Climate gradients are associated with divergent fungal and bacterial assembly processes and soil multifunctionality in a central Asian dryland

Abstract

Drylands are crucial to global biogeochemical cycles, yet the linkages between soil multifunctionality (SMF) and microbial diversity, community assembly, and network architecture remain poorly understood. Here, we investigated these relationships across three grassland types—desert steppe, temperate desert, and temperate steppe—in the drylands of Kazakhstan. Despite contrasting environmental conditions, microbial α diversity did not differ significantly among grassland types and was more strongly associated with climatic factors than with grassland type per se. Fungal and bacterial communities exhibited distinct assembly patterns. Fungal communities were governed by dispersal limitation (57.46%–57.69%) and homogeneous selection (13.16%–16.40%), with community similarity correlated with both environmental and geographic distances. In contrast, bacterial communities were shaped predominantly by ecological drift (40.38%–42.88%), and their similarity was linked primarily to geographic distance. Microbial co-occurrence network also varied systematically across grassland types, shifting from more modular structures in desert steppe to more connected configurations in temperate desert and temperate steppe. Structural equation modeling showed that climatic variables, particularly mean annual temperature, exerted the strongest direct effects on SMF, whereas community assembly processes influenced SMF only indirectly through their regulation of microbial diversity. Overall, climatic gradients, rather than microbial richness per se, emerged as the primary correlates of SMF, with fungal and bacterial communities exhibiting divergent assembly processes and network organization along the aridity gradient. These findings provide an integrative, association-based framework for understanding how environmental gradients related to aridification correspond to microbial ecological organization and SMF in drylands.

Graphical abstract

Keywords

microbial diversity / microbial community assembly / co-occurrence network / soil multifunctionality / dryland ecosystems

Highlight

	● Climatic variables most strongly associate with SMF in Kazakh drylands.
	● Microbial alpha diversity varied little among desert steppe (DS), temperate desert (TD), and temperate steppe (TS).
	● Fungal and bacterial communities exhibit contrasting assembly processes across grassland types.
	● Microbial networks shift from modular (DS) to connected (TD and TS) configurations.
	● SMF variation primarily associates with climatic background and spatially structured microbial organization.

References

[1] Aguilar, P., Sommaruga, R., 2020. The balance between deterministic and stochastic processes in structuring lake bacterioplankton community over time. Molecular Ecology29, 3117–3130.

[2] Aitchison, J., 1982. The statistical analysis of compositional data. Journal of the Royal Statistical Society Series B: Statistical Methodology44, 139–160.

[3] Bahram, M., Hildebrand, F., Forslund, S.K., Anderson, J.L., Soudzilovskaia, N.A., Bodegom, P.M., Bengtsson-Palme, J., Anslan, S., Coelho, L.P., Harend, H., Huerta-Cepas, J., Medema, M.H., Maltz, M.R., Mundra, S., Olsson, P.A., Pent, M., Põlme, S., Sunagawa, S., Ryberg, M., Tedersoo, L., Bork, P., 2018. Structure and function of the global topsoil microbiome. Nature560, 233–237.

[4] Benjamini, Y., Yekutieli, D., 2001. The control of the false discovery rate in multiple testing under dependency. Annals of Statistics29, 1165–1188.

[5] Berdugo, M., Delgado-Baquerizo, M., Soliveres, S., Hernández-Clemente, R., Zhao, Y.C., Gaitán, J.J., Gross, N., Saiz, H., Maire, V., Lehmann, A., Rillig, M.C., Solé, R.V., Maestre, F.T., 2020. Global ecosystem thresholds driven by aridity. Science367, 787–790.

[6] Bhagwat, T., Kuemmerle, T., Soofi, M., Donald, P.F., Hölzel, N., Salemgareev, A., Stirnemann, I., Urazaliyev, R., Baumann, M., Kamp, J., 2024. A novel, post-Soviet fire disturbance regime drives bird diversity and abundance on the Eurasian steppe. Global Change Biology30, e17026.

