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Faculty of Applied Biological Sciences, Gifu University, 1-1 Yanagido, Gifu 501-1193, Japan
b
11-D Jones Road, 10250 Penang, Malaysia
c
Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand
Received 27 February 2007; received in revised form 24 July 2007; accepted 4 December 2007
Available online 8 December 2007
Abstract
We review 72 published articles to elucidate characteristics of biomass allocation and productivity of mangrove forests and also introduce
recent progress on the study of mangrove allometry to solve the site- and species-specific problems. This includes the testing of a common
allometric equation, which may be applicable to mangroves worldwide. The biomass of mangrove forests varies with age, dominant species, and
locality. In primary mangrove forests, the above-ground biomass tends to be relatively low near the sea and increases inland. On a global scale,
mangrove forests in the tropics have much higher above-ground biomass than those in temperate areas. Mangroves often accumulate large amounts
of biomass in their roots, and the above-ground biomass to below-ground biomass ratio of mangrove forests is significantly low compared to that of
upland forests (ANCOVA, P < 0.01). Several studies have reported on the growth increment of biomass and litter production in mangrove forests.
We introduce some recent studies using the so-called summation method and investigate the trends in net primary production (NPP). For crown
heights below 10 m, the above-ground NPP of mangrove forests is significantly higher (ANOVA, P < 0.01) than in those of tropical upland forests.
The above-ground litter production is generally high in mangrove forests. Moreover, in many mangrove forests, the rate of soil respiration is low,
possibly because of anaerobic soil conditions. These trends in biomass allocation, NPP, and soil respiration will result in high net ecosystem
production, making mangrove forests highly efficient carbon sinks in the tropics.
# 2007 Elsevier B.V. All rights reserved.
Keywords: Common allometric equation; Growth; Litter; Low T/R ratio; Net ecosystem production; Net primary production
Contents
1.
2.
3.
4.
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . .
Recent progress of studies on mangrove allometry.
Trends in mangrove biomass . . . . . . . . . . . . . . . .
Trends in mangrove productivity . . . . . . . . . . . . .
Acknowledgement . . . . . . . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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1. Introduction
Forests form a major component of the carbon reserves in the
worlds ecosystems (Whittaker and Likens, 1975) and greatly
influence the lives of other organisms as well as human societies.
Tree biomass provides various benefits, including safe habitats,
food, and timber (Nagelkerken et al., 2008; Walters et al., 2008).
* Corresponding author. Tel.: +81 58293 2860; fax: +81 58293 2860.
E-mail address: komiyama@gifu-u.ac.jp (A. Komiyama).
0304-3770/$ see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.aquabot.2007.12.006
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128
129
131
134
136
136
129
130
Table 1
Allometric equations for various mangroves based on DBH (cm)
Above-ground tree weight (Wtop in kg)
Avicennia germinans
Wtop = 0.140DBH2.40 r2 = 0.97, n = 45, Dmax = 4 cm, Fromard et al. (1998) a
Wtop = 0.0942DBH2.54 r2 = 0.99, n = 21, Dmax: unknown, Imbert and Rollet (1989) a
A. marina
Wtop = 0.308DBH2.11 r2 = 0.97, n = 22, Dmax = 35 cm, Comley and McGuinness (2005)
Laguncularia racemosa
Wtop = 0.102DBH2.50 r2 = 0.97, n = 70, Dmax = 10 cm, Fromard et al. (1998) a
Wtop = 0.209DBH2.24 r2 = 0.99, n = 17, Dmax: unknown, Imbert and Rollet (1989) a
Rhizophora apiculata
Wtop = 0.235DBH2.42 r2 = 0.98, n = 57, Dmax = 28 cm, Ong et al. (2004)
Rhizophora mangle
Wtop = 0.178DBH2.47 r2 = 0.98, n = 17, Dmax: unknown, Imbert and Rollet (1989) a
Rhizophora spp.
