Plant Soil (2008) 310:103–112
DOI 10.1007/s11104-008-9633-1
REGULAR ARTICLE
Variation in small-scale spatial heterogeneity of soil
properties and vegetation with different land use in semiarid
grassland ecosystem
Zhiyong Zhou & Osbert Jianxin Sun &
Zhongkui Luo & Hongmei Jin & Quansheng Chen &
Xingguo Han
Received: 28 January 2008 / Accepted: 22 April 2008 / Published online: 30 May 2008
# Springer Science + Business Media B.V. 2008
Abstract Soil properties (i.e. soil organic carbon, SOC;
soil organic nitrogen, SON; and soil C/N ratio) and
vegetation in a semiarid grassland of Inner Mongolia,
northern China, were studied with the method of
geostatistical analysis. We examined the spatial hetero-
geneity of soil and plants, and possible impacts of land
use on their heterogeneity and on the relationship
between soil resources and plant richness. Land use
affected small scale spatial heterogeneity in plants and
soil. SOC, SON and C/N ratio displayed autocorrela-
tion over a range of ∼2 m under most circumstances on
sites where livestock grazing had been excluded. The
uncontrolled grazing site (UG, i.e. unregulated grazing
by excessive livestock) displayed an increased range of
spatial autocorrelation and the total amount of vari-
ability in soil nitrogen over the other land use types.
Plant life forms and plant species exhibited spatial
autocorrelation over a range of about 2 m on the
Responsible Editor: Tibor Kalapos.
Z. Zhou : O. J. Sun (*)
Key Laboratory for Silviculture and Conservation
of Ministry of Education, College of Forest Science,
Beijing Forestry University,
Beijing 100083, China
e-mail: sunjianx@bjfu.edu.cn
Z. Luo : H. Jin : Q. Chen : X. Han
State Key Laboratory of Vegetation and Environmental Change,
Institute of Botany, Chinese Academy of Sciences,
Beijing 100093, China
grazing exclusion (GE) and mowed (MW) sites, while
pattern of spatial autocorrelation for several less com-
mon species on the UG site were difficult to predict.
Plant species richness was positively related with spatial
heterogeneity of SOC, SON and C/N on both GE and
MW sites, and with only SOC heterogeneity on the UG
site. These suggest that spatial soil heterogeneity plays
an active role in maintaining plant species richness.
However, we call for caution in generalization of the
control of spatial soil heterogeneity over plant richness
when multiple modes of disturbances are present, as we
found in this study that higher total amount of variation
in soil nitrogen and C/N ratio on the over-grazed UG
site did not lead to increased plant species richness,
and that land use had apparent effects on the patterns
of spatial heterogeneity in both vegetation and soil.
Keywords Geostatistics . Heterogeneity . Land use .
Species richness . Soil resources .
Semiarid grassland
Abbreviations
AB annuals and biennials
UG uncontrolled grazing
GE grazing exclusion
MW mowed
PR perennial rhizomatous grasses
PB perennial bunchgrasses
PF perennial forbs
SS shrubs and semi-shrubs
MSH the magnitude of spatial heterogeneity
104
Introduction
Spatial heterogeneity is considered as a ubiquitous
feature of natural ecosystems (Palmer 2003) and one
of the major drivers of biological processes (Milne
1991; Kumar et al. 2006). It is defined as spatially
structured variability of an ecological feature, which
may be a categorical or quantitative, explanatory or
dependent variable (Wagner and Fortin 2005). Coex-
istence theory and spatial-heterogeneity theory postu-
late that landscape with greater spatial heterogeneity
in environment, especially in a resource, can support
more plant species to coexist (Davies et al. 2005;
Kumar et al. 2006). This hypothesis leads to the
notion that species diversity is positively related with
spatial heterogeneity in extrinsic conditions (e.g. varia-
tion of soil properties and aspect) (Davies et al. 2005;
Kumar et al. 2006). But this spatial-heterogeneity
hypothesis contradicts the favorable-conditions hy-
pothesis (Davies et al. 2005), and has been questioned
by many ecologists who suggest that soil resource
heterogeneity has no positive effect on plant species
diversity (Bakker et al. 2003; Reynolds et al. 2007).
Favorable-conditions hypothesis considers that higher