[7] Blondel, V.D., Guillaume, J.L., Lambiotte, R., Lefebvre, E., 2008. Fast unfolding of communities in large networks. Journal of Statistical Mechanics: Theory and Experiment2008, P10008.

[8] Bowman, E.A., Arnold, A.E., 2021. Drivers and implications of distance decay differ for ectomycorrhizal and foliar endophytic fungi across an anciently fragmented landscape. The ISME Journal15, 3437–3454.

[9] Bremner, J.M., Mulvaney, C.S., 1982. Nitrogen—total. In: Page, A.L., ed. Methods of Soil Analysis: Part 2 Chemical and Microbiological Properties. 2nd ed. Madison: American Society of Agronomy and Soil Science Society of America, 595–624.

[10] Chase, J.M., Myers, J.A., 2011. Disentangling the importance of ecological niches from stochastic processes across scales. Philosophical Transactions of the Royal Society B: Biological Sciences366, 2351–2363.

[11] Chen, Q.L., Ding, J., Zhu, D., Hu, H.W., Delgado-Baquerizo, M., Ma, Y.B., He, J.Z., Zhu, Y.G., 2020. Rare microbial taxa as the major drivers of ecosystem multifunctionality in long-term fertilized soils. Soil Biology and Biochemistry141, 107686.

[12] Chen, R.R., Wang, X.T., 2022. BOOK REVIEW: Analytical methods for soil and agro-chemistry (in Chinese). Edited by Zhu, H. Z., He, P. A., Chen, C. Z., Zhou, H. M., Su, D. C., Xu, J. M., Qin, H. Y., Bao, S. D., Lu, R. K. Jiang, S. H. Soil Science Society of China Beijing, China Agricultural Science and Technology Press, 2000, pp. 638. ISBN: 9787801199256. European Journal of Soil Science73( e13280), .

[13] Chen, S.T., Cao, P.X., Li, T., Wang, Y.Y., Liu, X., 2023a. Microbial diversity patterns in the root zone of two Meconopsis plants on the Qinghai-Tibet Plateau. PeerJ11, e15361.

[14] Chen, W.Q., Wang, J.Y., Chen, X., Meng, Z.X., Xu, R., Duoji, D., Zhang, J.H., He, J., Wang, Z.G., Chen, J., Liu, K.X., Hu, T.M., Zhang, Y.J., 2022a. Soil microbial network complexity predicts ecosystem function along elevation gradients on the Tibetan Plateau. Soil Biology and Biochemistry172, 108766.

[15] Chen, X., Li, Q., Chen, D.D., He, F.Q., Huo, L.L., Zhao, L., Xiao, C.W., 2022b. Reasonable management of perennial planting grassland contributes to positive succession of soil microbial community in Sanjiangyuan of Qinghai-Tibetan Plateau, China. Journal of Plant Ecology15, 359–371.

[16] Chen, X.D., Li, H., Condron, L.M., Dunfield, K.E., Wakelin, S.A., Mitter, E.K., Jiang, N., 2023b. Long-term afforestation enhances stochastic processes of bacterial community assembly in a temperate grassland. Geoderma430, 116317.

[17] Coyte, K.Z., Schluter, J., Foster, K.R., 2015. The ecology of the microbiome: networks, competition, and stability. Science350, 663–666.

[18] de Beurs, K.M., Henebry, G.M., 2004. Land surface phenology, climatic variation, and institutional change: analyzing agricultural land cover change in Kazakhstan. Remote Sensing of Environment89, 497–509.

[19] de Menezes, A.B., Prendergast-Miller, M.T., Richardson, A.E., Toscas, P., Farrell, M., Macdonald, L.M., Baker, G., Wark, T., Thrall, P.H., 2015. Network analysis reveals that bacteria and fungi form modules that correlate independently with soil parameters. Environmental Microbiology17, 2677–2689.

[20] de Menezes, A.B., Richardson, A.E., Thrall, P.H., 2017. Linking fungal-bacterial co-occurrences to soil ecosystem function. Current Opinion in Microbiology37, 135–141.

