Ecological Engineering 73 (2014) 454460

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Ecological Engineering
journal homepage: www.elsevier.com/locate/ecoleng

Dynamics of decomposition and nutrient release of leaf litter in

Kandelia obovata mangrove forests with different ages in Jiulongjiang
Estuary, China
Ting Li a,b , Yong Ye a, *
a
Key Laboratory of the Ministry of Education for Coastal and Wetland Ecosystem, College of the Environment and Ecology, Xiamen University, Xiamen,Fujian,
China
b
South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China

A R T I C L E I N F O A B S T R A C T

Article history: Rates of in-situ decomposition and nutrient (organic C, N and P) release of leaf litter were seasonally
Received 7 June 2014 compared among three planted Kandelia obovata mangrove forests (K12, K24 and K48 with forest ages of
Received in revised form 12 September 2014 12, 24 and 48 years, respectively) and one natural mature K. obovata forest (KM) in Jiulongjiang Estuary,
Accepted 29 September 2014
China. The average values of half-time (T50) of leaf litter decomposition in spring, summer, autumn and
Available online xxx
winter were 29.8, 18.7, 23.9 and 47.4d, respectively. Decomposition rates were lower in the older forests
(with T50 values of 30.1 and 31.1d averaged by all seasons in K48 and KM, respectively) than in the
Keywords:
younger ones (with T50 values 29.8 and 28.8d averaged by all seasons in K12 and K24, respectively),
Mangrove
Kandelia obovata
especially in summer and autumn. The annual mean T50 of nutrient release of leaf litter during
Leaf litter decomposition followed an order of KM > K48 > K24 > K12. During leaf litter decomposition, C releases
Decomposition were very similar to dry weight losses, while N releases were slower and P releases were much faster than
Nutrient release dry weight losses. With the development of restored mangrove forests, decomposition and nutrient
Forest age release of leaf litter became slow, which may increase the chance of leaf litter being exported into the
surrounding waters.
2014 Elsevier B.V. All rights reserved.

1. Introduction
Mangrove forests are extremely open and productive ecosys-
tems providing large amounts of energy and organic matter to
estuarine and coastal systems via litter fall and decomposition
(Lugo and Snedaker, 1974; Mackey and Smail, 1996; Ye et al., 2011).
Litter fall has been estimated to account for 3060% of total
primary production of mangrove forests (Bunt et al., 1979). The
high productivity was often attributed to rapid litter decomposi-
tion and efcient recycling of nutrient elements in mangrove
forests (Bosire et al., 2005). Most of mangrove primary production,
mainly consisting of mangrove leaves, becomes available to
consumers after senescence and breakdown. Decomposition of
litter fall is one of the basic functions of forest ecosystems (Harley,
1971; Ananda et al., 2008). The rate and extent of in situ
decomposition governs how much of organic matter and nutrients
is recycled within the mangrove forest, and how much is exported
to near-shore waters (Boulton and Boon, 1991 Zhou et al., 2010).
* Corresponding author. Tel.: +86 592 2880249; fax: +86 592 2880249.
E-mail address: yeyong@xmu.edu.cn (Y. Ye).
The importance of mangrove litter fall and its decomposition in the
maintenance of debris-based food webs in the coastal waters and
their signicance for coastal sheries were indicated (Golley et al.,
1962; Odum et al., 1975; Lee, 1995; Ashton et al., 1999).
Despite the value and importance of mangrove forests, these
ecosystems have been under severe pressure from rapidly
increasing human population, large scale deforestation practices
and conversion of forests into aquaculture farms (Alongi 2002; Ye
et al., 2011; Gross et al., 2014). In order to compensate the losses of
mangrove forests and to enhance biological functions of coastal
zones, vegetation restoration projects have been carrying out in
Jiulongjiang Estuary, China in the past decades (Chen et al., 2007;
Chen and Ye, 2011), but to achieve the ultimate goal of ecosystem
function restoration there still has a long way to go and the changes
in litter decomposition with forest restoration stages should be
fully understood.
Decomposition of mangrove leaf litter has been studied in
numerous subtropical and tropical regions (e.g. Valk and Attiwill,
1984; Woodroffe and Moss, 1984; Benner et al., 1986; Robertson,
1988; Tam et al., 1990; Ananda et al., 2008). Faster decomposition
rates result in nutrient retention, while slow decomposition rates
increases the chance of leaf litter being exported (Ashton et al.,