resource levels may sustain more species to survive
(Levine and D’Antonio 1999; Hawkins et al. 2003).
Existence of such controversial views calls for
continued efforts in fundamental research targeting
explicitly the issue of interrelationship between spatial
heterogeneity and species diversity. In ecosystems
that lack complex vertical structures, such as grasslands,
soil resources could be a more important driver of
species diversity than other spatial features.
Grasslands are often used as a model ecosystem for
research on ecosystem function and biodiversity
(Tilman and Downing 1994; Wardle et al. 1999;
Harpole and Tilman 2007), as well as on the impacts
of variations in biotic and abiotic factors and their
interactions on ecological processes (Hutchings et al.
2003). The grasslands of northern China form an
integral part of the Eurasian grassland biome and have
been the focus of extensive research concerning the
impacts of global change and sustainable utilization
of land resources (Kang et al. 2007; Xiao et al. 2007).
Our previous research indicates that land use can
impose significant effects on biogeochemistry and
small-scale relationship between species diversity and
productivity of the region (Zhou et al. 2006, 2007).
Spatial heterogeneity has also been found to play a
Plant Soil (2008) 310:103–112
central role in shaping the community structure and
successional pattern of regional grassland ecosystems
(Wang et al. 1998; Bai et al. 2002; Cheng et al. 2007).
Recognition of the role of land use in regulating
ecosystem processes of grassland ecosystems is an
important step in formulating land management
strategies toward sustainable utilization and conser-
vation of grassland resources.
Development of geostatistics has provided an
effective tool in assisting assessment of spatial hetero-
geneity characteristics in ecological processes because
the parameters from the semivariogram model offer an
index to quantify the magnitude and scale of spatial
heterogeneity in a variate (Gross et al. 1995; Lane and
BassiriRad 2005). Many studies have been carried out
to determine spatial heterogeneous characteristics of a
variety of terrestrial ecosystems, especially on soil prop-
erties, by using this technique (Schlesinger et al. 1996;
Augustine and Frank 2001; Hutchings et al. 2003).
In a previous study, we found that different land
use had significant effects on plant species diversity
in the steppe grasslands of northern China (Zhou et al.
2006). Many processes may exert vital effect on this
heterogeneous procedure in natural system, such as
spatial interactions between biotic and abiotic factors
and subsequently differential responses of organisms
and the organisms themselves (Milne 1991; Kumar et
al. 2006). It is little known how land use may affect
spatial heterogeneity in vegetation and soil resources.
Would different land use result in different patterns of
spatial heterogeneity in soil resources and plant
communities? Would spatial heterogeneity in plant
community be possibly related to spatial heterogene-
ity in soil resources? To address these questions, we
studied the effects of three contrasting land use types,
i.e. uncontrolled grazing (UG, defined as unregulated
grazing by excessive livestock), grazing exclusion by
fencing (GE), and mowed (MW), on the small scale
(10×10 m) spatial heterogeneity in both soil resources
and plant communities by using 0.5×0.5 m grid-cell
sampling of plants and soil and the geostatistical
method of semivariance analysis in semiarid grass-
land ecosystem of northern China. Our objectives
were to determine: (1) the effect of three contrasting
land use types on the small-scale spatial heterogeneity
in vegetation and soil organic carbon and nitrogen;
and (2) possible linkage between species richness and
spatial soil heterogeneity. We hypothesized that the
pattern of small-scale plant heterogeneity would
Plant Soil (2008) 310:103–112
change with land use in grassland ecosystems, and
that greater soil resource heterogeneity would support
greater plant species richness.
Materials and methods
Study sites and experimental design
Field sites for this study are located in Duolun County
of Inner Mongolia (latitude 41°46′–42°39′ N, longitude