[21] de Vries, F.T., Griffiths, R.I., Bailey, M., Craig, H., Girlanda, M., Gweon, H.S., Hallin, S., Kaisermann, A., Keith, A.M., Kretzschmar, M., Lemanceau, P., Lumini, E., Mason, K.E., Oliver, A., Ostle, N., Prosser, J.I., Thion, C., Thomson, B., Bardgett, R.D., 2018. Soil bacterial networks are less stable under drought than fungal networks. Nature Communications9, 3033.

[22] Delgado-Baquerizo, M., Maestre, F.T., Reich, P.B., Jeffries, T.C., Gaitan, J.J., Encinar, D., Berdugo, M., Campbell, C.D., Singh, B.K., 2016. Microbial diversity drives multifunctionality in terrestrial ecosystems. Nature Communications7, 10541.

[23] Delgado-Baquerizo, M., Reich, P.B., Trivedi, C., Eldridge, D.J., Abades, S., Alfaro, F.D., Bastida, F., Berhe, A.A., Cutler, N.A., Gallardo, A., García-Velázquez, L., Hart, S.C., Hayes, P.E., He, J.Z., Hseu, Z.Y., Hu, H.W., Kirchmair, M., Neuhauser, S., Pérez, C.A., Reed, S.C., Santos, F., Sullivan, B.W., Trivedi, P., Wang, J.T., Weber-Grullon, L., Williams, M.A., Singh, B.K., 2020. Multiple elements of soil biodiversity drive ecosystem functions across biomes. Nature Ecology & Evolution4, 210–220.

[24] Dixon, P., 2003. VEGAN, a package of r functions for community ecology. Journal of Vegetation Science14, 927–930.

[25] Dray, S., Legendre, P., Peres-Neto, P.R., 2006. Spatial modelling: a comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM). Ecological Modelling196, 483–493.

[26] Feng, K., Zhang, Z.J., Cai, W.W., Liu, W.Z., Xu, M.Y., Yin, H.Q., Wang, A.J., He, Z.L., Deng, Y., 2017. Biodiversity and species competition regulate the resilience of microbial biofilm community. Molecular Ecology26, 6170–6182.

[27] Fierer, N., Jackson, R.B., 2006. The diversity and biogeography of soil bacterial communities. Proceedings of the National Academy of Sciences of the United States of America103, 626–631.

[28] Friedman, J., Alm, E.J., 2012. Inferring correlation networks from genomic survey data. PLoS Computational Biology8, e1002687.

[29] Gardner, W.H., 1986. Water content. In: Klute, A., ed. Methods of Soil Analysis: Part 1 Physical and Mineralogical Methods. 2nd ed. Madison: American Society of Agronomy and Soil Science Society of America, 493–544.

[30] Gloor, G.B., Macklaim, J.M., Pawlowsky-Glahn, V., Egozcue, J.J., 2017. Microbiome datasets are compositional: and this is not optional. Frontiers in Microbiology8, 2224.

[31] Gong, H.Y., Wang, H.X., Wang, Y., Zhang, S., Liu, X., Che, J.C., Wu, S., Wu, J., Sun, X.M., Zhang, S.G., Yau, S.T., Wu, R.L., 2024. Topological change of soil microbiota networks for forest resilience under global warming. Physics of Life Reviews50, 228–251.

[32] Green, J.L., Holmes, A.J., Westoby, M., Oliver, I., Briscoe, D., Dangerfield, M., Gillings, M., Beattie, A.J., 2004. Spatial scaling of microbial eukaryote diversity. Nature432, 747–750.

[33] Guo, G.X., Kong, W.D., Liu, J.B., Zhao, J.X., Du, H.D., Zhang, X.Z., Xia, P.H., 2015. Diversity and distribution of autotrophic microbial community along environmental gradients in grassland soils on the Tibetan Plateau. Applied Microbiology and Biotechnology99, 8765–8776.