http://dx.doi.org/10.1016/j.ecoleng.2014.09.102
0925-8574/ 2014 Elsevier B.V. All rights reserved.
 T. Li, Y. Ye / Ecological Engineering 73 (2014) 454460
Fig. 1. Maps of (A) Caoputou, the study area, and (B) positions of the four studied
mangrove forests K48, K24 and K12 for K. obovata forests aged 48, 24 and 12 years,
and KM for one natural mature K. obovata forest.
1999 Zhou et al., 2010). Most published information is on genera
such as Rhizophora and Avicennia, but data on species such as
Kandelia obovata commonly found in subtropical regions are scanty
(Tam et al., 1998), and studies associated with the progress of
vegetation restoration are even rare. Therefore, the present study
focused on the question how decomposition and nutrient
dynamics of leaf litter of K. obovata, a species widely distributed
and commonly used in mangrove restoration in China, differ
among mangrove forests with forest ages.
2. Study area and methods
2.1. Study area
From March 2010 to February 2011, eld experiments were
carried out along the southern coastline of Jiulongjiang Estuary
near Caoputou Village (24
24 0 N, 117
55 0 E), Fugong Town, Longhai
County, Zhangzhou City, Fujian Province of China (Fig. 1A). The
climate of this region is a southern subtropical maritime (Table 1),
with annual mean air temperature of 21.0
C. In this region, winter
lasts from December to February, and the low air temperatures
usually appear in December or January. Spring months (from
March to May) are wet season with relatively long rain periods and
most of rainfall during hot seasons (summer from June to August or
autumn from September to November) is derived from typhoons.
To protect the sea bank in this area, several mangrove
plantations (mainly the species K. obovata) have been successfully
carried out since 1960s so that there are planted K. obovata
mangrove forests with different ages. In this study, the selected
three K. obovata forests were planted in 1962 (K48), 1986 (K24) and
1998 (K12) with ages of 48, 24 and 12 years (up to the year 2010),
Table 1
Monthly mean climate parameters from December 2009 to November 2010.
Parameter Dec Jan Feb Mar Apr
Temperature (
C) 15.6 15.1 16.2 18.1 19.6
Rainfall(mm) 29.7 41.5 110.3 18.2 78.1
Data from the local weather bureau of Longhai County about 17 km from the study area.
455
respectively (Fig. 1B). In addition, one natural mature K. obovata
forest (KM) was also selected as the succession endpoint of
restored mangroves (Fig. 1B). The oors of these forests are not
inundated during neap tides. Tides are semidiurnal with an
average range of 4 m and the mean salinity of open water adjacent
to the mangrove forests is about 17.1 psu (Chen et al., 2007). These
four forests were subjected to roughly equivalent tidal elevation
from eld observation and are inundated by high tides for 68 days
every month.
In order to completely represent the litter decomposition
condition in these K. obovata forests, mudat of each forest was
divided into three tide stripes, each ranging around 50 m. Sediment
and vegetation surveys for these four forests were conducted in
each tide strip of each forest in April 2010. One sediment samples
(525 cm deep) was collected in the middle zone of each stripe to
determine moisture, total organic C content, total N content and
pH value (Table 2). Density, height and basal diameter of adult trees
were recorded within one quadrat of 25 m2 in the middle zone of
each stripe (Table 3).
2.2. Leaf litter decomposition experiments
Decomposition experiments were seasonally conducted in each
of the four chosen K. obovata forests by the litterbag method (Fell
et al., 1984) in March 2010, June 2010, September 2010 and
February 2011, representing spring, summer, autumn and winter,
respectively. In each season, newly fallen leaves with yellow color
were randomly picked from each forest. The leaves were washed
clean and air-dried for 12 h so that no surface water remained and
then about every 20 g leaves were placed into each nylon litter bag
(18  25 cm2) with mesh size of 1 mm. The litter bags were securely
tied to aerial roots of K. obovata trees so that they were kept to lay
on the mud surface during the experiments. Totally 63 bags were
prepared for each forest, with 21 bags (7 bags a group xed on
3 random position) at each tide stripe. Three litter bags were
sampled at each tide stripe (one at each position) of each following
time, days 0 (not incubated on mud surface), 3, 7, 14, 21, 28 and
35 in spring, summer and autumn, and days 0, 7, 14, 21, 28, 42
and 56 in winter. These samples were brought to the laboratory
and gently washed in a sieve to remove sediment. The washed leaf
litter was immediately dried at 60
C for 48 h, the nal dry weight
was recorded, then ground in a mill and passed through the
100-mesh sieve. The powders were stored under desiccant
conditions for chemical analyses.
2.3. Chemical analyses
Subsamples of leaf litter were digested with sulfuric acid and
hydrogen peroxide. N contents were determined by the micro
Kjeldahl method and P contents were assayed by the ascorbic acid-
antimony reducing phosphate colorimetric method. Organic C
contents were analyzed by the method from Walkley and Black
(1934). The initial dry weight values of leaf litter were converted by
the moisture contents and fresh weight of leaf litter in the day
0 bags. All of the chemical analyses were done in laboratory at
about 20
C. Three samples were tested for each sampling event at
each stripe.
May Jun Jul Aug Sep Oct Nov
24.1 26.5 30.1 29.4 27.2 23.8 19.7
268.9 119.3 63.4 8.2 157.8 192.8 14.5
456 T. Li, Y. Ye / Ecological Engineering 73 (2014) 454460

