Rangeland Ecol Manage 62:345–350 | July 2009

Differences in Net Primary Productivity Among Contrasting Habitats in Artemisia

ordosica Rangeland of Northern China
Chunping Li,1 Osbert Jianxin Sun,2 Chunwang Xiao,3 and Xingguo Han4
Authors are 1PhD student, 3Associate Professor, and 4Professor, State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese
Academy of Sciences, Beijing 100093, People’s Republic of China; and 2Professor, Key Laboratory for Silviculture and Conservation of Ministry of Education,
College of Forest Science, Beijing Forestry University, Beijing 100083, People’s Republic of China.

Abstract
Artemisia ordosica Krasch. is a semishrub native to the Ordos Plateau of Inner Mongolia, northern China, and forms a unique
and dominant vegetation type in the sandland of the region. To determine the variation of productivity in A. ordosica rangeland,
we investigated net primary production (NPP), fine root turnover, soil microbial C (Cmic), and soil organic carbon density
(SOCd) on sand dunes differing in mobility (i.e., fixed, semifixed, and shifting sand dunes) in Mu Us sandland. We found that,
on an area basis, the NPP, SOCd, Cmic, and fine root turnover rates all increased with increasing vegetation cover. However, the
ratios of root NPP to total NPP (RMRN) increased with declining vegetation cover. Total NPP varied markedly among habitats
and ranged from 18.3 g ? m22 ? yr21 for communities on the shifting sand dunes to 293.8 g ? m22 ? yr21 for communities on the
fixed sand dunes; whereas the rates of fine root turnover varied from 0.16 ? yr21 to 0.54 ? yr21. Our study demonstrated that
habitat change in sandland has significant impacts on ecosystem productivity by affecting many related aspects of NPP. From
the perspective of biomass production, protection of the semifixed dunes from degradation should be taken as a higher priority
than trying to convert shifting sand dunes to semifixed sand dunes; whereas conversion of semifixed sand dunes to fixed sand
dunes would appear to be a much easier task than restoring shifting sand dunes.

Resumen
Artemisia ordosica Krasch. es una subarbustiva nativa de la Planicie de Ordos del interior de Mongolia, al norte de China,
forma un único y dominante tipo de vegetación en la región de las tierras arenosas. Para determinar la variación in la
productividad de los pastizales con hábitat de A. ordorisca, investigamos la producción primaria neta (NPP), cambios en raı́ces
finas, C microbial en el suelo (Cmic), y densidad de carbono orgánico en el suelo (SOCd) en las dunas de arena de diferente
movilidad (ej., fijas, semifijas, y dunas de arena cambiantes) en las tierras arenosas de Mu Us. Encontramos que, en base a área,
la NPP, SOCd, Cmic, y la tasa de cambio de raı́z fina se incrementaron al aumentar la cobertura de vegetación. Sin embargo, las
tasas de NPP de la raı́z a total NPP (RMRN) se incrementaron al disminuir la cobertura vegetal. La NPP total varió
notablemente entre hábitats, dentro del rango de 18.3 g ? m2 ? yr para comunidades en las dunas de arena cambiantes, a
293.8 g ? m2 ? yr en las comunidades en las dunas de arena fijas; mientras que las tasas de cambio de raı́ces finas variaron de
0.16 ? yr21 a 0.54 ? yr21. Nuestro estudio demostró que el cambio de hábitat en las tierras arenosas tuvo un impacto significativo
en la productividad, afectando muchos aspectos relacionados con la NPP. Desde el punto de vista de producción de biomasa, la
protección de las dunas semifijas de la degradación debe considerarse de mayor prioridad que tratar de convertir las dunas
cambiantes a semifijas; mientras que la conversión de las dunas semifijas a dunas fijas puede parecer una tarea mucho más fácil
que la restauración de las dunas de arena cambiantes.
Key Words: Artemisia ordosica, net primary production (NPP), Ordos Plateau, soil organic carbon (SOC) density, turnover rate