[34] Guo, X., Feng, J.J., Shi, Z., Zhou, X.S., Yuan, M.T., Tao, X.Y., Hale, L., Yuan, T., Wang, J.J., Qin, Y.J., Zhou, A.F., Fu, Y., Wu, L.Y., He, Z.L., Van Nostrand, J.D., Ning, D.L., Liu, X.D., Luo, Y.Q., Tiedje, J.M., Yang, Y.F., Zhou, J.Z., 2018. Climate warming leads to divergent succession of grassland microbial communities. Nature Climate Change8, 813–818.

[35] Guo, Z.H., Huang, X.B., Wang, T.L., Su, J.R., Li, S.F., 2024. Altitudinal influences on soil microbial diversity and community assembly in topsoil and subsoil layers: insights from the Jinsha River Basin, Southwest China. Forests15, 1746.

[36] Hanson, C.A., Fuhrman, J.A., Horner-Devine, M.C., Martiny, J.B.H., 2012. Beyond biogeographic patterns: processes shaping the microbial landscape. Nature Reviews Microbiology10, 497–506.

[37] Hernandez, D.J., David, A.S., Menges, E.S., Searcy, C.A., Afkhami, M.E., 2021. Environmental stress destabilizes microbial networks. The ISME Journal15, 1722–1734.

[38] Hooper, D.U., Adair, E.C., Cardinale, B.J., Byrnes, J.E.K., Hungate, B.A., Matulich, K.L., Gonzalez, A., Duffy, J.E., Gamfeldt, L., O’Connor, M.I., 2012. A global synthesis reveals biodiversity loss as a major driver of ecosystem change. Nature486, 105–108.

[39] Hu, W.G., Ran, J.Z., Dong, L.W., Du, Q.J., Ji, M.F., Yao, S.R., Sun, Y., Gong, C.M., Hou, Q.Q., Gong, H.Y., Chen, R.F., Lu, J.L., Xie, S.B., Wang, Z.Q., Huang, H., Li, X.W., Xiong, J.L., Xia, R., Wei, M.H., Zhao, D.M., Zhang, Y.H., Li, J.H., Yang, H.X., Wang, X.T., Deng, Y., Sun, Y., Li, H.L., Zhang, L., Chu, Q.P., Li, X.W., Aqeel, M., Manan, A., Akram, M.A., Liu, X.H., Li, R., Li, F., Hou, C., Liu, J.Q., He, J.S., An, L.Z., Bardgett, R.D., Schmid, B., Deng, J.M., 2021. Aridity-driven shift in biodiversity-soil multifunctionality relationships. Nature Communications12, 5350.

[40] Isobe, K., Allison, S.D., Khalili, B., Martiny, A.C., Martiny, J.B.H., 2019. Phylogenetic conservation of bacterial responses to soil nitrogen addition across continents. Nature Communications10, 2499.

[41] Jousset, A., Bienhold, C., Chatzinotas, A., Gallien, L., Gobet, A., Kurm, V., Küsel, K., Rillig, M.C., Rivett, D.W., Salles, J.F., van der Heijden, M.G.A., Youssef, N.H., Zhang, X.W., Wei, Z., Hol, W.H.G., 2017. Where less may be more: how the rare biosphere pulls ecosystems strings. The ISME Journal11, 853–862.

[42] Kurtz, Z.D., Müller, C.L., Miraldi, E.R., Littman, D.R., Blaser, M.J., Bonneau, R.A., 2015. Sparse and compositionally robust inference of microbial ecological networks. PLoS Computational Biology11, e1004226.

[43] Lauber, C.L., Strickland, M.S., Bradford, M.A., Fierer, N., 2008. The influence of soil properties on the structure of bacterial and fungal communities across land-use types. Soil Biology and Biochemistry40, 2407–2415.

[44] Lefcheck, J.S., 2016. PIECEWISESEM: piecewise structural equation modelling in R for ecology, evolution, and systematics. Methods in Ecology and Evolution7, 573–579.

[45] Li, J., Wang, X., Yuan, M.H., Duan, W.H., Xia, J.Y., Zhang, X.S., Zhao, Y.F., Wang, J.W., 2025. Effect of soil microbial community on ecosystem multifunctionality in an alpine grassland. Catena249, 108714.