Table 2 3. Results
Sediment parameters (525 cm deep) in the studied K. obovata forests (mean  SE,
n = 3). TN (%) and TOC (%) are contents of total nitrogen and total organic carbon in
sediments. 3.1. Leaf litter decomposition

Forest pH TN (%) TOC (%)

There were signicantly interactive effects of forest age, season
K12 7.05  0.48 0.080  0.011 1.59  0.08 and decomposition time on dry weight losses during leaf litter
K24 6.35  0.48 0.092  0.012 1.73  0.14
decomposition (Table 4), and the highly signicant effect of
K48 6.16  0.34 0.101  0.023 1.92  0.21
KM 5.50  0.74 0.088  0.025 1.89  0.50
decomposition time suggests that dry weight loss changed
signicantly in leaf litter over the incubation period. Decomposi-
tion rates varied not only within the incubation period, but also
among forests and seasons. Statistical signicance of forest by
Table 3
time and season by time interaction suggest that rate and pattern
Characteristics of plant communities of the studied K. obovata forests (n = 3).
of decomposition of K. obovata leaf litter were signicantly
Forest Stem diameter (cm) Stem height (m) Tree density (ind hm2) different at harvest times over the total incubation period, when
K12 5.4  0.2 4.5  0.2 10978  1487 the younger forests compared with the older forests or the hotter
K24 6.7  0.1 5.9  0.4 15387  12 seasons compared with the colder seasons.
K48 9.7  0.4 7.1  0.7 10233  318
Leaf litter of K. obovata in each forest degraded rapidly in the
KM 12.0  1.8 7.0  0.4 9733  400
rst 3 weeks, with about 30% of litter dry weight loss from the litter
bag (Fig. 2). The decomposition rate of leaf litter reached its
maximum within two weeks. At the end of the incubation
(57 weeks), leaf materials of K. obovata were difcult to be
identied and the residues were intermingled with sediment
inside the bags, so the eld decomposition experiment is hard to
2.4. Statistical analysis
proceed any longer with this litter bag method. The decomposition
patterns observed in different seasons and different forests were
The relationship between the percentage of initial dry weight
similar, with very rapid early losses.
remaining in litter bags and sampling time was best tted by the
Such decomposition patterns were better described by simple
negative single exponential model (with correction factor):
negative exponential 2 Equations than simple linear regression
X t aekt equations as R (coefcient of determination) values of the
exponential regression were higher than the linear regression
where Xt is the percentage of the initial material remaining after (Tam et al., 1998), so exponential equations were used to create
decomposition time (t), a is the correction factor and k is a
tting curves of leaf litter decomposition (Table 5). Despite the
decomposition coefcient. The times required for the decomposi- similarity in litter decomposition pattern among the four K.
tion of the half the initial material (T50) were determined by the
obovata forests, mean half-time of leaf decomposition (T50) values
equation T50 = ln (a/50)/k. Differences in litter decomposition rates
were signicantly different among the four forests in each season
and changes in nutrients (organic C, N, P) among the four forests
except for winter (p < 0.001, F = 20.659 in spring; p = 0.005,
(K12, K24, K48 and KM) and four seasons (spring, summer, autumn
F = 9.802 in summer; p = 0.002, F = 13.699 in autumn; p > 0.05 in
and winter) with decomposition time series of 1, 2, 3, 4 and
winter according to one-way ANOVA). Mean T50 was 29.8 d in
5 weeks were tested by repeated measures two-way analysis of
spring, 18.7d in summer, 23.9 d in autumn and 47.4 d in winter with
variance (ANOVA) with decomposition time as within-subject
an order of increasing degradation rate as summer > autumn >
factor, forest age and season as between-subject factors. One-way
spring > winter. There were higher decomposition rates and more
ANOVA test was employed to evaluate any difference in T50 among
signicant differences among seasons at the early periods of the
the four forests in each season. The same test was used to assess
experiment than the remaining periods. Decomposition rates were
any difference in litter decomposition rate, initial concentrations of
lower in the older forests (with T50 values of 30.1 and 31.1d
nutrients and nutrient release in leaf litter among the four forests.
averaged by all seasons in K48 and KM, respectively) than in the
All analyses were performed by SPSS16.0 and Excel 2003 for
younger ones (with T50 values 29.8 and 28.8d averaged by all
Windows.
seasons in K12 and K24, respectively), especially in summer and
autumn. Averaged by all of the forests and seasons, T50 value of dry
weight loss during decomposition of leaf litter was 29.9d.