115°50′–116°55′ E, elevation 1,150–1,800 m a.s.l.),
northern China. The long-term annual mean air
temperature, the monthly mean air temperatures of the
coldest month (January) and the warmest month (July)
of the year are 1.6°C, −18.3°C and 18.7°C. Mean annual
precipitation is 385 mm, of which 67% falls between
June and August. Soil in the top 40-cm layer is
classified as chestnut (FAO-UNESCO 1974), below
40 cm a mixture of sandy soil and gravels. Vegetation
consists predominantly of common grassland plants of
the steppe zone including Stipa krylovii Roshev.
(perennial buchgrass; C3), Agropyron cristatum (L.)
Gaertner (perennial rhizomatous grass; C3), Allium
bidentatum Fisch. ex Prokh. & Ikonn.-Gal. (perennial
forb; C3), Artemisia frigida Willd. (semi-shrub; C3), and
Cleistogenes squarrosa (perennial bunchgrass; C4), etc.
Three study sites were selected, which included
uncontrolled grazing (UG); grazing exclusion by
fencing (GE); and mowed (MW). The UG site had
been heavily grazed since 1979, with an estimated
75% of above-ground biomass consumed by livestock
(mainly cattle and sheep) each year. The GE site was
established in 2001 by constructing a fence around
21 ha of previously grazed grassland. The MW site,
which also was previously grazed grassland, had been
subjected to mechanical mowing in late August each
year since 2001, with ∼80% of aboveground biomass
harvested as forage. Before 1978 the study area was
not utilized or managed and the major disturbance
resulted mainly from antelope and rabbit (occasionally
sheep) grazing, or wildfire. More detailed descriptions
of the study sites are given in Zhou et al. (2006, 2007).
In late summer 2005, we established a 10×10 m
plot on each of the three sites representing different
land-use types. Plots were laid out on horizontal and
apparently topographically uniform lands in order to
avoid anisotropic effects in subsequent heterogeneous
analysis. Each plot was sub-divided into 400 0.5×
105
0.5 m grid-cells, and plants and soil were sampled
from 100 cells spatially separated and evenly distrib-
uted across the plot (i.e. the sampling cells were set
for every other row and column among the grid-cells
in each plot). Measurements were made on above-
ground biomass by species, soil organic carbon and
nitrogen, and species richness.
Vegetation survey and sampling
On each 10×10 m plot, aboveground living tissues were
clip-harvested at ground level and sorted by species
within each of the 100 0.5×0.5 m grid-cells designated
as the sampling cells as described above between 15 and
25 August in 2005. Timing of the plant sampling
corresponded with the peak biomass period. All plant
samples were oven-dried to a constant mass at 65°C for
a minimum of 48 h. Biomass was determined by plant
species and then grouped into categories of total
community and life forms in our data analysis. The life
forms include: annuals and biennials (AB); perennial
rhizomatous grasses (PR); perennial bunchgrasses (PB);
perennial forbs (PF); shrubs and semi-shrubs (SS).
Diversity trait of plant community was described
simply as species richness, i.e. the number of species
over measurement area.
Soil sampling and chemical analysis
Soil samples were collected from the same 100 0.5×
0.5 grid-cells used for plant sampling. On each grid-
cell, soil samples were collected to a depth of 20 cm
from three locations distributed triangularly near the
cell center. Same sampling scheme was maintained
consistently for all cells where soil samples were
collected. The soil samples were taken immediately
after plant harvesting and removal of surface litters by
using a 4 cm diameter soil sampler. Three soil
samples within the same grid-cell were pooled as a
single composite sample for subsequent processing.
After being air-dried in a ventilation room, those soil
samples were cleared of roots and organic debris and
ground to pass 2-mm sieves for chemical analysis.
Soil samples were analyzed for organic carbon and
nitrogen. Soil organic carbon (SOC) was analyzed
following a modified Mebius method (Nelson and
Sommers 1982): i.e. 0.5 g soil was digested with 5 ml
of 1 N K2Cr2O7 and 10 ml of 98% H2SO4 at 150°C for
about 30 min, followed by titration of the digest liquid
106 Plant Soil (2008) 310:103–112

