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Journal of Arid Land  2023, Vol. 15 Issue (1): 63-76    DOI: 10.1007/s40333-023-0001-z
Research article     
Biocrust-induced partitioning of soil water between grass and shrub in a desert steppe of Northwest China
YANG Xinguo1,2,3,*(), WANG Entian1,2,3, QU Wenjie1,2,3, WANG Lei1,2,3
1Northwest National Key Laboratory Breeding Base for Land Degradation and Ecological Restoration, Ningxia University, Yinchuan 750021, China
2Key Laboratory of Restoration and Reconstruction of Degraded Ecosystems, Ministry of Education, Ningxia University, Yinchuan 750021, China
3Ecology and Environment College, Ningxia University, Yinchuan 750021, China
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Maintaining the stability of exotic sand-binding shrub has become a large challenge in arid and semi-arid grassland ecosystems in northern China. We investigated two kinds of shrublands with different BSCs (biological soil crusts) cover in desert steppe in Northwest China to characterize the water sources of shrub (Caragana intermedia Kuang et H. C. Fu) and grass (Artemisia scoparia Waldst. et Kit.) by stable 18O isotopic. Our results showed that both shrublands were subject to persistent soil water deficiency from 2012 to 2017, the minimum soil depth with CV (coefficient of variation) <15% and SWC (soil water content) <6% was 1.4 m in shrubland with open areas lacking obvious BSC cover, and 0.8 m in shrubland covered by mature BSCs. For C. intermedia, a considerable proportion of water sources pointed to the surface soil. Water from BSCs contributed to averages 22.9% and 17.6% of the total for C. intermedia and A. scoparia, respectively. C. intermedia might use more water from BSCs in rainy season than dry season, in contrast to A. scoparia. The relationship between shrub (or grass) and soil water by δ18O shown significant differences in months, which partly verified the potential trends and relations covered by the high variability of the water source at seasonal scale. More fine roots at 0-5 cm soil layer could be found in the surface soil layer covered by BSCs (8000 cm/m3) than without BSCs (3200 cm/m3), which ensured the possibility of using the surface soil water by C. intermedia. The result implies that even under serious soil water deficiency, C. intermedia can use the surface soil water, leading to the coexistence between C. intermedia and A. scoparia. Different with the result from BSCs in desert areas, the natural withdrawal of artificial C. intermedia from desert steppe will be a long-term process, and the highly competitive relationship between shrubs and grasses also determines that its habitat will be maintained in serious drought state for a long time.

Key wordsdesert steppe      biological soil crusts      water resource      Caragana intermedia      Artemisia scoparia     
Received: 13 October 2022      Published: 31 January 2023
Corresponding Authors: YANG Xinguo     E-mail:
Cite this article:

YANG Xinguo, WANG Entian, QU Wenjie, WANG Lei. Biocrust-induced partitioning of soil water between grass and shrub in a desert steppe of Northwest China. Journal of Arid Land, 2023, 15(1): 63-76.

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Fig. 1 Location of the experimental site (a), landscape and habitat characteristics of the shrubland (b and c). (b), S-CK, shrubland without BSCs (biological soil crusts) cover; (c), S-BSC, shrubland with BSCs cover.
Fig. 2 Soil water contents in 0-300 cm (a-o) layer from 2012 to 2017. S-CK, shrubland with BSCs (biological soil crusts) cover; S-BSC, shrubland with BSCs cover; CK, natural grassland without replanted Caragana intermedia.
Fig. 3 Daily time series of relative increment in surface soil moisture (ISSM) in S-CK and S-BSC (biological soil crust) plots in the growth seasons (from May to October) during 2016-2017. (a), ISSM in S-CK and S-BSC; (b), SWC (soil water content) in 0-5 and 5-10 cm soil layers in S-CK; (c) SWC in 0-5 and 5-10 cm soil layers in S-BSC. S-CK, shrubland without BSCs cover; S-BSC, shrubland with BSCs cover.
Species Index Water source
(0-2 cm)
(2-30 cm)
(30-70 cm)
(>70 cm)
Caragana intermedia
Average (%) 22.9 34.3 22.8 20.0
CV (%) 62.5 58.6 63.4 41.5
Range (%) 0.0-55.6 9.5-80.4 2.7-63.3 2.2-33.1
Artemisia scoparia
Average (%) 17.9 41.6 22.1 18.0
CV (%) 81.3 73.9 108.2 67.1
Range (%) 3.7-47.2 8.3-90.7 1.2-76.1 3.0-36.0
Table 1 Average proportional water uptake from four potential soil sources for Caragana intermedia and Artemisia scoparia
Fig. 4 Water uptake percentage from different potential sources for Caragana intermedia and Artemisia scoparia in the rainy and dry seasons during 2016-2017. Water sources were defined as BSCs (biological soil crusts) ranging from surface (0-2 cm), shallow (2-30 cm), middle (30-70 cm) to deep (>70 cm). SWC, soil water content. Bars are standard errors.