Table 4
3.2. Nutrient releases
Results of repeated measures ANOVA for dry weight loss during decomposition of
leaf litter in the studied K. obovata forests.
The nutrient (organic C, N, P) contents in K. obovata leaf litter
Source of variation df MS F p were analyzed during the decomposition processes. The initial
Within-subject values of the contents of C, N and P in leaf litter at the beginning of
Time 4 0.951 3017.056 0.000 decomposition decreased with the increasing forest age and C:N
Time  age 12 0.004 11.358 0.000
Time  season 12 0.021 66.027 0.000
ratios of younger forests (K12 and K24) were signicantly higher
Time  age  season 36 0.002 7.746 0.000 than those of older forests (K48 and KM), but there were not
Error 128 0.000 signicant differences in any of these parameters between K48 and
Between-subject KM (Table 6). There were signicant effects of forest age, season
Age 3 0.009 14.474 0.000
and decomposition time on nutrient releases duing leaf litter
Season 3 0.551 902.692 0.000
Age  season 9 0.006 9.876 0.000 decomposition (Tables 79). Nutrient releases of organic C, N and P
Error 32 0.001 during decomposition experiments were closely paralleled with
Within-subject factor is decomposition time; between-subject factors are forest age
dry weight losses (Figs. 35). The loss percentages of C were very
and season. Decomposition time refers to the number of weeks (1, 2, 3, 4 and 5) leaf similar to those of dry weight losses, while the loss percentages of
litter bags were in the eld. N were slightly lower and the loss percentages of P were faster than
 T. Li, Y. Ye / Ecological Engineering 73 (2014) 454460 457

80 K12 80 K24

Dry weight loss (%)

60 60

40 40

20 20

0 0
0 7 14 21 28 35 42 49 56 63 0 7 14 21 28 35 42 49 56 63

80 K48 80 KM
Dry weight loss (%)

60 60

40 40 Spring
Summer
20 20 Autumn
Winter
0 0
0 7 14 21 28 35 42 49 56 63 0 7 14 21 28 35 42 49 56 63
Time (days) Time (days )

Fig. 2. Loss percentages of initial dry weight during decomposition of leaf litter in the studied K. obovata forests (n = 3).