with standardized FeSO4. Total nitrogen was measured
with the Kjeldahl digestion procedure (Gallaher et al.
1976) and NH4+–N was determined colorimetrically by
the alkali method with a Tector Kjeltec System 1026
Distilling unit. SON was calculated as the difference
between the Kjeldahl–N and residual NH4+–N.
Geostatistical analysis
Semivariance analysis in geostatistics was used to
examine the pattern of spatial heterogeneity in SOC,
SON, C/N and spatial distribution of plant species and
plant life forms. Spherical, exponential, and linear
models were applied for fitting the semivariance data
by using the GS+ software program, version 5.0
(Gamma Design Software 1992). The “best fit” model
was judged as one with the least sum of squares
(Robertson et al. 1993). We presented only results of
spherical model as other two models are mechanisti-
cally poor representation of the spatial pattern of land
features. The magnitude of spatial heterogeneity
(MSH) in a variate and the distance, “range (A0)”,
over which the spatial dependence is expressed was
determined by: (1) the “sill” (C+C0), which is the
total sample variation, at which the semivariogram
levels off for the patterned data; (2) the “nugget” (C0),
which is the variation that is found at a scale finer
than the field sampling; and (3) the structure variance
(C), which is the difference between the sill and
nugget. MSH is then measured by the proportion of
total sample variation explained by spatially struc-
tured variance (C/[C+C0]). Spatially structured het-
erogeneity does not exist among sample points if the
value of MSH approaches zero; whilst proportion of
the total sample variance increases with increasing
spatial dependence between sample points (Robertson
et al. 1993; Schlesinger et al. 1996; Gross et al. 1995;
Lane and BassiriRad 2005). For soil properties (i.e.
SOC, TN, and C/N ratio), plant species, and plant life
forms, semivariogram was calculated following the
method of Schlesinger et al. (1996). We opted for
fixed observation scale at 10×10 m for practical
purpose. The lag interval was 1 m and the active lag
interval was 10.18 m, which was 72% of the
maximum distance. The number of pairs per lag
interval ranged from 231 to 809, with an average of
527 pairs per interval.
Coefficient of variation (CV) in soil properties was
measured within the north–south 0.5×10 m strips
containing the grid-cells for plant and soil sampling.
Results
Spatial heterogeneity of SOC, SON and C/N
For soil variables studied, the “best-fit” model to their
semivariograms was spherical on three sites with
exception of SOC and SON on the GE site (Table 1).
Calculated ranges of spatial autocorrelation for most
soil variables varied from 1.63 m (SON on the MW
site) to 2.05 m (SOC on the UG). The UG site had
exceptionally longer spatial dependence distance
(14.69 m) in SON than the other sites. MSH for all
soil variables on the GE, MW and UG sites ranged
from 56% (SON on the UG site) to 98% (SON on the