Fig. 5 Fine root length density in 0-10 cm soil layer for Caragana intermedia in plots with (BSCs reserved) and without biological soil crusts (BSCs removed). Fine roots were defined as roots with a diameter within 0.5-2.0 mm. *, P<0.05 level. Bars are standard errors.
Plot Shrub morphological characters Biodiversity index
CA (m2) SH (m) BD (cm) CD (%) R H D E
S-CK 2.87±0.30 1.25±0.43 1.79±0.63 23.5±4.08 2.33±0.12 3.38±0.36 0.23±0.17 1.25±0.12
S-BSC 2.51±0.45 0.99±0.28 1.47±0.42 21.8±5.22 1.55±0.06 2.43±0.40 0.40±0.09 0.98±0.15
Table S1 Shrub morphological characters and biodiversity in S-CK and S-BSC
Plot Silt and clay (%) SOC (g/kg) TN (g/kg) TP (g/kg)
S-CK 11.08±5.08a 3.94±0.35a 0.29±0.07a 0.29±0.06a
S-BSC 15.44±7.04a 4.04±0.27a 0.23±0.06a 0.33±0.02a
Table S2 Information of surface soil (0-20 cm) in S-CK and S-BSC
Fig. S1 Upper-direct growth of roots in 0-20 cm soil layer for C. intermedia in the rainy season in 2018
Fig. S2 Soil water content and its CV (coefficient of variation) values in different soil layers during 2012-2017. CV-S1, CV values of soil water contents during 2012-2017 in S-CK plot; CV-S2, CV values of soil water contents during 2012-2017 in S-BSC plot; CV-CK, CV values of soil water contents during 2012-2017 in CK plot; AV-S1, average values of soil water contents during 2012-2017 in S-CK plot (shrubland without BSCs cover); AV-S2, average values of soil water contents during 2012-2017 in S-BSC plot (shrubland with BSCs cover); AV-CK, average values of soil water contents during 2012-2017 in CK plot (grassland without shrub).
Fig. S3 Root growth of A. scoparia from May to September
Fig. S4 Percentage of water uptake from four potential soil water sources for Caragana intermedia and Artemisia scoparia in the rainy and dry seasons during 2016-2017. Water sources were defined as BSCs (biological soil crusts) ranging from surface (0-2 cm), shallow (2-30 cm), middle (30-70 cm) to deep (>70 cm). Different uppercase letters indicate significant differences between C. intermedia and different water sources at P<0.05 level. Different lowercase letters indicate significant differences between A. scoparia and different water sources at P<0.05 level. * indicates significant differences between C. intermedia and A. scoparia at P<0.05 level. Bars are standard errors.
[1]   Angert A L, Huxman T E, Chesson P, et al. 2009. Functional tradeoffs determine species coexistence via the storage effect. Proceedings of the National Academy of Sciences, 106(28): 11641-11645.
[2]   Asbjornsen H, Mora G, Helmers M J. 2007. Variation in water uptake dynamics among contrasting agricultural and native plant communities in the Midwestern US. Agriculture, Ecosystems & Environment, 121(4): 343-356.
[3]   Chamizo S, Cantón Y, Miralles I, et al. 2012. Biological soil crust development affects physicochemical characteristics of soil surface in semiarid ecosystems. Soil Biology & Biochemistry, 49: 96-105.
doi: 10.1016/j.soilbio.2012.02.017
[4]   Chamizo S, Cantón Y, Lázaro R, et al. 2013. The role of biological soil crusts in soil moisture dynamics in two semiarid ecosystems with contrasting soil textures. Journal of Hydrology, 489: 74-84.
doi: 10.1016/j.jhydrol.2013.02.051
[5]   Chen L, Su Y, Li Y F, et al. 2019. Effects of heterogeneous habitats on phenotypic plasticity of Artemisia scoparia in the desert steppe of China. Acta Ecologica Sinica, 39(10): 3547-3556.
[6]   Chen Y L, Zhang Z S, Huang L, et al. 2017. Co-variation of fine-root distribution with vegetation and soil properties along a revegetation chronosequence in a desert area in northwestern China. CATENA, 151: 16-25.
doi: 10.1016/j.catena.2016.12.004
[7]   Cobos D R, Chambers C. 2010. Calibrating ECH2O soil moisture sensors, Application Note. Pullman: Decagon Devices Inc., 1-7.
[8]   Dawson T E. 1993. Water sources of plants as determined from xylem-water isotopic composition:perspectives on plant competition, distribution, and water relations. In: Ehleringer J R, Hall A E, Farquhar G D. Stable Isotopes and Plant Carbone Water Relations. San Diego: Academic Press, 465-496.