INTRODUCTION
Global warming and more frequent occurrence of drought are
expected to accelerate desertification processes of the world
(Emanuel et al. 1985; Pickup 1998; Thomas et al. 2005). One of
the regions that is in great risk of expanding desert is the Ordos
Plateau in the middle of northern China, where under strong
influence of continental climate, rising temperatures are predicted
Research was jointly funded by a grant from the National Natural Science Foundation of
China (30521002) and a grant from the Chinese Academy of Sciences (KZCX2-XB2-01).
Correspondence: Dr Osbert Jianxin Sun, Key Laboratory for Silviculture and Conservation of
Ministry of Education, College of Forest Science, Beijing Forestry University, Beijing 100083,
People’s Republic of China. Email: sunjianx@bjfu.edu.cn
Current address: Chunping Li, College of Life Sciences, Northwest Sci-Tech University of
Agriculture & Forestry, Yangling, Shannxi 712100, People’s Republic of China.
Manuscript received 3 August 2007; manuscript accepted 23 November 2008.
to decrease soil water (Fu and An 2002; Editorial Board of the
State Assessment Report on Climate Change of China 2007),
making the region more vulnerable to desertification.
The Ordos Plateau has a predominantly semiarid climate and
rich deposits of loess and sands. Desertification, coupled with soil
erosion, has severely impacted the ecosystem productivity and
socio-economic development of the region. Mu Us sandland on
Ordos Plateau contains three major types of habitat based on
vegetation cover and mobility: fixed sand dunes, semifixed sand
dunes, and shifting sand dunes. In the past, anthropogenic
disturbance such as grazing and cultivation had been largely
responsible for a transformation of fixed sand dunes to semifixed
and shifting sand dunes; however, recent efforts in ecological
restoration through air-seeding afforestation and other biological
and mechanical sand-fixation schemes have, to a certain extent,
given rise to a reversal of the desertification process.

RANGELAND ECOLOGY & MANAGEMENT 62(4) July 2009 345

 An important aspect of alterations in ecosystem structure and
function is changes in biomass and net primary productivity
(NPP) with habitat. A shift of NPP in quantity and allocation is
making it very difficult to compute the global carbon budget
when the impact of human-induced global climate change is to
be assessed (Intergovernmental Panel on Climate Change
2007). Of particular importance is the belowground carbon
pool, which is estimated to sequester more than half of the
world’s carbon and plays an active role in offset fossil fuel
emissions (Lal 2004). In arid or semiarid ecosystems, the
percentage of belowground carbon is even higher because root
system turnover rate is low (Schlesinger et al. 1990; Gill and
Jackson 2000) and more assimilated carbon is allocated to
belowground compartments (Rice et al. 1998). Despite dryland
ecosystems covering nearly one-third of the earth’s land
surface, the fundamental knowledge about the responses of
the carbon cycle to environmental change is still limited and is
poorly documented relative to other main biomes.
The Artemisia ordosica Krasch. community is a unique
vegetation type endemic to Ordos Plateau. A. ordosica is a
semishrub that occurs in habitats ranging from fixed to shifting
sand dunes (Zhang 1994). The species is of great importance to
the regional socio-economic activities as a source of firewood
and fodder plants for livestock, in addition to its role in
ecosystem functioning. Research conducted in the region has
helped to gain some insight in understanding the succession
dynamics (Wang and Liang 1997; Wang et al. 1997) and
general productivity (Wang and Li 1994; Liu 1998; Cheng et al.
2001) of A. ordosica communities. However, detailed infor-
mation is lacking on the responses of above- and belowground
NPP patterns in A. ordosica rangeland to changing habitats
caused by natural and anthropogenic disturbances, which is key
to understanding the adaptive strategy of the vegetation to
long-term environmental perturbations.
We conducted a 2-yr study on carbon cycling in A. ordosica
communities in three types of habitat, i.e., fixed, semifixed, and
shifting sand dunes, in the Mu Us sandland. The objectives of
this study were to 1) assess NPP variation in A. ordosica
