Co-existence mechanisms of plant species within “soil islands” habitat of desert steppe

SONG Naiping 1 WANG Xing 1 CHEN Lin 1 XUE Yi 1 CHEN Juan 1 SUI Jinming 1 WANG Lei 1 YANG Xinguo 1

(1.Breeding Base for State Key Laboratory of Land Degradation and Ecological Restoration in Northwest China, Key Laboratory for Restoration and Reconstruction of Degraded Ecosystem in Northwest China of Ministry of Education, Ningxia University, Yinchuan 750021)

【Abstract】Extensive fragmentation has led to the original desert steppe soils of hard sierozem to become isolated in the desertified lands, forming the so-called “soil islands”. We aimed to reveal the effect of fragmentation on the co-existence of plant species within “soil island” habitat. To this end, we characterized the plant communities on three large (200–300 m2), medium (about 100 m2) and small (about 50 m2) soil islands each, in the Wanjigou Village of Yanchi County in Ningxia Hui Autonomous Region of China. We analyzed the similarity of plant communities within and outside those soil islands, the breadth and overlap of plant ecological niche, the species co-existence pattern and its influencing factors using Jaccard similarity coefficient, niche overlapping degree formula, null model analysis, and meta-analysis. Our results showed that: (1) with increased fragmentation, the plant diversity within the soil islands decreased, and the dominant plant species changed from Stipa breviflora to a combination of Artemisia scoparia and S. breviflora, and the plant communities within soil islands became more similar to communities outside; (2) most species showed relatively small ecological niche overlap between within and outside soil islands, with the overlap showing concentrated distribution within soil islands but scattered distribution outside the islands; (3) environmental filtering was the main driver of species co-existence patterns. As the soil islands became larger, the effect of environmental factors on species co-existence decreased, and the key environmental factor that explained species composition changed from clay and fine sand to coarse sand. Notably, competition was a key driver of co-existence patterns within small soil islands. In summary, soil islands provided important habitats for representative plant species in the desert steppe and environmental factors influenced species co-existence patterns. Both processes were disrupted by fragmentation. Maintaining soil islands larger than 200 m2 would be critical for restoring representative plant species and preserving plant diversity of the desert steppe ecosystems.

【Keywords】 habitat fragmentation; soil islands; niche overlap; species co-existence; desert steppe;


【Funds】 National Natural Science Foundation of China (41461046) Construction Project of Top Disciplines in Ningxia Hui Autonomous Region (NXYLXK2017B06)

Download this article


    Albrecht M, Gotelli NJ (2001) Spatial and temporal niche partitioning in grassland ants. Oecologia, 126, 134–141.

    Bao SD (2000) Soil Agro-chemistrical Analysis. China Agriculture Press, Beijing (in Chinese).

    Bergholz K, May F, Giladi I, Ristow M, Ziv Y, Jeltsch F (2017) Environmental heterogeneity drives fine-scale species assembly and functional diversity of annual plants in a semi-arid environment. Perspectives in Plant Ecology, Evolution and Systematics, 24, 138–146.

    Boeye J, Kubisch A, Bonte D (2014) Habitat structure mediates spatial segregation and therefore coexistence. Landscape Ecology, 29, 593–604.

    Borcard D, Gillet F, Legendre P (translated by Lai JS) (2014) Numerical Ecology with R, pp. 332–334. Higher Education Press, Beijing (in Chinese).

    Briers R (2006) Ecology: From individuals to ecosystems. Freshwater Biology, 51, 1787–1788.

    Brown CD, Boutin C (2009) Linking past land use, recent disturbance, and dispersal mechanism to forest composition. Biological Conservation, 142, 1647–1656.

    Chai YF, Yue M (2017) Research advances in plant community assembly mechanisms. Acta Ecologica Sinica, 36, 4557–4572 (in Chinese with English abstract).

    Chen L, Mi XC, Ma KP (2014) Niche differentiation and its consequence on biodiversity maintenance in forest communities. Chinese Bulletin of Life Sciences, 26, 112–117 (in Chinese).

    Chu CJ, Wang YS, Liu Y, Jiang L, He FL (2017) Advances in species coexistence theory. Biodiversity Science, 25, 345–354 (in Chinese with English abstract).

    Collins CD, Banks-Leite C, Brudvig LA, Foster BL, Cook WM, Damschen EI, Andrade A, Austin M, Camargo JL, Driscoll DA (2017) Fragmentation affects plant community composition over time. Ecography, 40, 119–130.

    Duan HL, Zhao A, Yao Z (2017) Analysis of wetland plant–soil relationships and population niches in Chayegang marshland near Henghu farm in the Poyang Lake region during the dry season. Acta Ecologica Sinica, 37, 3744–3754 (in Chinese with English abstract).

    Gao M, He P, Zhang X (2014) Relative roles of spatial factors, environmental filtering and biotic interactions in fine-scale structuring of a soil mite community. Soil Biology & Biochemistry, 79, 68–77.

