Applied Soil Ecology 34 (2006) 266–275

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Differential responses of litter decomposition to increased soil

nutrients and water between two contrasting grassland
plant species of Inner Mongolia, China
Ping Liu a,b, Jianhui Huang a, Xingguo Han a, Osbert J. Sun a,*, Zhiyong Zhou a,b
a
Laboratory of Quantitative Vegetation Ecology, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
b
Graduate University of Chinese Academy of Sciences, Beijing 100049, China
Received 1 August 2005; received in revised form 22 December 2005; accepted 28 December 2005

Abstract
Soil chemistry and physical conditions are the key factors controlling litter decomposition. We studied the effects of increased
soil nitrogen (N), phosphorus (P), and water on the decomposition rates and associated nutrient dynamics of two dominant grassland
plant species (i.e. Allium bidentatum Fisch. ex Prokh. & Ikonn.-Gal. and Stipa krylovii Roshev.) with contrasting life forms and
tissue chemistry in a typical steppe of Inner Mongolia, China. The treatments included addition of urea at the rates equivalent to 0, 8,
16, and 32 gN/m2, and additions of mixed urea and triple superphosphate at the rates equivalent to 0, 8 gN/4 gP, 16 gN/8 gP, and
32 gN/16 gP/m2, with and without water added. We found marked differences between the two species in the rates, as well as in the
responses to water and addition rates of N and P, of litter decomposition. Additions of N alone or in mixture with P stimulated the
rate of litter decomposition in both species. Adding water significantly increased the values of decay constant, k, in A. bidentatum,
but not in S. krylovii. N and P concentrations in litters of A. bidentatum and S. krylovii all increased corresponding to increases in the
rates of N or mixed N and P additions. Our results clearly indicate that the decomposition of high quality litter is more likely to be
limited by soil moisture regimes, while that of low quality litter is more sensitive to nutrient availability. Our findings suggest that
plant species with different litter qualities should be taken into consideration when we are to model the carbon cycle and nutrient
dynamics in grassland ecosystems and that A. bidentatum is expected to contribute more than S. krylovii to the carbon cycle and
nutrient dynamics of the semi-arid grasslands of Inner Mongolia.
# 2006 Elsevier B.V. All rights reserved.

Keywords: Litter decomposition; Grassland ecosystems; Allium bidentatum; Stipa krylovii

1. Introduction development and the availability of nitrogen (N),

Decomposition of plant litter plays an important role
in nutrient cycling and carbon (C) fluxes of the
terrestrial ecosystems (Swift et al., 1979; Berg and
McClaugherty, 1989; Sun et al., 2004). The rate and
process of litter decomposition greatly influence soil
* Corresponding author. Tel.: +86 10 62836510.
E-mail addresses: osbertsun@ibcas.ac.cn,
osbertsun@ns.ibcas.ac.cn (O.J. Sun).
phosphorus (P) and other nutrients to plants and soil
microorganisms (Huang et al., 1998; Liu et al., 2000).
Understanding the key factors and processes that
control the rate of litter decomposition under different
environmental conditions and in different habitats is
therefore fundamental to quantitative analysis of C and
nutrient cycling of terrestrial ecosystems.
Decomposition is primarily driven by microbial
activities and can be best predicted by environmental
factors such as temperature and precipitation, as well as