Table 1 Analysis of spatial structure for soil organic carbon (SOC), soil organic nitrogen (SON) and soil C/N ratio in a semiarid
grassland ecosystem of northern China by different land use with a 1-m lag interval

Variable Land use Model Range (A0, m) Nugget (C0 ×10−3) Sill [(C+C0)×10−3] MSH r2

SOC GE S 1.40 1.00 36.10 0.97 0.01

MW S 1.97 0.90 33.10 0.97 0.31
UG S 2.05 1.10 27.40 0.96 0.42
SON GE S 1.53 0.23 8.32 0.97 0.04
MW S 1.63 0.14 5.60 0.98 0.20
UG S 14.69 8.44 19.38 0.56 0.67
C/N GE S 1.66 0.46 14.92 0.97 0.16
MW S 1.95 0.80 30.80 0.97 0.56
UG S 1.91 2.40 57.20 0.96 0.11

GE Grazing exclusion without biomass removal; MW grazing exclusion with biomass removal by mowing; UG unregulated grazing
by excessive livestock; S spherical model; MSH magnitude of spatial heterogeneity
Plant Soil (2008) 310:103–112 107

MW site). But the UG site had lower MSH in SOC, lation beyond the limit of the plot size (Table 3).
SON and soil C/N ratio than the MW site (Table 1). MSH ranged from 80% (A. frigida on the MW site) to
98% (most species on the GE and MW sites, and S.
Spatial heterogeneity of vegetation krylovii and A. frigida on the UG site).

Spherical model provided the best fit to the semi-
variograms for all of the five plant life forms on the
GE site, and for all but the PR on the UG site; whilst
on the MW site, spherical model provided significant
fit to the semivariograms for all but the PR and SS
plant life forms. Most plant life forms displayed a
range of spatial autocorrelation at or less than 2 m on
all the three sites (Table 2). The range of spatial
autocorrelation for the AB on the UG site was beyond
the measurable scale (the analysis gave a default
value of 30.99). MSH ranged from 85% (i.e. AB on
the UG site) to 99% (i.e. AB on the MW site, PR on
the GE site, and SS on the UG site).
At individual level, spherical model gave the best
fit to the semivariograms for most plant species, while
no model was found to produce significant fit for C.
squarrosa, A. bidentatum and Carex korshinskyi on
the UG site and for C. korshinskyi on the GE site. The
range of spatial autocorrelation varied from 1.52 m
(A. frigida on the UG site) to 3.01 m (S. krylovii on
the MW site) with exception of A. frigida on the MW
site, which displayed the range of spatial autocorre-
Coefficient of variation (CV) in SOC, SON and C/N
Value of CV varied with the soil variable and the site
(Fig. 1): it was similar for SOC among the GE, MW,
and UG sites, but twice high on the UG site than on
the MW and GE sites for SON, and differed sig-
nificantly (p<0.05) among the three sites for soil C/N
ratio which ranked in the order of UG > MW > GE.
Relationship between species richness and soil
heterogeneity
Species richness was found to be related significantly
(p<0.05) and positively with SOC heterogeneity (as
indicated by the value of CV) across all the three sites
(Fig. 2a,d,g) and with SON and soil C/N ratio
heterogeneity (as indicated by the value of CV) on
the GE and MW sites (Fig. 2b,e,c,f). There was also a
tendency of negative relationship between species
richness and SON heterogeneity (as indicated by the
value of CV) on the UG site, albeit lack of statistical
significance (Fig. 2h).

Table 2 Analysis of spatial structure for plant life forms in a semiarid grassland ecosystem of northern China by land use, with a 1-m
lag interval

Life form Land use Model Range (A0, m) Nugget (C0 ×10−3) Sill [(C+C0)×10−3] MSH r2

AB GE S 1.98 49 782 0.94 0.35

MW S 1.63 0.01 4.98 0.99 0.24
UG S 30.99 8.20 53.40 0.85 0.78
PR GE S 1.70 14 971 0.99 0.21
MW S 1.42 110 397 0.97 0.03
UG S 1.56 4.20 289 0.99 0.07
PB GE S 1.95 6.60 258 0.97 0.71
MW S 2.29 9.60 318 0.97 0.57
UG S 2.07 14.00 784 0.98 0.67
PF GE S 1.99 8.70 275 0.97 0.41
MW S 1.97 10.80 234 0.95 0.25
UG S 1.92 63.00 859 0.93 0.19
SS GE S 1.88 4.30 253 0.98 0.58
MW S 2.04 11.70 232 0.95 0.09
UG S 1.45 5.00 440 0.99 0.14