Table 5 Table 7
Simple negative exponential regression equations (X = aekt) on % of litter mass Results of repeated measures ANOVA for C losses during decomposition of leaf litter
remaining in litter bags (X) against time (t) during decomposition of leaf litter in the in the studied K. obovata forests.
studied K. obovata forests.
Source of variation df MS F p
Season Forest R2 k T50 (n = 3)
Within-subject
Spring K12 0.969 0.023 32.2  1.27c Time 4 0.934 4103.597 0.000
K24 0.962 0.026 26.9  0.69a Time  age 12 0.004 16.024 0.000
K48 0.938 0.023 29.9  0.41b Time  season 12 0.028 122.040 0.000
KM 0.993 0.023 30.3  0.71b Time  age  season 36 0.002 8.452 0.000
Summer K12 0.982 0.038 16.9  0.72a Error 128 0.000
K24 0.965 0.031 18.1  1.19ab Between-subject
K48 0.969 0.031 19.1  0.99bc Age 3 0.008 8.408 0.000
KM 0.986 0.032 20.7  0.51c Season 3 0.535 573.446 0.000
Autumn K12 0.914 0.025 22.0  1.33a Age  season 9 0.010 10.626 0.000
K24 0.944 0.025 23.2  0.58ab Error 32 0.001
K48 0.926 0.024 24.1  0.51b
KM 0.947 0.023 26.1  0.52c
Winter K12 0.901 0.013 47.9  1.32a
K24 0.903 0.013 46.9  2.52a
K48 0.886 0.013 47.4  2.90a
KM 0.934 0.013 47.3  2.76a Table 8
Results of repeated measures ANOVA for N losses during decomposition of leaf litter
R, coefcient of determination; k, decomposition constant; T50, half life time.
in the studied K. obovata forests.
Different letters after T50 data in the same season indicate statistically signicant
differences among forests in each season at the level of 0.05. Source of variation df MS F p
Within-subject
Time 4 1.057 557.205 0.000
Table 6
Time  age 12 0.006 3.208 0.000
Initial contents of nutrients in leaf litter at the beginning of decomposition in the
Time  season 12 0.042 22.126 0.000
studied K. obovata forests.
Time  age  season 36 0.003 1.710 0.016
Forest C (%) N (%) P (%) C:N Error 128 0.002
Between-subject
K12 45.28  0.84b 1.06  0.02c 0.122  0.003b 42.80  0.29c
Age 3 0.179 62.354 0.000
K24 44.50  0.65b 0.91  0.03b 0.109  0.003a 49.01  1.41b
Season 3 0.429 149.330 0.000
K48 43.15  0.51a 0.72  0.04a 0.104  0.003a 60.06  3.46a
Age  season 9 0.024 8.412 0.000
KM 43.29  0.17a 0.70  0.05a 0.101  0.009a 61.71  4.26a
Error 32
Different letter indicate statistically signicant differences among forests at the
level of 0.05.

those of dry weight losses. More rapid releasing of organic C, N and

P from decomposed leaves were found in summer and autumn
than in winter and spring, indicating more nutrients were released
and exported to the surrounding waters in hot seasons. For N
concentrations in the residual litter materials during decomposi-
tion (Fig. 4), there were general increases from the 2nd to the 3rd or
the 4th weeks of leaf litter decomposition in Spring, Summer and
Autumn. After these periods, the N remaining in litter materials
declined steadily. P remaining in the residual litter material rapidly
decreased during the rst 3 weeks and then decreased gradually
(Fig. 5). After 35 days decomposition, the nutrients releases from
leaf litter were around 54% for organic C, 50% for N and 66% for P in
spring, 68% for C, 62% for N and 76% for P in summer, 57% for C, 53%
for N and 68% for P in autumn, and 41% for C, 29% for N and 49% for P
in winter.
The decomposition patterns of nutrients in K. obovata leaf litter
were also described by simple negative exponential 2 Equations
458 T. Li, Y. Ye / Ecological Engineering 73 (2014) 454460