[9]   Eggemeyer K D, Awada T, Harvey F E, et al. 2009. Seasonal changes in depth of water uptake for encroaching trees Juniperus virginiana and Pinus ponderosa and two dominant C4 grasses in a semiarid grassland. Tree Physiology, 29(2): 157-169.
doi: 10.1093/treephys/tpn019 pmid: 19203941
[10]   Ehleringer J R, Phillips S L, Schuster W S F, et al. 1991. Differential utilization of summer rains by desert plants. Oecologia, 88: 430-434.
doi: 10.1007/BF00317589 pmid: 28313807
[11]   Grossiord C, Sevanto S, Dawson T E, et al. 2017. Warming combined with more extreme precipitation regimes modifies the water sources used by trees. New Phytologist, 213(2): 584-596.
doi: 10.1111/nph.14192 pmid: 27612306
[12]   Huang L, Zhang Z S. 2015. Stable isotopic analysis on water utilization of two xerophytic shrubs in a revegetated desert area: Tengger Desert, China. Water, 7(3): 1030-1045.
doi: 10.3390/w7031030
[13]   Jia Z Q, Zhu Y, Liu L. 2012. Different water use strategies of juvenile and adult Caragana intermedia plantations in the Gonghe basin, Tibet Plateau. PLoS ONE, 7(9): e45902, doi: 10.1371/journal.pone.0045902.
doi: 10.1371/journal.pone.0045902
[14]   Kidron G J, Monge, H C, Vonshak A, et al. 2012. Contrasting effects of microbiotic crusts on runoff in desert surfaces. Geomorphology, 139-140: 484-494.
[15]   Kidron G J. 2014a. Do mosses serve as sink for rain in the Negev Desert? A theoretical and experimental approach. CATENA, 121: 31-39.
doi: 10.1016/j.catena.2014.05.001
[16]   Kidron G J. 2014b. The negative effect of biocrusts upon annual-plant growth on sand dunes during extreme droughts. Journal of Hydrology, 508: 128-136.
doi: 10.1016/j.jhydrol.2013.10.045
[17]   Kowaljow E, Fernández R J. 2011. Differential utilization of a shallow-water pulse by six shrub species in the Patagonian steppe. Journal of Arid Environments, 75(2): 211-214.
doi: 10.1016/j.jaridenv.2010.10.004
[18]   Le Roux X, Bariac T, Mariotti A. 1995. Spatial partitioning of the soil water resource between grass and shrub components in a West African humid savanna. Oecologia, 104: 147-155.
doi: 10.1007/BF00328579 pmid: 28307351
[19]   Li X R. 2005. Influence of variation of soil spatial heterogeneity on vegetation restoration. Science in China Series D: Earth Sciences, 48: 2020-2031.
doi: 10.1360/04yd0139
[20]   Li X R, Tian F, Jia R, et al. 2010. Do biological soil crusts determine vegetation changes in sandy deserts? Implications for managing artificial vegetation. Hydrology Process, 24(25): 3621-3630.
doi: 10.1002/hyp.7791
[21]   Li X R, Zhang Z S, Huang L, et al. 2013. Review of the ecohydrological processes and feedback mechanisms controlling sand-binding vegetation systems in sandy desert regions of China. Chinese Science Bulletin, 58: 1483-1496.
doi: 10.1007/s11434-012-5662-5
[22]   Li X R, Zhang Z S, Tan H J, et al. 2014. Ecological restoration and recovery in the wind-blown sand hazard areas of northern China: relationship between soil water and carrying capacity for vegetation in the Tengger Desert. Science China Life Sciences, 57(5): 539-548. (in Chinese)
doi: 10.1007/s11427-014-4633-2 pmid: 24699917
[23]   Li X R, Zhang D H, Zhang F, et al. 2017. The eco-hydrological threshold for evaluating the stability of sand-binding vegetation in different climatic zones. Ecological Indicatiors, 83: 404-415.
[24]   Liang H R, Ji M, Ren J M. 2008. Phonological characteristics of nine varieties of Caragana. Journal of Inner Mongolia Forest Science and Technology, 34(3): 25-27. (in Chinese)
[25]   Liu J S, Xu X, Zhang Y, et al. 2010. Effect of rainfall interannual variability on the biomass and soil water distribution in a semiarid shrub community. Science China Life Sciences, 53(6): 729-737. (in Chinese)
doi: 10.1007/s11427-010-4014-4 pmid: 20602276
[26]   Liu L Y, Jia Z Q, Zhu Y J, et al. 2012. Water use strategy of different stand ages of Caragana intermedia in alpine sandland. Journal of Arid Land Resource & Environment, 26(5): 119-125. (in Chinese)
[27]   Liu W, Wang P, Li J, et al. 2014. Plasticity of source-water acquisition in epiphytic, transitional and terrestrial growth phases of Ficus tinctoria. Ecohydrology, 7(6): 1524-1533.