AB Annuals and biennials; PR perennial rhizomatous grasses; PB perennial bunchgrasses; PF perennial forbs; SS shrubs and
semishrubs; GE grazing exclusion without biomass removal; MW grazing exclusion with biomass removal by mowing; UG
unregulated grazing by excessive livestock; S spherical model; MSH magnitude of spatial heterogeneity
108 Plant Soil (2008) 310:103–112

Table 3 Analysis of spatial structure for plant species in a semiarid grassland ecosystem of northern China by land use, with a 1-m
lag interval

Plant species Land use Model Range (A0, m) Nugget (C0 ×10−3) Sill [(C+C0)×10−3] MSH r2

Stipa krylovii GE S 1.84 4.40 283 0.98 0.72

MW S 3.01 44.00 392 0.89 0.63
UG S 2.13 20.00 865 0.98 0.73
Artemisia frigida GE S 1.85 4.10 250 0.98 0.54
MW S 30.99 118 579 0.80 0.81
UG S 1.52 6.00 375 0.98 0.19
Potentilla acauli GE S 1.89 17.00 805 0.98 0.27
MW S 1.88 35.00 923 0.96 0.43
UG S 2.14 104 1,225 0.92 0.27
Cleistogenes squarrosa GE S 1.81 3.10 163 0.98 0.66
MW S 1.85 4.10 203 0.98 0.46
UG S 1.34 0.70 36.10 0.98 0.00
Allium bidentatum GE S 2.20 12.00 339 0.97 0.47
MW S 2.09 8.70 300 0.97 0.64
UG S 1.71 2.90 39.60 0.93 0.03
Carex korshinskyi GE S 1.43 1.60 53.30 0.97 0.03
MW S 1.75 1.50 75.10 0.98 0.29
UG S 1.36 0.40 190 1.00 0.00

GE Grazing exclusion without biomass removal; MW grazing exclusion with biomass removal by mowing; UG unregulated grazing
by excessive livestock; S spherical model; MSH magnitude of spatial heterogeneity

Discussion heterogeneity of vegetation (Augustine and Frank

Geostatistics has been extensively used for examining
the degree of spatially structured, local variability in
soil resources (Robertson et al. 1993; Schlesinger et
al. 1996; Augustine and Frank 2001) and the spatial
2001). The scale and magnitude of spatial variation
have been shown to differ significantly among
different systems. For example, Palmer (1990) found
that most elements showed spatial autocorrelation
within a distance of 5 m in forest soils, whereas the

Fig. 1 Coefficient of varia- 35

tion [CV=(SD/mean)×
100%; n=10; ±SE] for soil GE b
organic carbon (SOC), soil 30 MW
organic nitrogen (SON), and FG
Coefficient of variation (%)

soil C/N ratio in a semiarid c

25
grassland of northern China
by land use. Vertical bars
a b
indicate standard errors of 20 a a
means (n=10). Means with
different letters above the a
histogram bars indicate sig- 15 a
a
nificant differences between
soil variables with land use 10
at p<0.05. GE Grazing ex-
clusion without biomass re-
moval; MW grazing 5
exclusion with biomass re-
moval by mowing; UG un- 0
regulated grazing by
excessive livestock SOC SON C:N

Soil variables
Plant Soil (2008) 310:103–112 109
Fig. 2 Relationship GE MW UG
between species richness 30
and heterogeneity [i.e. a d g
coefficient of variation, 25
CV=(SD/mean)×100%] of