Table 9 releases during decomposition of leaf litter. Averaged by data in the

Results of repeated measures ANOVA for P losses during decomposition of leaf litter
four forests, T50 values of releases of C, N and P during
in the studied K. obovata forests.
decomposition of leaf litter were 29.4, 40.4 and 21.2d.
Source of Variation df MS F p
Within-subject
Time 4 1.016 495.825 0.000
4. Discussion
Time  age 12 0.004 1.712 0.071
Time  season 12 0.017 8.097 0.000 Mangrove litter has been reported to be quickly decomposed
Time  age  season 36 0.001 0.622 0.950 with rapid decreases in dry weight and high leaching rates in the
Error 128 0.002
rst few weeks of exposure in the eld (Valk and Attiwill, 1984;
Between-subject
Age 3 0.023 4.859 0.007 Mlinge et al., 2005). In the present study, the highest
Season 3 0.559 116.932 0.000 decomposition rate occurred in the 2nd week in winter and in
Age  season 9 0.010 2.167 0.052 the rst week in other seasons, and then followed by slower
Error 32 0.005 decreases for the rest of the experiment period. This quick
Within-subject factor is decomposition time; between-subject factors are forest age decomposition at early stage might be related to leaching of
and season. Decomposition time refers to the number of weeks (1, 2, 3, 4 and 5) leaf soluble organic materials and inorganic compounds such as
litter bags were in the eld.

110 K12 K24 110

K48 KM
90 90
Spring
% C remaining

S ummer
70 70
50 50
30 30
10 10
0 10 20 30 40 0 10 20 30 40
110 110
90 90
Autumn Winter
% C remaining

70 70
50 50
30 30
10 10
0 10 20 30 40 0 20 40 60
Time (days) Time (days)

Fig. 3. Remaining percentages of organic C during decomposition of leaf litter in the studied K. obovata forests (n = 3).

120 K12 K24 120

K48 KM
100 100
% N remaining

S pring S ummer
80 80

60 60

40 40

20 20
0 10 20 30 40 0 10 20 30 40

120 120
100 Autumn 100
Winter
% N remaining

80 80
60 60
40 40
20 20
0 10 20 30 40 0 20 40 60
Time (days) Time (days)

Fig. 4. Remaining percentages of N during decomposition of leaf litter in the studied K. obovata forests (n = 3).

(Table 10). The four forests had signicantly different T50 values of
releases of C, N and P during decomposition of leaf litter, with
larger differences in N releases than those in C and P releases.
Generally, T50 values of nutrient releases increased with forest age,
indicating that the longer the forest grows, the slower the nutrients
soluble sugars (Steinke et al., 1993). A slower loss of dry weight
reected the loss of more resistant materials (Steinke et al., 1990).
The decomposition pattern in the present study was similar to
those observed in previous studies (Twilley et al., 1986; Mackey
and Smail, 1996; Wafar et al., 1997; Imgraben and Dittmann, 2008).
 T. Li, Y. Ye / Ecological Engineering 73 (2014) 454460 459

110 K12 K24 110

K48 KM
90 90
S pring

% P remaining
70 S ummer
70

50 50

30 30

10 10
0 10 20 30 40 0 10 20 30 40

110 110
90 Autumn 90
Winter
% P remaining

70 70
50 50
30 30
10 10
0 10 20 30 40 0 20 40 60
Time (days) Time (days)

Fig. 5. Remaining percentages of P during decomposition of leaf litter in the studied K. obovata forests (n = 3).