Wtop = 0.128DBH2.60 r2 = 0.92, n = 9, Dmax = 32 cm, Fromard et al. (1998) a
Wtop = 0.105DBH2.68 r2 = 0.99, n = 23, Dmax = 25 cm, Clough and Scott (1989)a
Bruguiera gymnorrhiza
Wtop = 0.186DBH 2.31 r2 = 0.99, n = 17, Dmax = 25 cm, Clough and Scott (1989)a
Bruguiera parviflora
Wtop = 0.168DBH2.42 r2 = 0.99, Dmax = 25 cm, n = 16, Clough and Scott (1989)a
Ceriops australis
Wtop = 0.189DBH2.34 r2 = 0.99, n = 26, Dmax = 20 cm, Clough and Scott (1989)a
Xylocarpus grnatum
Wtop = 0.0823DBH2.59 r2 = 0.99, n = 15, Dmax = 25 cm, Clough and Scott (1989)a
Common equation
Wtop = 0.251pD2.46 r2 = 0.98, n = 104, Dmax = 49 cm, Komiyama et al. (2005)
Wtop = 0.168pDBH2.47 r2 = 0.99, n = 84, Dmax = 50 cm, Chave et al. (2005)
Avicennia marina
WR = 1.28DBH1.17 r2 = 0.80, n = 14, Dmax = 35 cm,
Comley and McGuinness (2005)
Bruguiera spp.
WR = 0.0188(D2H)0.909 r2: unknown, n = 11, Dmax = 33 cm,
Tamai et al. (1986)
c.f., H = D/(0.025D + 0.583)
Bruguiera exaristata
WR = 0.302DBH2.15 r2 = 0.88, n = 9, Dmax = 10 cm,
Comley and McGuinness (2005)
Ceriops australis
WR = 0.159DBH1.95 r2 = 0.87, n = 9, Dmax = 8 cm,
Comley and McGuinness (2005)
R. apiculata
WR = 0.00698DBH2.61 r2 = 0.99, n = 11, Dmax = 28 cm,
Ong et al. (2004)
c.f., Wstilt = 0.0209DBH2.55 r2 = 0.84, n = 41
Rhizophora stylosa
WR = 0.261DBH1.86 r2 = 0.92, n = 5, Dmax = 15 cm,
Comley and McGuinness (2005)
Rhizophora spp.
WR = 0.00974(D2H)1.05 r2: unknown, n = 16, Dmax = 40 cm,
Tamai et al. (1986)
c.f., H = D/(0.02D + 0.678)
Xylocarpus granatum
WR = 0.145DBH2.55 r2 = 0.99, n = 6, Dmax = 8 cm,
Poungparn et al. (2002)
Common equation
WR = 0.199p0.899D2.22 r2 = 0.95, n = 26, Dmax = 45 cm,
Komiyama et al. (2005)
131
132
Table 2
List of above-ground (ABG) and below-ground biomass (BGB) of worldwide mangrove forests
Location
Condition or age
Species
ABG
(t ha1)
BGB
(t ha1)
H
(m)
BA
(m2 ha1)
Reference
48480 N, 1008350 E
18100 N, 1278570 E
18100 N, 1278570 E
18100 N, 1278570 E
278240 S, 153880 E
98580 N, 988380 E
18100 N, 1278570 E
98N, 988E
98N, 988E
98N, 988E
08210 N, 1038480 E
48480 N, 1008350 E
88150 N, 798500 E
>80
Primary forest
Primary forest
Primary forest
Secondary forest
Primary forest
Primary forest
Primary forest
Primary forest
Primary forest
Concession area
>80
Fringe
460.0
436.4
406.6
356.8
341.0
298.5
299.1
281.2
281.2*
281.2*
279.0
270.0
240
180.7
110.8
196.1
121.0
272.9**
177.2
11.76
106.3**
68.1**
22.4
26.4
21.2
16.4
15.5
10.6
21.7
7.2
35.9
36.2
25.1
31.30
22.8
24.0
31.30
31.30
22.1
43.8
Indonesia (Halmahera)
India (Andaman Island)
Malaysia (Matang)
Sri Lanka
18100 N, 1278570 E
128N
48N
88150 N, 798500 E
Primary forest
Primary forest
28-year-old
Fringe
R. apiculata forest
Rhizophora forest
R. apiculata stand
Avicennia
216.8
214.0
211.8
193.0
98.8
22.5
15.0
10.3
18.7
15.7
29.7
08210 N, 1038480 E
18100 N, 1278570 E
88150 N, 798500 E
Concession area
Primary forest
Fringe
B. sexangula stand
R. stylosa forest
Mixed forest