doi: 10.1002/eco.1475
[28]   Lu T, Zhao X N, Gao X D, et al. 2017. Soil water use strategy of dominant species in typical natural and planted shrubs in loess hilly region. Chinese Journal of Plant Ecology, 41(2): 175-185. (in Chinese)
doi: 10.17521/cjpe.2016.0253
[29]   Niu X W, Ding Y C, Zhang Q, et al. 2003. Studies on the characteristics of Caragana root development and some relevant physiology. Acta Botanica Boreali-Occidentalia Sinica, 23(5): 860-865. (in Chinese)
[30]   Ogle K, Reynolds J. 2004. Plant responses to precipitation in desert ecosystems: Integrating functional types, pulses, thresholds, and delays. Oecologia, 141: 282-294.
pmid: 15007725
[31]   Phillips D L, Gregg J W. 2003. Source partitioning using stable isotopes: Coping with too many sources. Oecologia, 136: 261-269.
pmid: 12759813
[32]   Schenk H J. 2008. Soil depth, plant rooting strategies and species' niches. New Phytologist, 178(2): 223-225.
doi: 10.1111/j.1469-8137.2008.02427.x pmid: 18371001
[33]   Song N P, Yang M X, Wang L, et al. 2014. Annual dynamic change of soil moisture in artificial Caragana forest in desert steppe region. Chinese Journal of Ecology, 33(4): 2618-2624. (in Chinese)
[34]   Yang D D, Zhao W, Chen L, et al. 2018. Seasonal conversion of the effects of biological soil crusts on surface hydrology in the artificial Caragana intermedia shrublands. Acta Botanica Boreali-Occidentalia Sinica, 38(7): 1349-1356. (in Chinese)
[35]   Yang X G, Zhao W, Chen L, et al. 2015. Evolution characteristics of soil and vegetation in artificial Caragana forest in desert steppe. Ecology & Environment, 24(4): 590-594. (in Chinese)
[36]   Zhang L, Wu B, Ding G D, et al. 2010. Root distribution characteristics of Salix psammophyla and Caragana korshinskii in Mu Us Sandy Land. Journal of Arid Land Resource & Environment, 24(3): 158-161. (in Chinese)
[37]   Zhang P P, Zhao Y G, Wang Y, et al. 2014. Impact of biological soil crusts on soil water repellence in the hilly Loess Plateau region, China. Chinese Journal of Applied Ecology, 25(3): 657-663. (in Chinese)
[38]   Zhang Z S, Chen Y, Zhang J G, et al. 2006. Root growth dynamics of Caragana korshinskii using mini-rhizotrons. Chinese Journal of Plant Ecology, 30(3): 457-464. (in Chinese)
doi: 10.17521/cjpe.2006.0061
[39]   Zhang Z S, Chen Y L, Xu B X, et al. 2015. Topographic differentiations of biological soil crusts and hydraulic properties in fixed sand dunes, Tengger Desert. Journal of Arid Land, 7(2): 205-215.
doi: 10.1007/s40333-014-0048-y
[40]   Zhao W, Yang M X, Chen L, et al. 2015. Structure and dynamics of grassaceous layer vegetation in artificial Caragana intermediate shrubland in desert steppe. Journal of Zhejiang University: Science in Agriculture & Life, 41(6): 723-731. (in Chinese)
[41]   Zheng X R, Zhao G Q, Li X Y, et al. 2015. Application of stable hydrogen isotope in study of water sources for Caragana microphylla bushland in Nei Mongol. Chinese Journal of Plant Ecology, 39(2): 184-196. (in Chinese)
doi: 10.17521/cjpe.2015.0018
[42]   Zhou H, Zheng X J, Tan L S, et al. 2013. Differences and similarities between water sources of Tamarix ramosissima, Nitraria sibirica and Reaumuria soongorica in the southeastern Junggar Basin. Chinese Journal of Plant Ecology, 37(7): 665-673. (in Chinese)
doi: 10.3724/SP.J.1258.2013.00069
[43]   Zhu Y J, Jia Z Q, Lu Q, et al. 2010. Water use strategy of five shrubs in Ulanbuh Desert. Sciencetia Silvae Sinicae, 46(4): 15-21. (in Chinese)
[44]   Zuo X A, Zhao X Y, Zhao H L. 2009. Spatial heterogeneity of soil properties and vegetation-soil relationships following vegetation restoration of mobile dunes in Horqin Sandy Land, Northern China. Plant and Soil, 318: 153-167.
doi: 10.1007/s11104-008-9826-7
[45]   Zuo Z, Wang J L, Zhang Y P, et al. 2006. Investigates about pea shrub resource utilization and its feed exploits processing at the present situation in Ningxia-based on the Yanchi County. Pratacultural Science, 23(3): 17-22. (in Chinese)
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