0929-1393/$ – see front matter # 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.apsoil.2005.12.009
 P. Liu et al. / Applied Soil Ecology 34 (2006) 266–275
litter quality (Meentemeyer, 1978; Berg et al., 1993;
Aerts, 1997; Taylor et al., 1989; Wang et al., 2000;
Moretto et al., 2001), but soil chemistry and physical
conditions can also influence the rate of litter
decomposition (Gijsman et al., 1997; Seneviratne
et al., 1999). As decomposer microbes require nutrients
from either litter material or surrounding soils to
maintain their life activities (Ocio et al., 1991;
Sinsabaugh et al., 1993), soil nutrient availability has
long been suggested as one of the controlling factors
affecting the rate of litter decomposition (Swift et al.,
1979). However, results of the studies concerning the
effects of increased soil N and P on the rate of litter
decomposition and nutrient dynamics to date have been
controversial. For example, some studies (Hunt et al.,
1988; Fenn, 1991; Ostertag and Hobbie, 1999) found
that increased N and P could stimulate litter decom-
position, while others found no (i.e. Prescott et al.,
1999; Dukes and Field, 2000) or depressing effects
(Söderström et al., 1983; Magill and Aber, 1998). Berg
et al. (1982) and Berg and Matzner (1997) found a
positive response to N in the initial decomposition
phase but a negative response in the later stages.
Kwabiah et al. (1999) suggested that responses of plant
litter decomposition to soil nutrients were determined
by litter quality. Such inconsistency in the relationships
between litter decomposition and soil nutrients,
therefore, calls for continued investigations of the
subject.
Water availability can influence the rates of litter
decomposition and nutrient release through its effects
on the activities of the decomposer communities
(Orchard and Cook, 1983; Berg, 1986; Clein and
Schimel, 1994). Water supply in the form of rainfall can
also affect decomposition by facilitating leaching and
breakdown of surface litter (Swift et al., 1979).
Moreover, water availability may affect litter decom-
position indirectly by altering the litter quality in terms
of lignin and nutrient concentrations of plants (Pastor
and Post, 1988; Austin and Vitousek, 2000).
Global change will inevitably alter nutrient cycling
processes in terrestrial ecosystems along with changes
in C and water fluxes. The accelerated rates of N
mineralization likely to occur with global climate
warming (Rustad et al., 2001) and increased N
deposition due to anthropogenic activities (Bobbink
et al., 1998) could result in N enrichment under many
land use and land cover types, including grassland
ecosystems. The global precipitation pattern is antici-
pated to change under the climate change scenarios,
especially in arid and semi-arid areas where summer
precipitation is expected to increase (Melillo, 1990). All
267
these changes would potentially result in changes in
litter decomposition that could affect C and nutrient
cycles, and affect how we simulate the C cycling
process in the context of global change. However, there
is a lack of regionally based, species-specific informa-
tion on the rates of litter decomposition for most of the
terrestrial ecosystems in China.
The increased intensity of land use on the grasslands
of northern China has caused severe land degradation
since the late 1970s, resulting in decreased productivity
and ecosystem stability. Much of the area is now in
various stages of degradation, where soil N, P and water
are considered to be the principal limiting factors to net
primary productivity. The recycling of nutrients through
plant litter decomposition is an essential mechanism to
maintain productivity (Tiessen et al., 1994). To examine
how increased soil nutrient availability, predominantly
N and P, and water would affect the litter decomposition
rates and nutrient dynamics of two dominant grassland
plant species with contrasting life forms and tissue
chemistry (i.e. Allium bidentatum and Stipa krylovii),
we conducted two field experiments with varying
addition rates of N, P, and water. Our objectives were to
determine: (1) how increased soil N would affect the
decomposition rates and nutrient dynamics of A.
bidentatum and S. krylovii, and (2) how simultaneous
increases in soil N and P, with or without increased
water availability, would affect the rates of decomposi-
tion of the two species. Our underlying hypotheses were
that species with contrasting life forms and tissue
chemistry would differ significantly in the rates of litter
decomposition, and that increased soil nutrient and
water would stimulate litter decomposition in the
degraded semi-arid grassland ecosystems.
2. Materials and methods
2.1. Study site
This study was conducted at a field site of the Duolun
Restoration Ecology Experimentation and Demonstra-
tion Station in Inner Mongolia, China (latitude 428020 N;
longitude 1168160 E; altitude 1344 m a.s.l.). The mean