Leaf litter decomposition can be inuenced by not only seasons the rst 3 weeks of decomposition and N quantity declined much
(Mackey and Smail, 1996) but also spatial scales such as tidal levels slower than organic C and P. This may be due to the rapid releases
(Lee, 1989; Mackey and Smail, 1996; Dick and Osunkoya, 2000; of soluble forms of P from litter by leaching (Steinke et al., 1993;
Mlinge et al., 2002). The decomposition rates in both summer and Tam et al., 1998).
winter from K. obovata forests in Jiulongjiang Estuary in China in The decomposition of leaf litter is greatly associated with the
the present study, with T50 of dry weight losses of 18.7 and 47.4d surrounding environment which can be largely changed with the
respectively, were much slower than those with corresponding development of a restored forest. At different stages of a restored
values of 13 and 42d from mangrove forests of the same species in mangrove ecosystem, many factors such as litter production,
Hong Kong (Tam et al., 1990), about 2
-latitude and 2
C annual microbiological activity, inundation condition, aerobic condition of
mean air temperature less than the area of the present study. sediment, faunal communities and litter nutritional values also
The decomposition rates of leaf litter in each of the four mangrove changed (Clough et al., 2000; Morrisey et al., 2003; Nga et al., 2005;
forests in the present study were more rapid in summer than those Chen et al., 2007; Ye et al., 2013). The present study showed that
in winter because activities of microbes increased in hot seasons, initial N and P levels in leaf litter declined as the increase in forest
as reported in other mangrove forests (Lin, 1999). age of K. obovata forests. This is in line with ndings from Nga et al.
Earlier studies indicated that initial nutrients level of litter, in (2005) that mangrove plants in younger forests (7 and 11 years) are
particular N content, has a controlling inuence on litter able to take up more N and P than those in older forests (17 and
decomposition rates (Melillo et al., 1982; Twilley et al., 1997). 24 years) and the formers produce a large quantity of higher
Decomposition rates generally increase for litter with high leaf N quality litter as input to the aquatic system. Hong and San (1993)
content (Pelegra et al., 1997). Fenchel et al. (1998) suggested that concluded that old trees produce much woody materials, such as
high initial N content or low C:N ratio in litter would result in high stems, branches and reproductive parts, while young trees produce
microbial assimilation and mineralization efciencies, and less more leaves. Clough et al. (2000) even indicated that the leaf area
immobilization effects, hence accelerate the decomposition of index (LAI) decreased with the increase in mangrove forest age,
litter fall. Certainly, we observed a steady enrichment of litter N in resulting in more closed canopy in young forests than in old ones.
Chen et al. (2007) showed that younger forests (4 and 7 years)
generally had more macro-benthic fauna species than older ones
Table 10
(19 and 43 years). In a conclusion, litter decomposition rate is
Simple negative exponential regression equations (X = aekt) on % of annual litter
nutrients remaining in litter bags (X) against time (t) during decomposition of leaf inevitably to show some differences with the development of a
litter in the studied K. obovata forests. restored mangrove forest, as is indicated by the present study that
litter decomposition rates are directly correlated to the age of the
Nutrient Forest R2 k T50 (n = 3)
mangrove forests. We found that leaf litter lose its dry weight and
C remaining K12 0.992 0.023 28.6  0.57a
nutrients (organic C, N, P) signicantly faster in younger forests
K24 0.985 0.022 28.8  1.09a
K48 0.985 0.022 29.2  0.63a (K12, K24) than in older forests (K48, KM), especially during the
KM 0.989 0.022 30.9  1.02b hotter and wetter seasons (summer and autumn). With the
N remaining K12 0.988 0.022 32.3  0.68a development of restored mangrove forests, slower decomposition
K24 0.949 0.020 36.9  0.84b and nutrient release of leaf litter may increase the chance of leaf
K48 0.903 0.014 51.1  3.24 d
KM 0.979 0.017 41.1  1.82c
litter being exported into ocean.
P remaining K12 0.979 0.032 19.4  0.74a
K24 0.962 0.028 21.3  0.43ab
K48 0.964 0.028 21.4  1.13ab
Acknowledgements
KM 0.978 0.029 22.5  1.60b
The work described in this paper is supported by the grants
R, coefcient of determination; k, decomposition constant; T50, half life time.
Different letters after T50 data for the same nutrient indicate statistically signicant from National Natural Science Foundation of China (Project no.
differences among forests at the level of 0.05.
460 T. Li, Y. Ye / Ecological Engineering 73 (2014) 454460
41076049 and 41276077). We thank Baipeng Pang and Yantao Gu
for their dedicating work in the eld and/or laboratory.
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