communities among the three habitat types, and 2) determine
the habitat effects on fine-root production, turnover rate, soil
organic carbon density, and soil microbial biomass carbon.
Findings from this study will be useful to those who manage
rangelands in the arid and semiarid areas of northern China
and who predict the carbon budget in this region under future
environments.
MATERIALS AND METHODS
Study Site
This study was conducted from July 2004 to September 2005
on sites near the Ordos Sandland Ecological Research Station
of the Chinese Academy of Sciences (lat 39u299N, long
110u119E, 1 295 m above sea level). The climate typically is
continental and semiarid (aridity index 0.3). Annual mean
precipitation is ,358 mm, of which 60–80% falls between
June and August. Annual mean temperature varies from 6.0uC
to 8.5uC; the monthly mean temperature fluctuates between
20uC and 24uC in July and is approximately 210uC in January.
Much of the region is covered by loose sands of varying depth.
346
Cover of other species, which consist of Hedysarum mongo-
licum Turcz., Chenopodium aristatum L., and Agriophyllum
squarrosum (L.) Moq., is 1–5% in all habitats. Fixed,
semifixed, and shifting sand dunes are the three typical
landscapes on Ordos Plateau. Percentage of vegetation cover
is the index often used to determine the type of sandland
because the dune activity correlates positively with the
vegetation cover.
Aboveground and Coarse Root Biomass and NPP Measurement
Three 10 m 3 10 m plots were established in each of the fixed,
semifixed, and shifting sand dunes near the research station.
For a detailed description of the characteristics of the three
habitats, please refer to the article by Li and Xiao (2007). The
height and the canopy diameter (D) in two perpendicular
directions were measured for all A. ordosica plants on each plot
at a monthly interval from May to September in 2005. The
aboveground and coarse root (D . 2 mm) biomass of individ-
ual A. ordosica plants was determined based on the site-specific
allometry (Li and Xiao 2007). The aboveground and coarse
root biomass of other plants was directly measured by
harvesting and weighing of oven-dried samples (65uC to
constant weight) from May to September in 2005. Above-
ground and coarse root NPP of A. ordosica communities were
estimated as the difference between maximum and minimum
biomass of the target components, which were measured in
May and September, respectively.
Fine Root Biomass and NPP Measurement
A soil corer (inner diameter of 8 cm) was used to collect root
samples for estimating fine root biomass at six random points
on each plot. Cores were collected at monthly intervals from
early May to late September 2005. Samples from each location
were taken to a depth of 100 cm and separated into 0–5-, 5–15-,
15–30-, 30–45-, 45–60-, 60–75-, 75–90-, and 90–100-cm layers.
The fine roots were washed and sorted into classes of 0–1- and
1–2-mm diameters. Live and dead root fragments were separated
by visual inspection based on attributes described by Persson
(1980) and Vogt and Persson (1991) in which the xylem of dead
roots appears darker and deteriorated, the degree of cohesion
between the cortex and periderm decreases, and root tips
become brittle and less resilient. Dry weights were determined
after oven-drying at 65uC to constant weight.
In order to estimate fine root (D # 2 mm) productivity, a
simplified ingrowth core method was used (Janssens et al.
2002). Similar to the sampling method described above, soil
core samples were collected at six random locations on
each plot in August 2004. Samples from each location were
taken to a depth of 100 cm and separated into 0–5-, 5–15-, 15–
30-, 30–45-, 45–60-, 60–75-, 75–90-, and 90–100-cm layers.
After removal of roots, the soils were placed back to the same
holes with the same stratification. We tried to retain the
original bulk density by filling each stratum with the
appropriate mass of soil and covering the surface with the
original litter. Those cores were sampled again in August 2005
for newly produced fine roots to calculate belowground net
primary productivity.
The rate of fine root turnover was estimated as the ratio of
the total fine root biomass produced in 1 yr over the mean
Rangeland Ecology & Management