[46] Liu, C., Cui, Y.M., Li, X.Z., Yao, M.J., 2021. microeco: an R package for data mining in microbial community ecology. FEMS Microbiology Ecology97, fiaa255.

[47] Liu, H., Ren, F.R., Wan, S.Q., Han, S.J., Zheng, J.Q., 2025. Nitrogen and water additions with or without mowing altered soil microbial community characteristics in a semi-arid steppe. Ecological Processes14, 3.

[48] Liu, Y.X., Chen, L., Ma, T.F., Li, X.F., Zheng, M.S., Zhou, X., Chen, L., Qian, X.B., Xi, J., Lu, H.Y., Cao, H.L., Ma, X.Y., Bian, B., Zhang, P.F., Wu, J.Q., Gan, R.Y., Jia, B.L., Sun, L.Y., Ju, Z.C., Gao, Y.Y., Wen, T., Chen, T., 2023. EasyAmplicon: an easy-to-use, open-source, reproducible, and community-based pipeline for amplicon data analysis in microbiome research. iMeta2, e83.

[49] Loreau, M., Hector, A., 2001. Partitioning selection and complementarity in biodiversity experiments. Nature412, 72–76.

[50] Ma, B., Wang, H.Z., Dsouza, M., Lou, J., He, Y., Dai, Z.M., Brookes, P.C., Xu, J.M., Gilbert, J.A., 2016. Geographic patterns of co-occurrence network topological features for soil microbiota at continental scale in eastern China. The ISME Journal10, 1891–1901.

[51] Ma, B., Wang, Y.L., Ye, S.D., Liu, S., Stirling, E., Gilbert, J.A., Faust, K., Knight, R., Jansson, J.K., Cardona, C., Röttjers, L., Xu, J.M., 2020. Earth microbial co-occurrence network reveals interconnection pattern across microbiomes. Microbiome8, 82.

[52] Ma, L.N., Zhang, C.X., Xu, X.F., Wang, C.W., Liu, G.F., Liang, C.Z., Zuo, X.A., Wang, C.J., Lv, Y.X., Wang, R.Z., 2022. Different facets of bacterial and fungal communities drive soil multifunctionality in grasslands spanning a 3500 km transect. Functional Ecology36, 3120–3133.

[53] Martiny, A.C., Treseder, K., Pusch, G., 2013. Phylogenetic conservatism of functional traits in microorganisms. The ISME Journal7, 830–838.

[54] Martiny, J.B.H., Bohannan, B.J.M., Brown, J.H., Colwell, R.K., Fuhrman, J.A., Green, J.L., Horner-Devine, M.C., Kane, M., Krumins, J.A., Kuske, C.R., Morin, P.J., Naeem, S., Øvreås, L., Reysenbach, A.L., Smith, V.H., Staley, J.T., 2006. Microbial biogeography: putting microorganisms on the map. Nature Reviews Microbiology4, 102–112.

[55] Nekola, J.C., White, P.S., 1999. The distance decay of similarity in biogeography and ecology. Journal of Biogeography26, 867–878.

[56] Newman, M.E.J., 2006. Modularity and community structure in networks. Proceedings of the National Academy of Sciences of the United States of America103, 8577–8582.

[57] Ning, D.L., Yuan, M.T., Wu, L.W., Zhang, Y., Guo, X., Zhou, X.S., Yang, Y.F., Arkin, A.P., Firestone, M.K., Zhou, J.Z., 2020. A quantitative framework reveals ecological drivers of grassland microbial community assembly in response to warming. Nature Communications11, 4717.

[58] Olesen, J.M., Bascompte, J., Dupont, Y.L., Jordano, P., 2007. The modularity of pollination networks. Proceedings of the National Academy of Sciences of the United States of America104, 19891–19896.

[59] Öpik, M., Davison, J., Moora, M., Zobel, M., 2014. DNA-based detection and identification of Glomeromycota: the virtual taxonomy of environmental sequences. Botany92, 135–147.