178.8
178.2
172.0
94.0
20.1
22.3
4.3
15.2
14.0
34.3
Indonesia (Halmahera)
Thailand (Phuket Southern)
Australia
Thailand (Trat Eastern)
India (Andaman Island)
Australia
Japan (Okinawa)
Japan (Okinawa)
Thailand (Satun Southern)
Indonesia (East Sumatra)
Sri Lanka
18100 N, 1278570 E
88N, 988E
338500 S, 151890 E
128120 N, 1028330 E
128N
338500 S, 151890 E
248230 N, 1248080 E
248230 N, 1248080 E
78220 N, 1008030 E
08210 N, 1038480 E
88150 N, 798500 E
Primary forest
15-year-old
Primary forest
Secondary forest
Primary forest
Primary forest
Primary forest
Primary forest
Secondary forest
Concession area
Riverine
Sonneratia forest
R. apiculata forest
A. marina forest
Mixed forest
Bruguiera and Ceriops forest
A. marina forest
R. mucronata forest
B. gymnorrhiza forest
C. tagal forest
B. parviflora stand
Mixed forest
169.1
159.0
144.5
142.2
124.0
112.3
108.1
97.6
92.2
89.7
85.0
38.5
147.3
50.3
160.3
87.5
15.9
8.0
7.0
10.8
12.5
7.0
5.5
5.5
5.2
18.8
4.4
21.2
19.0
6.0
31.0
32.9
15.2
9.2
20.0
08210 N, 1038480 E
88150 N, 798500 E
Concession area
Island habitat
B. sexangula stand
Rhizophora
76.0
71.0
17.1
3.9
5.0
11.4
88150 N, 798500 E
88150 N, 798500 E
Secondary forest
Riverine
Mixed forest
Mixed forest
62.2
57.0
28.0
6.5
4.5
11.4
13.1
08210 N, 1038480 E
08210 N, 1038480 E
Concession area
Concession area
B. parviflora stand
R. apiculata stand
42.9
40.7
19.5
29.5
4.0
2.5
Region
133
Remark: *ABG reported by Tamai et al. (1986). **Fine-root fraction was excluded from the BGB.
268N
USA (Florida)
98N
48250 S, 398300 E
198100 N, 648400 E
58300 N, 538100 E
58230 N, 528500 E
58230 N, 528500 E
58230 N, 528500 E
298480 S, 318030 E
48520 N, 528190 E
188N, 678E
258270 N, 808200 E
48250 S, 398300 E
58230 N, 528500 E
48520 N, 528190 E
258100 N, 808450 E
Panama
Kenya
Dominican Republic
French Guiana
French Guiana
French Guiana
French Guiana
South Africa
French Guiana
Puerto-rico
USA (Florida)
Kenya
French Guiana
French Guiana
USA (Florida)
Primary forest
Primary forest
50 years
Matured riverine
Matured coastal
Senescent forest
Matured riverine
Young stage
Primary forest
Pioneer stage 1 year
Pioneer stage
R. mangle stand
7.9
26.80
24.00
24.60
18.50
17.80
20.60
13.54
12.50
13.70
12.0
23.0
19.6
19.1
6.0
7.7
7.5
4.0
3.0
2.8
3.5
1.2
306.2
64.4
279.2
249.0
233.0
188.6
180.0
143.3
122.2
94.5
71.8
62.9
56.0
40.1
35.1
31.5
12.5
33.60
48520 N, 528190 E
French Guiana
Matured coastal
Lagucularia, Avicennia,
Rhizophora
Rhizophora forest
R. mucronata forest
R. mangle, Lagucularia, A. gernim
Lagucularia, Avicennia
Lagucularia, Avicennia, Rhizophora
Rhizophora, Avicennia
Rhizophora, Avicennia
B. gymnorrhiza, A. marina
Lagucularia
R. mangle
R. mangle and A. germinas fringe
C. tagal forest
Avicennia
Lagucularia
R. mangle stand
315.0
22.7
134
Fig. 4. The summation method used to calculate the net primary production
(NPP) and the net ecosystem production (NEP) of forests. See main text for
abbreviations.