Figure 1. Seasonal pattern of biomass density in Artemisia ordosica
communities of different habitats in Ordos Plateau in 2005. Error bars
represent 6 1 SE (n 5 3).
standing biomass of fine roots (Aber et al. 1985; Aerts et al.
1992). The ratio of root NPP to total NPP (RMRN, g ? g21) was
also calculated.
Calculation of Soil Organic Carbon Density
Soil samples were collected to a depth of 100 cm and separated
into eight layers (as described previously) from four random
locations on each plot in September 2005 with a soil corer
(inner diameter 4 cm). The samples were air-dried after the
surface organic materials or root fragments were completely
sieved off. Soil organic carbon (SOC) content was determined
for each sample by the dichromate oxidation method as
described in Bao (2000).
SOC density for a soil profile with a depth of d (cm), SOCd
(t ? C ? ha21), was calculated using the following equation:
X
n
ð1{Vi %Þ|Bi |Ci |Ti
SOCd ~ [1]
i~1
100
where n is the number of pedogenic horizons in the soil
survey, Vi% is the volumetric percentage of the soil fraction
. 2 mm (rock fragment), Bi is the bulk density (g ? cm23), Ti
is the thickness (cm) in the layer i, and Ci is the organic C
content (%). In this study, the depth of soil profiles for
SOCd calculations was set to 100 cm. Soil bulk density was
determined for each habitat type using three soil profiles.
Measurement of Soil Microbial Biomass C
Four soil samples were taken from the top 10 cm on each plot
with a soil corer (inner diameter of 4 cm) in September 2005.
Samples were stored in a chest cooler and taken to a laboratory
for soil microbial biomass C (Cmic) analysis using the
chloroform-fumigation method (Witt et al. 1999).
Statistical Analysis
One-way analysis of variance was used to detect the
significance of variation of the means of above- and below-
ground net primary production, root turnover rate, Cmic, SOC
62(4) July 2009
Figure 2. Above- and belowground NPP (columns) and belowground
NPP to total NPP (RMRN, line) in Artemisia ordosica communities of
different habitats in Ordos Plateau. Vertical bars represent 6 1 SE
(n 5 3), and different letters indicate significant differences of means for
the same variables between habitats at P , 0.05 by LSD test. NPP
indicates net primary production; RMRN, ratio of root net primary
production to total net primary production; LSD, least
significant difference.
density, and vertical distribution of fine roots on the three types
of habitats. Least significant difference (LSD) was used in the
multiple comparisons among habitats. Significant differences
were determined at P , 0.05 level unless otherwise stated. All
statistical analyses were performed with SPSS 13.0 for
Windows.
RESULTS
Habitat Effects on Above- and Belowground Biomass and NPP
The three types of habitats differed significantly in the seasonal
dynamics of biomass. Generally, biomass increased with time
within a single growing season at the community level, with the
rate of increase declining markedly from the fixed sand dunes
to the shifting sand dunes. Biomass increased from 189 to
444 g ? m22 on the fixed sand dunes, 88 to 236 g ? m22 on the
semifixed sand dunes, and 16 to 35 g ? m22 on the shifting sand
dunes (Fig. 1).
The above- and belowground NPP displayed a declining
trend from the fixed sand dunes to the shifting sand dunes
(Fig. 2). The aboveground NPP of the fixed sand dunes was
219.5 g ? m22 ? yr21, approximately twice that of the semifixed
sand dunes (119.1 g ? m22 ? yr21) and 10 times that of the
shifting dunes (11.5 g ? m22 ? yr21). Belowground NPP was
74.3 g ? m22 ? yr21, 42.2 g ? m22 ? yr21, and 6.8 g ? m22 ? yr21
for the fixed, semifixed, and shifting sand dune habitats,
respectively. However, the ratio of root NPP to total NPP
(RMRN, g ? g21) was significantly greater on the shifting sand
dunes than on the fixed and semifixed sand dunes (Fig. 2).
Habitat Effects on Fine Root NPP and Its Vertical Distribution
Fine root NPP of the two diameter classes displayed decreasing
patterns in response to the changing habitat types (Fig. 3a).
Namely, the 1–2 mm fine root NPP dropped from 4.9 g ?
m22 ? yr21 on the fixed sand dunes to 0.06 g ? m22 ? yr21 on the
shifting sand dunes; the , 1 mm fine root NPP dropped from
12.8 g ? m22 ? yr21 to 0.8 g ? m22 ? yr21, respectively. In all
347