annual, minimum, and maximum air temperatures for
the area are 1.6,
18.3, and 18.7 8C. The mean annual
precipitation is 385 mm, occurring mainly from July to
September (accounting for 67% of the total), with
Penman evaporative potential of 1748 mm. Soils of the
region are commonly referred to as chestnut type
(Calcic Kastanozems) in the Chinese classification, and
are classified as Calcic–orthic Aridisol in the US soil
taxonomy classification system (Yuan et al., 2005).
268 P. Liu et al. / Applied Soil Ecology 34 (2006) 266–275
The area has suffered from severe degradation since
1979 as a result of overgrazing. The primary vegetation
is of the typical steppe, with perennial herb A.
bidentatum and bunch grass species S. krylovii being
among the dominant species of the plant community.
2.2. Experimental design and treatments
We conducted two decomposition experiments on a
fenced site from July and October 2004, both lasting for
100 days. The first experiment contained four levels of
N addition rates (0, 8, 16, and 32 gN/m2; designated as
N0, N8, N16, and N32, respectively), and the second
experiment contained four addition rates of mixed N
and P (0, 8 gN/4 gP, 16 gN/8 gP, and 32 gN/16 gP/m2;
designated as N0P0, N8P4, N16P8, and N32P16, respec-
tively) with and without increased water availability.
Both of the experiments were deployed based on a
randomized block design with four blocks and each plot
size of 4 m  4 m. N was added as granular urea, and P
in the form of granulated triple superphosphate, which
were applied 5 days before the litterbags were deployed.
In treatments with increased water availability, water
was applied on 15 July, 10 August, 29 August, and 22
September, following major rain events in the previous
week. For each water application, 15 mm of water was
added, simulating an average increase of rainfall by
30% of the corresponding period.
2.3. Litter decomposition
In early July 2004, litters of A. bidentatum
(predominantly leaves) and S. krylovii (predominantly
culms) were collected from the study area, including
freshly fallen and senesced tissues attached to the
plants. After being air-dried to constant mass, they were
clipped into fragments of 10 cm in length, and placed
into the 10 cm  15 cm polyethylene litterbags (mesh
size 1 mm2) that each contained 3 g of air-dried litter
material. Each treatment plot contained nine litterbags
of the same species. A total of 864 litterbags were
prepared and deployed onto the treatment plots on 9
July 2004.
The remaining litter was retrieved 35, 70, and 100
days after initial litterbag deployment. For each
sampling time, three litterbags for each species from
each treatment plot were collected. In the laboratory,
extraneous matter such as other plant materials, rocks
and small animals were handpicked from the litterbags.
The retrieved litters were then oven-dried at 70 8C for
48 h, to determine the remaining dry mass. Five samples
for each litter type were oven-dried at 70 8C for 48 h at
the time of initial deployment to determine the ratio
between air-dried mass and oven-dried mass. This ratio
was used to convert the initial air-dried mass of the litter
to oven-dried mass.
2.4. Chemical analysis
After determination of the dry mass, litters of the
same plant species from the three within-plot replica-
tions were pooled for chemical analysis. Total C, N and
P concentrations were determined for the final samples
from the two litter decomposition experiments. Total C
was measured by the standard method of wet-
combustion, total N by semi-micro Kjeldahl method
and total P by molybdenum blue colorimetric method
(Bao, 1999). Five litter samples for each plant species
were also analyzed for total C, N and P to determine the
initial litter chemistry.
2.5. Data analysis
The value of decay constant, k, was determined by
fitting the following exponential function (Olson, 1963):
xt
¼ e
kt
x0
where, xt is the remaining litter mass after a given time
period t, x0 is the initial litter mass. The remaining litter
nutrients were calculated by multiplying the sample
mass by the respective nutrient concentration. Tests of
significance of the remaining mass were performed by
analysis of variance (ANOVA; three-way without water
treatment and four-way with water treatment). Data
analyses were performed using procedures of SPSS
(v.11.0). The least significant difference (LSD) was
used for comparisons of means with confidence level
of P < 0.05.
3. Results
3.1. Initial litter chemistry
Total C contents in the initial litters were similar
between A. bidentatum and S. krylovii (Table 1). Allium
bidentatum had twice the N and P concentrations, and
half the C:N and C:P ratios, of S. krylovii (Table 1).
3.2. Species difference in litter decomposition
Allium bidentatum litter decomposed faster than S.
krylovii in both experiments (Fig. 1). After 100 days of
decomposition, the remaining litter mass was 70% of
 P. Liu et al. / Applied Soil Ecology 34 (2006) 266–275 269

