorrhiza
(Suzuki and Tagawa 1983; Komiyama et al. 1988; Deshar
et al. 2012), B. parviflora (Kusmana et al. 1992; Hossein
et al. 2008), B. sexangula (Kusmana et al. 1992), Ceriops
tagal (Komiyama et al. 1988), Kandelia obovata (Khan
et al. 2009), Rhizophora mucronata (Suzuki and Tagawa
1983), and R. apiculata (Putz and Chan 1986; Chandra
et al. 2011). Our study showed that aboveground biomass
of L. racemosa was much higher than that of several other
mangrove species, including B. parviflora and B. sexan-
gula, which grow in East Sumatra, Indonesia (Kusmana
et al. 1992), and K. obovata, which grow in Okinawa,
Japan (Khan et al. 2009). Moreover, the biomass of L.
racemosa was comparable with values recorded elsewhere
(see Table 4).
Biomass density (aboveground biomass/mean tree
height) of L. racemosa forest varied among stands
(Table 3). The biomass density of L. racemosa trees at
mature stands was high, ranging from 21.56 to 29.80 ton
ha¡1 m¡1, with a mean of 25.81 ton ha¡1 m¡1, which is
much higher than that of K. obovata (22.30 ton ha¡1 m¡1)
forest (Khan et al. 2009). On the other hand, the present
data of the biomass density of L. racemosa at young
stands was low, ranging from 4.07 to 14.31 ton ha¡1 m¡1,
with a mean of 9.16 ton ha¡1 m¡1. In most mangroves,
the biomass density appears to be higher than other types
of terrestrial forest, while the tropical zones (i.e., lower
latitudes) show lower biomass density and higher latitudes
show higher biomass density (Khan et al. 2009; Deshar
et al. 2012).
These variations in aboveground biomass may depend
on many environmental factors including stand character-
istics, species, as well as ecological circumstances and
geographical location (Komiyama et al. 2008). Previous
work (Table 4) shows that high total aboveground bio-
mass of Rhizopora apiculata growing in Matang, Malay-
sia ranges from 270 to 460 ton ha¡1 (Putz and Chan
(1986), and reached 436.4 ton ha¡1 for B. gymnorrhiza
and 356.8 ton ha¡1 for R. apiculata growing in Halma-
hera, Indonesia (Komiyama et al. 1987). In this study,
aboveground biomass of L. racemosa was much lower
when compared to the biomass of these other mangroves.
However, a comparison with the aboveground biomass
for other global mangroves worldwide (Table 4) indicates
that the aboveground biomass of L. racemosa forest in the
RAWN Park, Indonesia was much higher than that of
Kandelia candel in Hong Kong, China (Lee 1990), or of
R. mangle in Mexico (Day et al. 1996) and in Florida
(Ross 2001). Conversely, aboveground biomass of
L. racemosa was much lower than that of tropical fringe
and basin mangroves in the Dominican Republic
(Sherman et al. 2003). These differences in aboveground
biomass of mangroves occurring in different regions may
not only be due to differences in stand structure, but also
48 A. Kangkuso et al.

Table 4. List of aboveground biomass of mangrove forests in different regions of the world.

Aboveground Mean tree

Places Mangroves biomass (ton ha¡1) height (m) References

Neotropical
Dominican Republic Rhizophora mangle 233 Sherman et al. (2003)
Florida, USA R. mangle 56 Ross et al. (2001)
Mexico R. mangle 135 Day et al. (1996)
Subtropical
Japan (Okinawa) R. mucronata 108.1 5.5 Suzuki and Tagawa (1983)
Japan (Okinawa) Bruguiera gymnorrhiza 97.6 5.5 Suzuki and Tagawa (1983)
Japan (Okinawa) Kandelia obovata 80.5 3.61 Khan et al. (2009)
Hong Kong K. candel 128.6 Lee (1990)
Tropical
Malaysia (Matang) R. apiculata 270
460 Putz and Chan (1986)
Malaysia (Kuala Selangor) B. parviflora 144.47 Hossein et al. (2008)
Malaysia (Lawas) R. apiculata 116.79 12.16 Chandra et al. (2011)
Thailand (Ranong Southern) Rhizophora spp. forest 298.5 Komiyama et al. (1987)
Thailand (Satun Southern) Ceriops tagal 92.2 5.2 Komiyama et al. (2000)
Thailand (Ranong Southern) Rhizophora spp. 281.2 10.6 Komiayama et al. (1987)
Indonesia (Halmahera) B. gymnorrhiza 436.4 22.4 Tamai et al. (1986)
Indonesia (Halmahera) R. apiculata 356.8 21.2 Komiyama et al. (1988)
Indonesia (East Sumatra) B. sexangula 279.0 21.7 Komiyama et al. (1988)
Indonesia (Halmahera) R. stylosa 178.2 22.3 Kusmana et al. (1992)
Indonesia (Halmahera) Sonneratia sp 169.1 15.9 Komiyama et al. (1988)
Indonesia (East Sumatra) B. parviflora 89.7 18.8 Komiyama et al. (1988)
Indonesia (East Sumatra) B. sexangula 76.0 17.1 Kusmana et al. (1992)
Indonesia (Southeast Sulawesi) Lumnitzera racemosa (mature stands) Mean 173.15 8.2 Present study
Indonesia (Southeast Sulawesi) Lumnitzera racemosa (young stands) Mean 59.05 5.4 Present study