[60] Peter, H., Beier, S., Bertilsson, S., Lindström, E.S., Langenheder, S., Tranvik, L.J., 2011. Function-specific response to depletion of microbial diversity. The ISME Journal5, 351–361.

[61] Prăvălie, R., 2016. Drylands extent and environmental issues. A global approach. Earth-Science Reviews161, 259–278.

[62] Puente-Sánchez, F., Pascual-García, A., Bastolla, U., Pedrós-Alió, C., Tamames, J., 2024. Cross-biome microbial networks reveal functional redundancy and suggest genome reduction through functional complementarity. Communications Biology7, 1046.

[63] Pyarali, K., Peng, J., Disse, M., Tuo, Y., 2022. Development and application of high resolution SPEI drought dataset for Central Asia. Scientific Data9, 172.

[64] Qiu, L.P., Zhang, Q., Zhu, H.S., Reich, P.B., Banerjee, S., van der Heijden, M.G.A., Sadowsky, M.J., Ishii, S., Jia, X.X., Shao, M.A., Liu, B.Y., Jiao, H., Li, H.Q., Wei, X.R., 2021. Erosion reduces soil microbial diversity, network complexity and multifunctionality. The ISME Journal15, 2474–2489.

[65] Riddley, M., Hepp, S., Hardeep, F., Nayak, A., Liu, M.M., Xing, X., Zhang, H.L., Liao, J.Q., 2025. Differential roles of deterministic and stochastic processes in structuring soil bacterial ecotypes across terrestrial ecosystems. Nature Communications16, 2337.

[66] Salnikov, V., Talanov, Y., Polyakova, S., Assylbekova, A., Kauazov, A., Bultekov, N., Musralinova, G., Kissebayev, D., Beldeubayev, Y., 2023. An assessment of the present trends in temperature and precipitation extremes in Kazakhstan. Climate11, 33.

[67] Shade, A., Peter, H., Allison, S.D., Baho, D., Berga, M., Buergmann, H., Huber, D.H., Langenheder, S., Lennon, J.T., Martiny, J.B., Matulich, K.L., Schmidt, T.M., Handelsman, J., 2012. Fundamentals of microbial community resistance and resilience. Frontiers in Microbiology3, 417.

[68] Shen, L., Stopher, P.R., 2014. Review of GPS travel survey and GPS data-processing methods. Transport Reviews34, 316–334.

[69] Shipley, B., 2009. Confirmatory path analysis in a generalized multilevel context. Ecology90, 363–368.

[70] Stegen, J.C., Lin, X.J., Fredrickson, J.K., Konopka, A.E., 2015. Estimating and mapping ecological processes influencing microbial community assembly. Frontiers in Microbiology6, 370.

[71] Steven, B., McHugh, T.A., Reed, S., 2017. The response of arid soil communities to climate change. In: Steven, B., ed. The Biology of Arid Soils. Berlin, Boston: De Gruyter, 139–158.

[72] Stone, B.W., Li, J.H., Koch, B.J., Blazewicz, S.J., Dijkstra, P., Hayer, M., Hofmockel, K.S., Liu, X.J.A., Mau, R.L., Morrissey, E.M., Pett-Ridge, J., Schwartz, E., Hungate, B.A., 2021. Nutrients cause consolidation of soil carbon flux to small proportion of bacterial community. Nature Communications12, 3381.

[73] Tedersoo, L., Bahram, M., Põlme, S., Kõljalg, U., Yorou, N.S., Wijesundera, R., Ruiz, L.V., Vasco-Palacios, A.M., Thu, P.Q., Suija, A., Smith, M.E., Sharp, C., Saluveer, E., Saitta, A., Rosas, M., Riit, T., Ratkowsky, D., Pritsch, K., Põldmaa, K., Piepenbring, M., Phosri, C., Peterson, M., Parts, K., Pärtel, K., Otsing, E., Nouhra, E., Njouonkou, A.L., Nilsson, R.H., Morgado, L.N., Mayor, J., May, T.W., Majuakim, L., Lodge, D.J., Lee, S.S., Larsson, K.H., Kohout, P., Hosaka, K., Hiiesalu, I., Henkel, T.W., Harend, H., Guo, L.D., Greslebin, A., Grelet, G., Geml, J., Gates, G., Dunstan, W., Dunk, C., Drenkhan, R., Dearnaley, J., De Kesel, A., Dang, T., Chen, X., Buegger, F., Brearley, F.Q., Bonito, G., Anslan, S., Abell, S., Abarenkov, K., 2014. Global diversity and geography of soil fungi. Science346, 1256688.