Table 1
Initial litter chemistry of Allium bidentatum and Stipa krylovii in the degraded typical steppe of Inner Mongolia, China
Species C (g/kg) N (g/kg) P (g/kg) C:N ratio C:P ratio
Allium bidentatum 481.9  6.0 5.0  0.3 0.42  0.02 97  6 1273  65
Stipa krylovii 486.3  7.2 2.8  0.2 0.23  0.01 174  15 2720  101
Values are means  S.E. (n = 5).

the initial in A. bidentatum, but >80% in S. krylovii. The
patterns of litter decomposition over the duration of the
experiment were well described by the exponential
model, xt/x0 = e
kt (values of r2 ranged between 0.89
and 0.99, and P < 0.05; Table 2). The values of decay
constant, k, of A. bidentatum were about twice of S.
krylovii across all the treatments (Table 2).
3.3. Effects of nutrient addition and water on litter
decomposition

In the experiment I, addition of N alone to the

substrate soils only slightly affected the rate of litter
decomposition in both A. bidentatum and S. krylovii as
shown by three-way ANOVA and changes in the k
values, with two species showing the similar patterns of
responses (Tables 2 and 3).
Four-way ANOVA showed that the litter mass
remaining was significantly (P < 0.05) affected by
retrieval time, plant species, water treatment, addition
rates of mixed N and P, and some of the interactions in the
experiment II (Table 4). Increased rates of N and P

Table 2
Decay constant (k; y
1, r2 range 0.89–0.99, P < 0.05) for litters of
Allium bidentatum and Stipa krylovii as affected by additions of N
alone (experiment I) or in mixture with P with or without adding water
(experiment II) in the degraded typical steppe of Inner Mongolia,
China.
Treatment k
Allium bidentatum Stipa krylovii
Experiment I
N0 1.20  0.07 a 0.53  0.07 b
N4 1.24  0.10 a 0.58  0.07 ab
N8 1.30  0.10 a 0.63  0.10 ab
N16 1.33  0.08 a 0.67  0.06 a
Experiment II
No water added
N0P0 1.20  0.07 c 0.63  0.06 c
N8P4 1.23  0.11 bc 0.69  0.02 bc
N16P8 1.37  0.05 ab 0.77  0.07 ab
N32P16 1.48  0.13 a 0.85  0.10 a
Water added
N0P0 1.37  0.07 b 0.63  0.01 d
N8P4 1.41  0.15 ab 0.71  0.02 c
N16P8 1.48  0.09 ab 0.79  0.04 b
N32P16 1.56  0.09 a 0.87  0.05 a
Fig. 1. Effects of N addition or concurrent additions of N and P with or
without water added on litter decomposition of Allium bidentatum (open The values of k are means  S.E. (n = 4), which were derived based on
symbols) and Stipa krylovii (closed symbols). Vertical bars represent exponential decay model, xt/x0 = e
kt. Data in the same column
standard errors (n = 4). *P < 0.05; **P < 0.01; ***P < 0.001. followed by the same letter are not significantly different (P < 0.05).
270 P. Liu et al. / Applied Soil Ecology 34 (2006) 266–275

Table 3
F and P (shown in parentheses) values from three-way ANOVA for mass remaining (M, % of initial mass) and two-way ANOVA for residual N and P
(% of initial mass) during decomposition of litter materials in soils with N addition (experiment I), with retrieval time, plant species and N addition
(fertilizer) as main effects
Source of d.f. F
variation
M N P
Time (T) 2 79.3 (<0.001)
Species (S) 1 286.5 (<0.001) 14.3 (<0.001) 65.2 (<0.001)
Fertilizer (F) 3 2.4 (0.073) 34.1 (<0.001) 7.7 (0.001)
TS 2 10.5 (<0.001)
TF 6 0.9 (0.513)
SF 3 1.4 (0.241) 3.5 (0.030) 0.01 (0.999)
TSF 6 0.9 (0.504)