NEP includes the rate of respiration (Rh) produced by microand macro-organisms in the soil, and is calculated from the
following formula: NEP = GPP (Rm + Rh) = NPP Rh.
Using the above derivation, it becomes possible to estimate
Table 3
Net primary production (NPP) of the above-ground part and canopy height (H) of worldwide mangrove forests
Type
H (m)
=
Latitude
(8)
Y
(t ha1 yr1)
L
(t ha1 yr1)
NPP
(t ha1 yr1)
Country
References
Rhizophora
3.5
8.15
6.77
6.24
13.01
Sri Lanka
Rhizophora + Avicennia
3.5
8.15
5.62
5.52
11.14
Sri Lanka
Rhizophora
3.5
8.15
4.33
4.41
8.74
Sri Lanka
Avicennia
3.5
8.15
1.40
3.74
5.14
Sri Lanka
30
30
7
0.5
6
4
6
11
21
8.6
20
6
24
8.30
8.30
25.27
25.27
18.40
18.40
18.40
8.00
4.50
18.00
18.40
18.40
19.10
1.99
0.92
2.02
20.00
12.38
3.07
12.06
7.72
4.96
3.07
4.10
6.70
11.26
9.49
12.52
8.35
19.50
13.88
12.10
8.10
6.95
3.99
6.12
26.70
23.64
12.56
24.58
16.07
16.80
PNG
PNG
Florida
Florida
Mexico
Mexico
Mexico
Thailand
Malaysia
Puerto Rico
Mexico
Mexico
Dominica
Amarasinghe and
Balasubramaniam (1992)
Amarasinghe and
Balasubramaniam (1992)
Amarasinghe and
Balasubramaniam (1992)
Amarasinghe and
Balasubramaniam (1992)
Robertson et al. (1991)
Robertson et al. (1991)
Ross et al. (2001)
Ross et al. (2001)
Day et al. (1996)
Day et al. (1996)
Day et al. (1996)
Christensen (1978)
Ong et al. (1995)
Golley et al. (1962)
Day et al. (1987)
Day et al. (1987)
Sherman et al. (2003)
24
19.10
23.60
Dominica
7.50
5.00
22.90
17.70
Indonesia
Malaysia
Rhizophora + Bruguiera
Avicennia + Sonneratia
Rhizophora
Rhizophora
Rhizophora + Avicennia
Avicennia
Avicennia
Rhizophora
Rhizophora
Rhizophra
Avicennia
Rhizophora
Rhizophora
+ Lagunchularia
+ Avicennia
Rhizophora
+ Lagunchularia
+ Avicennia
Rhizophora
Rhizophora
7.2
31.5
The growth increment (Y) and litter production (L) were used in the summation method for NPP using 12 studies, while other eight studies used different methods
including the photosynthetic and the light absorption methods.
135
136
Euskirchen (2004), NEP estimates vary with forest age and are
high in 30- to 120-year-old stands. They showed that the mean
NEP of temperate forests was less than 4.5 t C ha1 yr1 and
may even be negative (1.9 t C ha1 yr1) in 0- to 10-year-old
stands.
These results suggest that mangrove forests are highly
efficient carbon sinks in the tropics. However, future studies
should cover detailed examination on each component of
carbon fluxes, especially the dynamics of the below-ground
fraction including fine-roots. Mangroves live under extremely
wet conditions and require a bottom-heavy tree form.
Relatively high primary production and low decomposition
processes in mangrove soils are considered to bring about
unusual carbon dynamics.
Acknowledgement
We thank to Dr. S. Katahata of Gifu University for his
critical reading of our manuscript.
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