Figure 3. Fine root a, NPP and b, turnover rates of two diameter
classes in Artemisia ordosica communities of different habitats in Ordos
Plateau. Error bars represent 6 1 SE (n 5 18), and different letters
indicate significant differences of means between habitats at P , 0.05 by
LSD test. NPP indicates net primary production; LSD, least significant
difference.
habitats, the NPP of fine roots , 1 mm was greater than that of
1–2 mm. As for the vertical distribution of root NPP, more
than 70% of the fine roots , 1 mm and 90% of the fine roots
1–2 mm were located in the upper 30 cm soil layer in all three
habitat types. Very little fine root NPP was observed below
60 cm (Fig. 4). The peak fine root NPP occurred at a depth of
about 30 cm along the soil profile, but the magnitude of the
fine root density differed markedly between different types of
habitats.
Habitat Effects on Fine Root Turnover
The turnover rate of fine roots , 1 mm was on average 3-fold
greater than that for total fine roots (# 2 mm diameter),
displaying a declining trend from the fixed sand dunes to the
shifting sand dunes (Fig. 3b). The difference in the turnover
rates was significant among habitat types and was markedly
greater between the semifixed and shifting sand dunes than
between the fixed and semifixed sand dunes.
Habitat Effects on SOCd and Cmic
SOCd differed significantly among the three habitat types, with
a declining trend along the habitat gradients from the fixed
sand dunes to the shifting sand dunes (Fig. 5). The differences
in SOCd were much greater between the fixed and semifixed
sand dunes than between the semifixed and shifting sand dunes.
348
Cmic mainly differed between the fixed sand dunes and the
other two types of habitats, and no significant difference was
found between the semifixed and shifting sand dunes (Fig. 5).
DISCUSSION
Habitat change caused by desertification could influence NPP
patterns because productivity is very sensitive to environmen-
tal perturbations (Huenneke et al. 2002). Our study in the
semiarid Mu Us sandland detected significant differences in
aboveground and belowground NPP in A. ordosica commu-
nities along habitat gradients representative of different levels
of land degradation; the declining NPP from the fixed sand
dunes to the shifting sand dunes reflected the deteriorating
abiotic and biotic conditions. The fixed sand dunes developed
relatively fertile soil containing higher soil organic nitrogen
and SOC in the succession process, whereas the activated sand
dunes suffered severe nutrient loss. Such difference in soil
fertility can account for variable productivity among different
habitats (Schlesinger et al. 1990; Busso 1997; Sharifi et al.
1999).
We found that NPP differed more between the semifixed and
shifting sand dunes than between the fixed and semifixed sand
dunes. This suggests that habitat change between semifixed and
shifting sand dunes would result in greater impacts than between
fixed and semifixed sand dunes on carbon accumulation in A.
ordosica communities. Transformation from semifixed to
shifting sands dunes would not only markedly reduce commu-
nity biomass and productivity, but also make the restoration
more difficult because biomass accumulation during the growing
season was shown to be far less on the shifting sand dunes.
Therefore, protecting the semifixed sand dunes from degrada-
tion should be taken as a higher priority than trying to convert
shifting sand dunes to semifixed sand dunes; conversion of
semifixed sand dunes to fixed sand dunes would appear to be a
much easier task than restoring shifting sand dunes.
Fine root NPP showed the same declining trend as
aboveground NPP at the community level, indicating that the
infertile soil condition on shifting sand dunes also limited the
fine root growth. However, for individual plants, the propor-
tion of fine root NPP over total NPP increased when habitat
type changed from the fixed and semifixed sand dunes to the
shifting sand dunes as shown by the results of RMRN.
Differences in habitat conditions are known to affect produc-
tion allocation in many plants, which likely is a necessary
adaptive strategy for providing sufficient nutrients and water to
plant growth and survival under poor resource conditions
(Nadelhoffer et al. 1985; Steele et al. 1997; Martı́nez et al.
1998; Snyman 2005).
Root turnover is a central component of ecosystem carbon
and nutrient cycling and is sensitive to many of the factors
considered in global change analysis (Aber et al. 1985;
Bloomfield et al. 1996). The turnover rate of A. ordosica roots
was found to increase with decreasing root diameter and was
greater on the fixed and semifixed sand dunes than on the
shifting sand dunes. Our estimate of the fine root turnover rate
for A. ordosica could be conservative due to potentially
underestimated NPP by not accounting for microbial decom-
position during the measurement period. Nonetheless, the
Rangeland Ecology & Management
Figure 4. Vertical distribution of root NPP density for two contrasting size classes in Artemisia ordosica communities under different habitats in
Ordos Plateau. Error bars represent 6 1 SE (n 5 18). NPP indicates net primary production.
values were in the general range found in many other studies
for shrubs in temperate zone (e.g., Caldwell et al. 1977;
Saterson and Vitousek 1984; Berendse et al. 1987; Martı́nez et
al. 1998; Gill and Jackson 2000; Lauenroth and Gill 2003), and
consistent with the result of Cmic. The patterns of Cmic closely
matched SOCd in relation to habitat types.
MANAGEMENT IMPLICATIONS
Our study demonstrated that habitat change in sandland has
significant impacts on ecosystem productivity by affecting
many related aspects of NPP. Thus, the level of productivity
provides direct guidance for land-use of different habitats.
Fixed sand dunes could be moderately used as rangeland
Figure 5. Soil organic carbon density (SOCd, left) at 1 m depth and soil
microbial biomass C (Cmic, right) in the 0–10 cm soil layer in Artemisia
ordosica communities of different habitats in Ordos Plateau. Error bars
represent 6 1 SE (n 5 12), and the same letters indicate no significant
differences between means of certain variables at P , 0.05 by LSD test.
LSD indicates least significant difference.
62(4) July 2009
because they have higher productivity and the exploitation of
semifixed sand dunes should be cautious because of its fragile
vegetation. Protection and overuse might trigger two opposite
successional pathways on semifixed sand dunes. From the
perspective of biomass production, protecting the semifixed
sand dunes from degradation should be taken as a higher
priority than trying to convert shifting sand dunes to semifixed
sand dunes; conversion of semifixed sand dunes to fixed sand
dunes would appear to be a much easier task than restoring
shifting sand dunes.
ACKNOWLEDGMENTS
The authors wish to thank the researchers at Ordos Sandland Ecological
Research Station of the Chinese Academy of Sciences for their technical
assistance throughout this study. We also thank Gang Li for his valuable
comments on the early version manuscript, and Zhenqing Yang for his help
with field work.
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