additions significantly (P < 0.05) accelerated the rate of
litter decomposition in both A. bidentatum and S. krylovii;
the values of k increased with increasing rates of N and P
additions in both species regardless water treatment
(Table 2). Adding water significantly (P = 0.001) inc-
reased the k value in A. bidentatum, but not in S. krylovii
(Table 2). The decomposition rates of S. krylovii litter
were more responsive to changes in the rates of mixed N
and P additions, as significant treatment effects of N and P
additions were shown for the remaining mass litter of the
species on the Day 70 and 100 after the commencement of
litter decomposition experiment (Fig. 1), and the changes
in the values of k with increasing rates of N and P additions
were greater in S. krylovii than in A. bidentatum (Table 2).
In both species, the k value responded more to the
concurrent changes in N and P (Experiment II) than
changes in N alone (Experiment I; Table 2).
3.4. Litter nutrient dynamics
At the end of the 100-day study, the N and P
concentrations in litters of A. bidentatum and S. krylovii
all increased with increases in the addition rates of N in
the experiment I (Fig. 2), and the addition rates of mixed
N and P in the experiment II (Fig. 3). Adding water
decreased the N and P concentrations in A. bidentatum
and S. krylovii in the experiment II (Fig. 3).
Two-way and three-way ANOVA showed that the
residual amounts of litter N and P were significantly
(P < 0.01) affected by plant species and addition rates
of N in the experiment I (Table 3), and by plant species,
addition rates of mixed N and P, and water treatment in
the experiment II (Table 4).
The absolute amounts of litter N increased in nearly
all treatments after decomposing for 100 days, in both

Table 4
F and P (shown in parentheses) values from four-way ANOVA for mass remaining (M, % of initial mass) and three-way ANOVA for residual N and P
(% of initial mass) during decomposition of litter materials in soils with mixed N and P additions with or without water added (experiment II), with
retrieval time, plant species, water and N and P additions (fertilizer) as main effects
Source of variation d.f. F
M N P
Time (T) 2 266.1 (<0.001)
Species (S) 1 930.3 (<0.001) 44.5 (<0.001) 120.9 (<0.001)
Water (W) 1 4.0 (0.047) 10.8 (0.002) 20.9 (<0.001)
Fertilizer (F) 3 23.2 (<0.001) 116.5 (<0.001) 59.7 (<0.001)
TS 2 28.2 (<0.001)
TW 2 1.1 (0.343)
SW 1 5.9 (0.016) 0.1 (0.789) 0.5 (0.464)
TSW 2 2.6 (0.075)
TF 6 1.1 (0.374)
SF 3 0.6 (0.586) 6.6 (0.001) 0.3 (0.825)
TSF 6 1.7 (0.115)
WF 3 2.9 (0.039) 0.2 (0.908) 0.8 (0.496)
TWF 6 0.6 (0.740)
SWF 3 2.7 (0.047) 0.2 (0.926) 0.2 (0.889)
TSWF 6 0.4 (0.879)
 P. Liu et al. / Applied Soil Ecology 34 (2006) 266–275 271

decomposition, whereas in S. krylovii, the absolute

amounts of litter P increased (Table 5). Similar to the
patterns of litter N and P concentrations, the absolute
amounts of N and P in the remaining litter from the two
experiments all increased with increasing addition rates
of N alone or mixed N and P (Table 5).
Adding water significantly reduced the absolute
amounts of litter N and P in A. bidentatum (P = 0.004
and <0.001, respectively) and the absolute amounts of
litter P in S. krylovii (P = 0.025; Table 5).