[74] Tedersoo, L., Mikryukov, V., Anslan, S., Bahram, M., Khalid, A.N., Corrales, A., Agan, A., Vasco-Palacios, A.M., Saitta, A., Antonelli, A., Rinaldi, A.C., Verbeken, A., Sulistyo, B.P., Tamgnoue, B., Furneaux, B., Ritter, C.D., Nyamukondiwa, C., Sharp, C., Marín, C., Dai, D.Q., Gohar, D., Sharmah, D., Biersma, E.M., Cameron, E.K., De Crop, E., Otsing, E., Davydov, E.A., Albornoz, F.E., Brearley, F.Q., Buegger, F., Gates, G., Zahn, G., Bonito, G., Hiiesalu, I., Hiiesalu, I., Zettur, I., Barrio, I.C., Pärn, J., Heilmann-Clausen, J., Ankuda, J., Kupagme, J.Y., Sarapuu, J., Maciá-Vicente, J.G., Fovo, J.D., Geml, J., Alatalo, J.M., Alvarez-Manjarrez, J., Monkai, J., Põldmaa, K., Runnel, K., Adamson, K., Bråthen, K.A., Pritsch, K., Tchan, K.I., Armolaitis, K., Hyde, K.D., Newsham, K.K., Panksep, K., Adebola, L.A., Lamit, L.J., Saba, M., Cáceres, M.E.d.S., Tuomi, M., Gryzenhout, M., Bauters, M., Bálint, M., Wijayawardene, N., Hagh-Doust, N., Yorou, N.S., Kurina, O., Mortimer, P.E., Meidl, P., Nilsson, R.H., Puusepp, R., Casique-Valdés, R., Drenkhan, R., Garibay-Orijel, R., Godoy, R., Alfarraj, S., Rahimlou, S., Põlme, S., Dudov, S.V., Mundra, S., Ahmed, T., Netherway, T., Henkel, T.W., Roslin, T., Fedosov, V.E., Onipchenko, V.G., Yasanthika, W.A.E., Lim, Y.W., Piepenbring, M., Klavina, D., Kõljalg, U., Abarenkov, K., 2021. The Global Soil Mycobiome consortium dataset for boosting fungal diversity research. Fungal Diversity111, 573–588.

[75] Tedersoo, L., Sánchez-Ramírez, S., Kõljalg, U., Bahram, M., Döring, M., Schigel, D., May, T., Ryberg, M., Abarenkov, K., 2018. High-level classification of the fungi and a tool for evolutionary ecological analyses. Fungal Diversity90, 135–159.

[76] Tripathi, B.M., Stegen, J.C., Kim, M., Dong, K., Adams, J.M., Lee, Y.K., 2018. Soil pH mediates the balance between stochastic and deterministic assembly of bacteria. The ISME Journal12, 1072–1083.

[77] Wagg, C., Bender, S.F., Widmer, F., van der Heijden, M.G.A., 2014. Soil biodiversity and soil community composition determine ecosystem multifunctionality. Proceedings of the National Academy of Sciences of the United States of America111, 5266–5270.

[78] Wagg, C., Schlaeppi, K., Banerjee, S., Kuramae, E.E., van der Heijden, M.G.A., 2019. Fungal-bacterial diversity and microbiome complexity predict ecosystem functioning. Nature Communications10, 4841.

[79] Walkley, A., Black, I.A., 1934. An examination of the Degtjareff method for determining soil organic matter, and a proposed modification of the chromic acid titration method. Soil Science37, 29–38.