CV of SOC (%)
20
soil properties (soil organic
carbon, SOC; soil organic 15
nitrogen, SON; and soil C/N
ratio) in a semiarid grass- 10
land ecosystem of northern
China by land use. GE 5 y = - 49.12 + 3.75 x y = 1.94 + 0.81 x y = - 15.18 + 1.81 x
2 2
(r = 0.76; p = 0.0009; n = 10) (r = 0.57; p = 0.012; n = 10) 2
( r = 0.49; p = 0.024; n = 10 )
Grazing exclusion without
0
biomass removal; MW 50 e h
b
grazing exclusion with bio- y = - 39.57+ 2.99 x y = 0.47 + 0.59 x
2
mass removal by mowing; 40 (r2 = 0.72; p = 0.002; n = 10) (r = 0.43; p = 0.039; n = 10 )
CV of SON (%)

UG unregulated grazing by
excessive livestock 30

20

10
y = 83.88 - 3.20 x
2
( r = 0.29; p = 0.108; n = 10 )
0
35
c f i
CV of soil C:N ratio (%)

30

25

20

15

10

5 y = - 61.92 + 4.47 x y = - 17.42 + 1.76 x y = 18.57 + 0.29 x

(r2 =0.43; p = 0.04; n = 10) 2
( r = 0.53; p = 0.018; n = 10 ) ( r2 = 0.007; p = 0.818; n = 10 )
0
10 12 14 16 18 20 22 2410 12 14 16 18 20 22 2410 12 14 16 18 20 22 24

Species richness (10 x 0.25 m2)

