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Dynamics of Decomposition and Nutrient Release of Leaf Litter in Kandelia Obovata Mangrove Forests With Different Ages in Jiulongjiang Estuary, China
Dynamics of Decomposition and Nutrient Release of Leaf Litter in Kandelia Obovata Mangrove Forests With Different Ages in Jiulongjiang Estuary, China
Ecological Engineering
journal homepage: www.elsevier.com/locate/ecoleng
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 The importance of mangrove litter fall and its decomposition in the
maintenance of debris-based food webs in the coastal waters and
Mangrove forests are extremely open and productive ecosys- their signicance for coastal sheries were indicated (Golley et al.,
tems providing large amounts of energy and organic matter to 1962; Odum et al., 1975; Lee, 1995; Ashton et al., 1999).
estuarine and coastal systems via litter fall and decomposition Despite the value and importance of mangrove forests, these
(Lugo and Snedaker, 1974; Mackey and Smail, 1996; Ye et al., 2011). ecosystems have been under severe pressure from rapidly
Litter fall has been estimated to account for 3060% of total increasing human population, large scale deforestation practices
primary production of mangrove forests (Bunt et al., 1979). The and conversion of forests into aquaculture farms (Alongi 2002; Ye
high productivity was often attributed to rapid litter decomposi- et al., 2011; Gross et al., 2014). In order to compensate the losses of
tion and efcient recycling of nutrient elements in mangrove mangrove forests and to enhance biological functions of coastal
forests (Bosire et al., 2005). Most of mangrove primary production, zones, vegetation restoration projects have been carrying out in
mainly consisting of mangrove leaves, becomes available to Jiulongjiang Estuary, China in the past decades (Chen et al., 2007;
consumers after senescence and breakdown. Decomposition of Chen and Ye, 2011), but to achieve the ultimate goal of ecosystem
litter fall is one of the basic functions of forest ecosystems (Harley, function restoration there still has a long way to go and the changes
1971; Ananda et al., 2008). The rate and extent of in situ in litter decomposition with forest restoration stages should be
decomposition governs how much of organic matter and nutrients fully understood.
is recycled within the mangrove forest, and how much is exported Decomposition of mangrove leaf litter has been studied in
to near-shore waters (Boulton and Boon, 1991 Zhou et al., 2010). 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
* Corresponding author. Tel.: +86 592 2880249; fax: +86 592 2880249. rates result in nutrient retention, while slow decomposition rates
E-mail address: yeyong@xmu.edu.cn (Y. Ye). 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 455
Fig. 1. Maps of (A) Caoputou, the study area, and (B) positions of the four studied 2.2. Leaf litter decomposition experiments
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.
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,
1999 Zhou et al., 2010). Most published information is on genera respectively. In each season, newly fallen leaves with yellow color
such as Rhizophora and Avicennia, but data on species such as were randomly picked from each forest. The leaves were washed
Kandelia obovata commonly found in subtropical regions are scanty clean and air-dried for 12 h so that no surface water remained and
(Tam et al., 1998), and studies associated with the progress of then about every 20 g leaves were placed into each nylon litter bag
vegetation restoration are even rare. Therefore, the present study (18 25 cm2) with mesh size of 1 mm. The litter bags were securely
focused on the question how decomposition and nutrient tied to aerial roots of K. obovata trees so that they were kept to lay
dynamics of leaf litter of K. obovata, a species widely distributed on the mud surface during the experiments. Totally 63 bags were
and commonly used in mangrove restoration in China, differ prepared for each forest, with 21 bags (7 bags a group xed on
among mangrove forests with forest ages. 3 random position) at each tide stripe. Three litter bags were
sampled at each tide stripe (one at each position) of each following
2. Study area and methods 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
2.1. Study area and 56 in winter. These samples were brought to the laboratory
and gently washed in a sieve to remove sediment. The washed leaf
From March 2010 to February 2011, eld experiments were litter was immediately dried at 60 C for 48 h, the nal dry weight
carried out along the southern coastline of Jiulongjiang Estuary was recorded, then ground in a mill and passed through the
near Caoputou Village (24 24 0 N, 117 55 0 E), Fugong Town, Longhai 100-mesh sieve. The powders were stored under desiccant
County, Zhangzhou City, Fujian Province of China (Fig. 1A). The conditions for chemical analyses.
climate of this region is a southern subtropical maritime (Table 1),
with annual mean air temperature of 21.0 C. In this region, winter 2.3. Chemical analyses
lasts from December to February, and the low air temperatures
usually appear in December or January. Spring months (from Subsamples of leaf litter were digested with sulfuric acid and
March to May) are wet season with relatively long rain periods and hydrogen peroxide. N contents were determined by the micro
most of rainfall during hot seasons (summer from June to August or Kjeldahl method and P contents were assayed by the ascorbic acid-
autumn from September to November) is derived from typhoons. antimony reducing phosphate colorimetric method. Organic C
To protect the sea bank in this area, several mangrove contents were analyzed by the method from Walkley and Black
plantations (mainly the species K. obovata) have been successfully (1934). The initial dry weight values of leaf litter were converted by
carried out since 1960s so that there are planted K. obovata the moisture contents and fresh weight of leaf litter in the day
mangrove forests with different ages. In this study, the selected 0 bags. All of the chemical analyses were done in laboratory at
three K. obovata forests were planted in 1962 (K48), 1986 (K24) and about 20 C. Three samples were tested for each sampling event at
1998 (K12) with ages of 48, 24 and 12 years (up to the year 2010), each stripe.
Table 1
Monthly mean climate parameters from December 2009 to November 2010.
Parameter Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov
Temperature ( C) 15.6 15.1 16.2 18.1 19.6 24.1 26.5 30.1 29.4 27.2 23.8 19.7
Rainfall(mm) 29.7 41.5 110.3 18.2 78.1 268.9 119.3 63.4 8.2 157.8 192.8 14.5
Data from the local weather bureau of Longhai County about 17 km from the study area.
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
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
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.
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).
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 soluble sugars (Steinke et al., 1993). A slower loss of dry weight
releases of C, N and P during decomposition of leaf litter, with reected the loss of more resistant materials (Steinke et al., 1990).
larger differences in N releases than those in C and P releases. The decomposition pattern in the present study was similar to
Generally, T50 values of nutrient releases increased with forest age, those observed in previous studies (Twilley et al., 1986; Mackey
indicating that the longer the forest grows, the slower the nutrients and Smail, 1996; Wafar et al., 1997; Imgraben and Dittmann, 2008).
T. Li, Y. Ye / Ecological Engineering 73 (2014) 454460 459
% 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 Melillo, J.M., Aber, J.D., Muratore, J.F., 1982. Nitrogen and lignin control of hardwood
for their dedicating work in the eld and/or laboratory. leaf litter decomposition dynamics. Ecology 63, 621626.
Mlinge, P., Atta, N., Tsuchiya, M., 2002. Nutrient dynamics and leaf litter
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