4. Discussion

On a smaller spatial scale, litter quality is considered

Fig. 2. Effects of N addition on litter N and P concentrations of Allium
bidentatum and Stipa krylovii after decomposing for 100 days in the
field. Vertical bars represent standard errors (n = 4).
A. bidentatum and S. krylovii (Table 5). The changes in
the absolute amounts of litter P, however, differed
between the two species: in A. bidentatum, the absolute
amounts of litter P decreased slightly after 100-day
as the most important factor influencing decomposition
rate (Melillo et al., 1982; Aerts and De Caluwe, 1997).
High quality litters are often characterized by higher N
concentrations and lower C:N ratios, and can decom-
pose faster in comparison with low quality litters
(Sanchez, 2001). In this study, the litters of the perennial
herb A. bidentatum had twice the N concentration, and
about half of the C:N ratio, of the bunch grass species S.
krylovii. The marked differences in the rate of litter
decomposition between these two species can, there-
fore, attribute to differences in litter quality.
We also found that increasing the rates of N additions
resulted in slight increase in the rates of litter
decomposition in the two grass species. This is in
general agreement with many other studies showing

Table 5
Residual amounts of N and P (% of initial mass) in Allium bidentatum and Stipa krylovii after decomposing for 100 days as affected by additions of N
alone (experiment I) or in mixture with P with or without adding water (experiment II)
Treatment Allium bidentatum Stipa krylovii
N P N P
Experiment I
N0 103.1  4.0 c 86.2  2.5 c 101.4  3.2 c 102.3  4.2 b
N8 105.3  3.8 c 91.0  4.5 bc 111.2  4.3 b 106.3  7.6 ab
N16 111.7  5.3 b 95.3  2.6 ab 120.2  9.3 b 111.0  7.6 ab
N32 119.6  4.0 a 98.8  2.6 a 134.5  5.7 a 114.6  8.5 a
Experiment II
No water added
N0P0 102.3  2.0 d 86.7  3.0 d 101.8  3.2 d 100.7  4.1 c
N8P4 109.2  4.5 c 96.4  3.9 c 114.4  5.9 c 110.8  6.0 b
N16P8 116.0  5.2 b 105.1  5.0 b 125.7  4.2 b 117.3  8.5 ab
N32P16 122.3  3.4 a 113.3  3.7 a 137.3  3.3 a 122.5  5.6 a
Water added
N0P0 96.4  3.1 d 83.0  4.1 c 97.4  5.0 d 96.7  5.4 c
N8P4 104.8  4.2 c 92.4  3.8 b 112.2  6.4 c 107.2  4.4 b
N16P8 112.1  3.7 b 97.6  4.7 ab 122.2  7.3 b 112.3  4.5 ab
N32P16 120.3  2.2 a 102.6  4.4 a 133.5  5.9 a 116.4  4.5 a
Values greater than 100 indicate nutrient immobilization, and less than 100 indicate net mineralization. Data in the same column followed by the
same letter are not significantly different (P < 0.05). Values are means  S.E. (n = 4).
272 P. Liu et al. / Applied Soil Ecology 34 (2006) 266–275

Fig. 3. Effects of mixed N and P additions with or without water added on litter N and P concentrations of Allium bidentatum and Stipa krylovii after
decomposing for 100 days in the field. Vertical bars represent standard errors (n = 4).