to differences in climatic factors. At any given latitude, well as community structure of mangroves (Lin 1999).
high values of biomass in mangroves indicate optimum The mangrove species grown as draft type in unsuitable
growing conditions such as low salinity, high fertility, and conditions or mangrove grown in sparse might be allo-
favorable climatic conditions (Saenger and Snedaker cated more aboveground biomass for branches and leaf
1993). Therefore, the present results imply that biomass rather than for stem. Our present study shows that alloca-
of mangroves might be attributed to the characteristics of tion of stem biomass in L. racemosa forest was much
mangrove stands (e.g., whether stands are crowded or higher than in R. apiculata forest (Table 6) in Lawas,
not), mangrove developmental stage (whether stands are Malaysia (Chandra et al. 2011), but it was much lower
mature or young), and habitat conditions, including salin- when compared to B. parviflora (Table 6) growing in
ity and nutrient availability. Langkawi Island, Malaysia (Norhayati and Latif 2001).
The proportion of biomass in each component of
L. racemosa trees (Table 5) varied (i.e., the proportion of Table 5. Proportion of biomass components for Lumnitzera
stem biomass in mature stands was estimated at < 70%, racemosa stands.
except in stand no. 3). On the other hand, most young
stands showed high proportions of stem biomass Proportion of biomass (%)
(> 70%), while only 20% was branch biomass and Stands Stem Branch Leaf
< 10% was leaf biomass. This means that allocation of
biomass in the stem and branches for L. racemosa forest 1 63.66 28.43 7.91
might be affected by stand age because, as shown in 2 67.30 25.39 7.31
Table 5, the branch biomass allocation of L. racemosa in 3 71.98 20.00 8.02
mature stands was much higher than in young stands. On 4 66.28 25.75 7.97
the other hand, stem biomass allocation in young stands 5 72.71 19.26 8.03
was much higher than in mature stands. However, above- 6 72.34 19.62 8.04
ground biomass allocation in mangrove forest may not 7 72.54 19.44 8.02
only depend on stand age, but also on habitat characteris- 8 75.54 16.48 7.98
tics or edaphic factors including water logging, nutrient 9 74.94 17.05 8.01
availability, and salinity (Robertson and Alongi 1992), as Mean 70.81 21.27 7.92

Table 6. Comparison of proportion of biomass for each component of mangroves in different places.
Location (countries) Species
Langkawi Island, Malaysia Bruguiera parviflora
Lawas, Malaysia Rhizophora apiculata
RAWN Park, Southeast Sulawesi, Indonesia Lumnitzera racemosa
Previous studies have shown that biomass allocation
below and aboveground varies among mangroves. For
example, nearly 50% of mangrove biomass is allocated to
the root for C. tagal growing in Satun, Thailand
(Komiyama et al. 2000) and for K. obovata growing in
Okinawa Island, Japan (Khan et al. 2009). On the other
hand, biomass allocation to the root of R. apiculata grow-
ing in Matang, Malaysia was < 25% (Gong and Ong
1990), while it was < 15% for B. parviflora growing in
Kuala Selangor, Malaysia (Hossein et al. 2008). These
trends suggest that biomass allocation in mangrove forests
is affected by various environmental factors including
stand structure, stand age, and habitat, as well as climatic
factors. Therefore, this study suggests the existence of
integrated mechanisms between the internal and external
factors that maintain the aboveground biomass of L. race-
mosa forest.
Conclusions
Our present study provides important analysis for eluci-
dating the aboveground biomass of mangroves in the
RAWN Park. This is critical information for the conserva-
tion and management activities of mangrove forest in this
national park, which is habitat for the most protected
endemic animal on Sulawesi Island, Indonesia, the anoa
(Bubalus sp). This allometric model for mangrove forest
will support further analysis on the productivity and blue
carbon production of L. racemosa as a dominant man-
grove species. The high aboveground biomass of L. race-
mosa indicates its vigor and high blue carbon production,
which helps maintain coastal region productivity. Lumnit-
zera racemosa is classified as Least Concern on the IUCN
Red List and there are currently no conservation measures
specifically for L. racemosa. The findings of this study
will therefore help inform the RAWN Park management
plan.
Acknowledgements
This research was supported by the Higher Education, Ministry
of Education and Culture, Republic of Indonesia with grant no.
0263/E5/2014. It was also partially supported by the Tokyo Insti-
tute of Technology, Japan. We would also like to thank the Rec-
tor, and Head of Research Center Halu Oleo University as well
as the Head of RAWN Park.
Disclosure statement
No potential conflict of interest was reported by the authors.
Forest Science and Technology 49
Proportion of biomass (%)
Stem Branch Leaf References
90.0 9.10 0.90 Norhayati and Latif (2001)
57.30 35.09 7.61 Chandra et al. (2011)
70.94 21.28 8.00 Present study
ORCID
Analuddin Kangkuso http://orcid.org/0000-0003-0258-9376
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