[80] Wang, H.X., Tian, D., Liu, H.Q., Wang, Z.Y., He, Y.J., Lu, J., Zhu, Y.L., Wei, S.X., Wang, H.L., Wu, L.C., Chen, L.J., 2024. Rare rather than abundant taxa of soil bacteria and fungi regulate soil multifunctionality in Eucalyptus plantations. CATENA245, 108303.

[81] Wang, J.M., Zhang, T.H., Li, L.P., Li, J.W., Feng, Y.M., Lu, Q., 2017a. The patterns and drivers of bacterial and fungal β-diversity in a typical dryland ecosystem of Northwest China. Frontiers in Microbiology8, 2126.

[82] Wang, J.N., Pan, Z., Yu, J.S., Zhang, Z., Li, Y.Z., 2023. Global assembly of microbial communities. mSystems8, e01289–22.

[83] Wang, X.B., Lü, X.T., Yao, J., Wang, Z.W., Deng, Y., Cheng, W.X., Zhou, J.Z., Han, X.G., 2017b. Habitat-specific patterns and drivers of bacterial β-diversity in China’s drylands. The ISME Journal11, 1345–1358.

[84] Wang, X.B., Van Nostrand, J.D., Deng, Y., Lü, X.T., Wang, C., Zhou, J.Z., Han, X.G., 2015. Scale-dependent effects of climate and geographic distance on bacterial diversity patterns across northern China’s grasslands. FEMS Microbiology Ecology91, fiv133.

[85] Yang, L.Y., Ning, D.L., Yang, Y.F., He, N.P., Li, X.Z., Cornell, C.R., Bates, C.T., Filimonenko, E., Kuzyakov, Y., Zhou, J.Z., Yu, G.R., Tian, J., 2022. Precipitation balances deterministic and stochastic processes of bacterial community assembly in grassland soils. Soil Biology and Biochemistry168, 108635.

[86] Yu, Y.Y., Shi, Y., Li, M.M., Wang, C., Zhang, L.L., Sun, Z.J., Lei, B.H., Miao, Y., Wang, W.J., Liu, B.H., Zheng, J.Q., Chu, H.Y., Han, S.J., 2022. Land-use type strongly affects soil microbial community assembly process and inter-kingdom co-occurrence pattern in a floodplain ecosystem. Applied Soil Ecology179, 104574.

[87] Yuan, Z.Q., Ali, A., Ruiz-Benito, P., Jucker, T., Mori, A.S., Wang, S.P., Zhang, X.K., Li, H., Hao, Z.Q., Wang, X.G., Loreau, M., 2020. Above- and below-ground biodiversity jointly regulate temperate forest multifunctionality along a local-scale environmental gradient. Journal of Ecology108, 2012–2024.

[88] Zhang, B.G., Wu, X.K., Zhang, G.S., Zhang, W., Liu, G.X., Chen, T., Qin, Y., Zhang, B.L., Sun, L.K., 2017. Response of soil bacterial community structure to permafrost degradation in the upstream regions of the Shule River Basin, Qinghai–Tibet Plateau. Geomicrobiology Journal34, 300–308.

[89] Zhang, S.H., Chen, Y.S., Zhou, X.B., Zhu, B., 2024a. Spatial patterns and drivers of ecosystem multifunctionality in China: arid vs. humid regions. Science of the Total Environment920, 170868.

[90] Zhang, T.Y., Liu, Y.D., Zimini, Y., Yuan, L.H., Wen, Z.M., 2024b. Effects of soil moisture and atmospheric vapor pressure deficit on the temporal variability of productivity in Eurasian grasslands. Remote Sensing16, 2368.

[91] Zhang, Z.Q., Lu, Y.H., Wei, G.H., Jiao, S., 2022. Rare species-driven diversity-ecosystem multifunctionality relationships are promoted by stochastic community assembly. mBio13, e00449–22.

[92] Zhou, J.Z., Ning, D.L., 2017. Stochastic community assembly: does it matter in microbial ecology?. Microbiology and Molecular Biology Reviews81, e00002-17.

Rights & permissions

Higher Education Press