small or no responses of litter decomposition to
increasing soil N input (Pastor et al., 1987; Theodorou
and Bowen, 1990; Downs et al., 1996). One of the
possible explanations could be that other nutrients, such
as phosphorus, were more limiting for decomposer
microbes as found by others (Ostertag and Hobbie,
1999; Cleveland et al., 2002). In our second experiment
with concurrent changes in N and P, the values of decay
constant varied more markedly than in the first
experiment. The above results suggest synergistic
effects of N and P on litter decomposition rates in
these two species that differ in litter quality.
Apart from the marked difference in the rates of litter
decomposition between the two species, Allium
bidentatum and S. krylovii also differed in the responses
of litter decomposition to increased soil water and
concurrent changes in soil N and P. The litter
decomposition of A. bidentatum was found to be more
responsive to soil water, whereas S. krylovii responded
more to concurrent changes in the soil N and P. The
stronger effect of soil water on the litter decomposition
of A. bidentatum than S. krylovii could be mainly due to
the differences in litter quality as reflected by both litter
N concentration and C:N ratio. The differences in
physical structures between the two litters could also
partly attribute to the different responses of litter
decomposition to soil water. Allium bidentatum is a
perennial herb with more fragile structures, while S.
krylovii is a bunch grass species with tissues that are
more fibrous.
Many studies have demonstrated that nutrient
additions can affect litter nutrient dynamics without
directly affecting the rate of decomposition (Hunt et al.,
1988; O’Connell, 1994; Downs et al., 1996). Such mode
of nutrient immobilization decouples nutrient miner-
alization from decomposition temporally, which may
have implications to nutrient retention in degraded
ecosystems. Organic N and P are much less mobile than
inorganic N and P and both plants and microbes can
make use of organic P by producing extracellular
phosphatases to mineralize common organic P com-
pounds into labile phosphates (McGill and Cole, 1981).
Incorporation of nutrients into decomposing litter
provides an important nutrient-retention mechanism
for the system (McGroddy et al., 2004).
In this study, we demonstrated marked differences in
the rates, as well as in the responses to changes in soil
nutrients and water, of litter decomposition between
 P. Liu et al. / Applied Soil Ecology 34 (2006) 266–275
two dominant grassland plant species that differ in life
forms and tissue chemistry. Our results clearly indicate
that the decomposition of high quality litter is more
likely to be limited by soil moisture regimes, while that
of low quality litter is more sensitive to nutrient
availability.
Our study area, Duolun County of Inner Mongolia,
belongs to the agro-pastoral ecotone of the Inner
Mongolia Plateau. Here S. krylovii is the main
construction species of the typical steppes, with A.
bidentatum as one of the predominant companion
species. In this agro-pastoral ecotone, sheep and cattle
grazing was once a major land use practice. Most of the
local grasslands were subjected to heavy grazing from
early 1980s to 2000, resulting in severe land degrada-
tion. Beginning in 2001, a national policy was put into
effect to ban grazing from the whole Duolun County in
an effort to reverse the land degradation problem.
Instead the grasses are now mowed for forage
production and officially free of grazing.
Existing literature on grassland vegetation dynamics
of northern China has mostly concerned with Leymus
chinensis and S. grandis steppes (e.g. Wang et al.,
1996a, 1996b, 2000a, 2000b; Liu et al., 2002). Although
specific information on the succession dynamics of S.
krylovii steppe during degradation and restoration is
lacking, the patterns should be similar to the S. grandis
steppe. In general, long-term overgrazing reduces the
status of S. krylovii into a sub-dominant species, while
A. bidentatum becomes a minor or rare species. Under
moderate grazing, the production of A. bidentatum and
S. krylovii decreases in a similar proportion as other
plants; Allium bidentatum would be more exposed and
hence likely to be more susceptible to grazing than other
plants in degraded grasslands because of its better
palatability to livestock. When protected from grazing,
the two species can both quickly restore their
dominance in the plant communities; but the recovery
is faster for A. bidentatum than Stipa spp. (Wang et al.,
1999). In a separate study, we found that in grassland
communities subjected to mowing, the maximum
biomass of A. bidentatum was 50% greater than S.
krylovii and ranked second in all plant species 5 years
after exclusion of grazing (unpublished data). The much
greater increases of A. bidentatum than S. krylovii
during the recovering of the degraded grassland
ecosystems emphasized greater importance of A.
bidentatum to the carbon cycle and nutrient dynamics
of the plant communities because of its faster
decomposition rate and greater sensitivity to soil water.
In conclusion, our findings suggest that plant species
with different litter qualities should be taken into
273
consideration when we are to model the carbon cycle
and nutrient dynamics in grassland ecosystems and that
A. bidentatum is expected to contribute more than S.
krylovii to the carbon cycle and nutrient dynamics of the
semi-arid grasslands of Inner Mongolia.
Acknowledgements
This research was supported by the National
Natural Science Foundation of China (30330150),
and by a startup fund to O.J. Sun from Institute of
Botany of the Chinese Academy of Sciences. We
thank the Duolun Restoration Ecological Experimen-
tation and Demonstration Station for access permis-
sion to the study site and technical assistance, and
Q.S. Chen for helps with field sampling and data
analysis.
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