range over which there was spatial dependence for and the total amount of variability (as indicated by the
NH4 and NO3 was up to 20 m in an old-field value of CV) in SON over the other land use types.
community in Michigan (Robertson 1987). Grassland While most studies have demonstrated spatial
under a uniform cover usually possesses a longer pattern of soil properties from plant individual or
distance of spatial autocorrelation in soils than in species level (Kleb and Wilson 1997) to landscape
woody areas (Schlesinger et al. 1996). In this study, scale (Augustine and Frank 2001) in a variety of
we found that soil organic matter and C/N ratio in a ecosystems, far less attentions are paid to spatial
grassland ecosystem, where the livestock grazing had distribution of plants and their relationships with
been excluded (i.e. on the GE, MW sites), displayed spatial heterogeneity in environment. In this study,
autocorrelation over a range of ∼2 m under most we quantified the spatial pattern of vegetation by life
circumstances, similar to the findings of Bai et al. forms and by dominance in the local plant commu-
(2002) for the same region. Spherical model did not nities. Most of the plant life forms and species
provide the best fit to the semivariograms for SOC exhibited spatial autocorrelation over a range of about
and SON on the GE site, indicating that the 2 m on the GE and MW sites, while the pattern of
patchiness did not occur at the scales studied on this spatial autocorrelation for several less abundant
site. The UG site represented a situation of over- species on the UG site were poorly described by any
grazing for our study area, which was found to of the models used in our study. Disturbance and
display an increased range of spatial autocorrelation environmental heterogeneity have been found to
110
affect mainly the temporal and spatial variability of
satellite species that contribute to the overall diversity
of tallgrass prairie communities (Collins and Barber
1985; Collins and Glenn 1990). Aboveground bio-
mass removal by mowing can meliorate the vertical
spatial heterogeneity of sunlight which plays an
important role for understory species colonization,
while reducing or preventing the dominance of
abundant plants (Howe 1999). When livestock graz-
ing are excluded, such as that on the GE site, the
standing dead litter may exert great impact on spatial
pattern of plant community. We found that the UG
site had greater magnitude of spatial heterogeneity for
the shrub and semi-shrub species than the GE and
MW sites. Moreover, different species can respond to
their environment factors at different scales and these
scales are correlated with the movement ranges of
organisms (Holland et al. 2004). Combined effects of
abiotic and biotic processes are therefore expected to
regulate the spatial distribution of plant species and
plant life forms.
There exist complex plant–soil feedbacks with-
in native plant communities. Species composition
(Milchunas and Lauenroth 1995) and spatial distri-
bution of plant individuals (Vinton and Burke 1995)
can influence resource availability and heterogene-
ity at a variety of spatial scales in grasslands. Soil
heterogeneity can exert strong effects on many
species physiological or morphological properties
(Einsmann et al. 1999). For example, heterogeneity
in soil resources can affect the location of root
biomass, because plant roots are able to proliferate
in nutrient-rich patches (Hutchings et al. 2003). It is
evident in this study that the spatial autocorrelation
range is of the similar patch scales for soil resources
and vegetation.
Greater heterogeneity of resources or other fitness-
constraining environmental factors are known to
result in higher species diversity. This underlies the
spatial-heterogeneity hypothesis, which predicts that
there exists a positive relationship between plant
species diversity and spatial heterogeneity in extrinsic
conditions (Davies et al. 2005). In this study, we used
the coefficient of variation, CV, as a surrogate variable
for spatial heterogeneity for exploring relationship
between spatial heterogeneity in soil resources and
plant species richness by land use. Although many
empirical studies have shown that varying resource
heterogeneity has no effect on plant species richness
Plant Soil (2008) 310:103–112
(Wijesinghe et al. 2005; Reynolds et al. 2007), or the
relationship is scale-dependent (Anderson et al. 2004),
we found that plant species richness was positively
related with the CVs of SOC, SON and soil C/N ratio,
especially on sites of grazing exclusion (i.e. the GE
site) and plant harvesting by mechanical mowing (i.e.
the MW site). The match between scale of plant
patches and that of soil resource patches may suggest
the possibility of this relationship (Wijesinghe et al.
2005). Such positive relationship has been attributed
to niche partitioning by some researchers (Davies et
al. 2005; Harpole and Tilman 2007). Competition
intensity is generally expected to be intense within
nutrient rich patches under heterogeneous conditions,
and plant species with precise foraging ability are
considered being at a competitive advantage when
growing in such environment, while the nutrient poor
patches would act as refuges for less precise foragers,
or less competitive species, from intense competition
in heterogeneous environment (Hutchings et al.
2003). It is the spatial heterogeneity, especially the
comparatively nutrient poor patches, in environment
that provides more chances for satellite species to
successfully colonize in grasslands when grazers are
removed. Additionally, SOC is mainly influenced by
spatial pattern of plant species composition and the
spatial-temporal distribution of litter quality and input
(Pastor et al. 1998), which partly explain the positive
relationship between specie richness and SOC.
Because of the computational limitation in geo-
statistical analysis, our study was conducted using
non-replicated plots over a relatively restricted area.
This deficiency in experimental design may cause
some degree of bias in the resulting data. Nonetheless,
it is clear in this study that spatial soil heterogeneity
plays an active role in maintaining plant species
richness. However, any generalization of the control
of spatial soil heterogeneity over plant diversity needs
to be made with caution when multiple modes of
disturbances are present. For example, our results
showed that the higher total amount of variation in
SON and C/N ratio on overgrazed UG site did not
lead to increased plant species richness, and that land
use had apparent effects on the pattern of spatial
heterogeneity in both vegetation and soil. This study
offers further evidence that the relationship between
spatial heterogeneity in extrinsic environment and
plant diversity is a complex balance among variant
ecological processes.
Plant Soil (2008) 310:103–112
Acknowledgements Principle funding for this research was
provided by the National Natural Science Foundation of China
(grant no. 30521002). We gratefully acknowledge the field and
laboratory assistance from Wenming Bai, Wenyan Yang, Wei
Zhao, Ping Liu, Xuelin Zhang, Hongtao Zhao, Guangmei Wang,
Yingzhi Gao, Jin Liu, Jiaqian Tian and Zhidan Yan. We thank
Wenting Xu for helpful advice on data analysis and the Duolun
Restoration Ecology Experimentation and Demonstration Station
for permission to conduct this research on its experimental field
sites. The constructive comments by two anonymous reviewers
have helped to improve this paper over an earlier version.
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