    Gonzalez M, Ladet S, Deconchat M, Cabanettes A, Alard D, Balent G (2010) Relative contribution of edge and interior zones to patch size effect on species richness: An example for woody plants. Forest Ecology and Management, 259, 266–274.

    Gotelli NJ, McCabe DJ (2002) Species co-occurrence: A meta-analysis of JM Diamond’s assembly rules model. Ecology, 83, 2091–2096.

    Gotelli NJ, Ulrich W (2010) The empirical Bayes approach as a tool to identify non-random species associations. Oecologia, 162, 463–477.

    Haddad NM, Brudvig LA, Clobert J, Davies KF, Gonzalez A, Holt RD, Lovejoy TE, Sexton JO, Austin MP, Collins CD (2015) Habitat fragmentation and its lasting impact on Earth’s ecosystems. Science Advances, 1, e1500052.

    Han DY, Yang YF (2014) Species abundance–distribution relationship and its interpretation in plant communities on the Songnen grasslands, China. Biodiversity Science, 22, 348–357 (in Chinese with English abstract).

    Hu G, Feeley KJ, Yu M (2016) Habitat fragmentation drives plant community assembly processes across life stages. PLoS ONE, 11, e0159572.

    Hu GL, Zhao WZ, Wang G (2011) Reviews on spatial pattern and sand-binding effect of patch vegetation in arid desert area. Acta Ecologica Sinica, 31, 7609–7616 (in Chinese with English abstract).

    Jiménez JJ, Decaëns T, Rossi JP (2012) Soil environmental heterogeneity allows spatial c-occurrence of competitor earthworm species in a gallery forest of the Colombian “Llanos”. Oikos, 121, 915–926.

    Krasnov BR, Shenbrot GI, Khokhlova IS (2011) Aggregative structure is the rule in communities of fleas: Null model analysis. Ecography, 34, 751–761.

    Li XR, Zhao Y, Hui R, Su JQ, Gao YH (2014) Progress and trend of development of restoration ecology research in the arid regions of China. Progress in Geography, 33, 1435–1443 (in Chinese with English abstract).

    Ma KM, Zhang JY, Guo XD, Fu BJ (2002) Plant diversity distribution of mountains in rural landscapes: The combined influences of topography and land use. Acta Phytoecologica Sinica, 26, 575–588 (in Chinese with English abstract).

    May R, Mclean A (1976) Theoretical Ecology: Principles and Applications, pp. 1147–1152. Oxford University Press, New York.

    Miguel JM, Martín-Forés I, Acosta-Gallo B, del Pozo A, Ovalle C, Sánchez-Jardón L, Castro I, Casado MA (2016) Non-random co-occurrence of native and exotic plant species in Mediterranean grasslands. Acta Oecologica, 77, 18–26.

    Patiño J, Whittaker RJ, Borges PAV, Fernández-Palacios JM, Ah-Peng C, Araújo MB, Ávila SP, Cardoso P, Cornuault J, de Boer EJ, de Nascimento L, Gil A, GonẐalez-Castro A, Gruner DS, Heleno R, Horta J, Illera GJ, Kaiser-Bunbury CN, Matthews TJ, Papadopoulou A, Pettorelli N, Price JP, Santos AMC, Steinbauer MJ, Triantis KA, Valente L, Vargas P, Weigelt P, Emerson BC (2017) A roadmap for island biology: 50 fundamental questions after 50 years of The Theory of Island Biogeography. Journal of Biogeography, 44, 963–983.

    Shang KK, Da LJ (2013) Microtopographic spatial differentiation pattern and coexistence mechanism of relict deciduous broadleaved trees: A review. Chinese Journal of Ecology, 32, 1912–1919 (in Chinese with English abstract).

    Walter H (1985) Vegetation of the Earth and Ecological Systems of the Geo-biosphere. Springer-Verlag, New York.

    Wu J, Yu XC, Liu ZM, Hou XZ, Qian JQ, Xing BZ (2017) Effects of “habitat island” area on plant species diversity and its distribution pattern in Horqin Sand Land. Chinese Journal of Ecology, 36, 43–53 (in Chinese with English abstract).

    Zhang JT (2004) Numerical Ecology. Science Press, Beijing (in Chinese).

    Zhang Q (2011) Biodiversity Patterns and Environmental Interpretation of Stipa breviflora Grassland in Inner Mongolia. PhD dissertation, Inner Mongolia University, Hohhot (in Chinese with English abstract).

    Zhou SR, Zhang DY (2006) Neutral theory in community ecology. Journal of Plant Ecology (Chinese Version), 30, 868–877 (in Chinese with English abstract).

    Zou XH (2015) Ecological Effects and Dynamic Mechanisms of Vegetattion Patches in Alkali-Saline Grassland Landscape on the Songnen Plains. PhD dissertation, Northeast Normal University, Changchun (in Chinese with English abstract).

This Article



Vol 26, No. 07, Pages 667-677

July 2018


Article Outline


  • 1 Materials and methods
  • 2 Results
  • 3 Discussion
  • Supplementary material
  • References