Abstract
Urbanization reshapes soil ecosystems, yet its impacts on the composition and interactions of soil microbial communities remain insufficiently understood. Here, we use high-throughput sequencing to profile bacteria, fungi, and protists across four representative habitat types of different urbanization level—campuses, roadside greenbelts, suburban unmanaged lands, and concrete pavements in four Chinese cities spanning a broad latitudes (Changchun, Beijing, Hangzhou, Haikou). Vegetated habitats consistently structure microbial communities, whereas concrete pavements form a distinct disturbed habitat with the lowest microbial richness. Bacterial richness is highest in campuses and roadside greenbelts, while fungal and protistan richness peaks in suburban unmanaged lands. Despite regional variation, communities under similar vegetation types show convergent compositions, jointly shaped by regional climate and land-use intensity. Bacterial assemblages are more sensitive to environmental variation than fungi and protists. Proteobacteria dominate bacteria communities, whereas fungal and protistan communities display marked habitat specificity. Co-occurrence networks complexity peaks in roadside greenbelts, remains stable in suburban unmanaged lands and collapses significantly in concrete pavements. Microbial diversity patterns are strongly associated with temperature, soil pH, and latitude. Together, these results demonstrate that environmental conditions and urbanization intensity drive the biogeography and interaction patterns of soil microbiomes, providing insights for ecological restoration and urban green-space management.
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Data availability
The raw sequencing data are available in the NCBI Sequence Read Archive (SRA) (http://trace.ncbi.nlm.nih.gov/Traces/sra/) under accession number PRJNA1309871 for roadside greenbelts, PRJNA1309878 for suburban unmanaged lands, PRJNA1309853 for concrete pavements, and PRJNA1309333 for urban campus. Processed data and analysis inputs are available in the Zenodo repository at https://doi.org/10.5281/zenodo.1881998271 and https://doi.org/10.5281/zenodo.1910416472. In addition, Supplementary Data 1 contains environmental, climatic, microbial diversity, and urbanization-related variables used in this study. Supplementary Data 2 contains OTU abundance matrices of bacterial, fungal, and protistan communities with taxonomic annotations. Supplementary Data 3 provides the source data underlying all figures and charts in the main manuscript. Any remaining information can be obtained from the corresponding author upon reasonable request.
Code availability
All custom R scripts used for data processing, statistical analyses, microbial network construction, and structural equation modelling in this study are publicly available on Zenodo at https://doi.org/10.5281/zenodo.18819982. Analyses were performed in R (v4.3.0), using packages including vegan, randomForest, ggplot2, igraph, piecewiseSEM, dplyr, and ggpubr. No restrictions apply to the access or use of the code.
References
Banerjee, S. & van der Heijden, M. G. A. Soil microbiomes and one health. Nat. Rev. Microbiol. 21, 6–20 (2023).
Ferreira, C. S. S., Pereira, P. & Kalantari, Z. Human impacts on soil. Sci. total Environ. 644, 830–834 (2018).
Fu, B. et al. Urbanization alters soil bacterial communities in southern China coastal cities. Ecotox. Environ. Safe. 250, 114492 (2023).
Jagadesh, M. et al. Revealing the hidden world of soil microbes: metagenomic insights into plant, bacteria, and fungi interactions for sustainable agriculture and ecosystem restoration. Microbiol. Res. 285, 127764 (2024).
Pecsi, E. L., Forbes, S. & Guillemette, F. Organic matter composition as a driver of soil bacterial responses to pig carcass decomposition in a canadian continental climate. J. geophys. Res. Biogeo. 129, e2024JG008355 (2024).
Hawkins, H. J. et al. Mycorrhizal mycelium as a global carbon pool. Curr. Biol.: CB 33, R560–R573 (2023).
Pauwels, R., Graefe, J. & Bitterlich, M. An arbuscular mycorrhizal fungus alters soil water retention and hydraulic conductivity in a soil texture specific way. Mycorrhiza 33, 165–179 (2023).
Biswas, J. K., Pramanik, S. & Kumar, M. Fish parasites as proxy bioindicators of degraded water quality of River Saraswati, India. Environ. Monit. Assess. 195, 818 (2023).
Zhao, D. et al. The influence of different types of urban land use on soil microbial biomass and functional diversity in Beijing China. Soil Use Manag. 29, 230–239 (2013).
Zhan, Y. et al. Urbanization can accelerate climate change by increasing soil N2O emission while reducing CH4 uptake. Glob. Change Biol. 29, 3489–3502 (2023).
Cunha, G. K. G. et al. Urbanization increases the risk of phosphorus loss in sandy soils of tropical ecosystems. Chemosphere 349, 140937 (2024).
Huang, J. et al. Urbanization in China drives soil acidification of Pinus massoniana forests. Sci. Rep. 5, 13512 (2015).
Chen, S. et al. Vegetation cover variations associated with climate change and human activities in Nanjing metropolitan area of China. Environ. Sci. Pollut. Res. 30, 38535–38549 (2023).
Zhang, L. et al. Spatiotemporal evolution characteristics and driving forces of vegetation cover variations in the Chengdu-Chongqing region of China under the background of rapid urbanization. Environ. Sci. Pollut. Res. 31, 22976–22993 (2024).
Li, J. et al. Urbanization-driven forest soil greenhouse gas emissions: Insights from the role of soil bacteria in carbon and nitrogen cycling using a metagenomic approach. Sci. Total Environ. 923, 171364 (2024).
Zheng, F. et al. Urbanization reduces the stability of soil microbial community by reshaping the diversity and network complexity. Chemosphere 364, 143177 (2024).
Yang, L. Y. et al. Diversity and activity of soil N2O-reducing bacteria shaped by urbanization. Environ. Sci. Technol. 58, 17295–17303 (2024).
Burton, V. J. et al. Land use and soil characteristics affect soil organisms differently from above-ground assemblages. BMC Ecol. Evol. 22, 135 (2022).
Huang, C. et al. Opposite effects of soil pH on bacteria and fungi β diversity in forests at a continental scale. J. environ. Manag. 370, 122428 (2024).
Zhang, Y. et al. Temperature fluctuation promotes the thermal adaptation of soil microbial respiration. Nat. Ecol. Evol. 7, 205–213 (2023).
Zuo, X. et al. Contrasting relationships between plant-soil microbial diversity are driven by geographic and experimental precipitation changes. Sci. Total Environ. 861, 160654 (2023).
Faust, K. & Raes, J. Microbial interactions: from networks to models. Nat. Rev. Microbiol. 10, 538–550 (2012).
Wagg, C. et al. Fungal-bacterial diversity and microbiome complexity predict ecosystem functioning. Nat. Commun. 10, 4841 (2019).
Hernandez, D. J. et al. Environmental stress destabilizes microbial networks. ISME J. 15, 1722–1734 (2021).
Berry, D. & Widder, S. Deciphering microbial interactions and detecting keystone species with co-occurrence networks. Front. Microbiol. 5, 219 (2014).
Deng, Y. et al. Molecular ecological network analyses. BMC Bioinform. 13, 113 (2012).
Weiss, S. et al. Correlation detection strategies in microbial data sets vary widely in sensitivity and precision. ISME J. 10, 1669–1681 (2016).
Gloor, G. B. et al. Microbiome datasets are compositional: and this is not optional. Front. Microbiol. 8, 2224 (2017).
Blondel, V. D. et al. Fast unfolding of communities in large networks. J. Stat. Mech. Theory Exp. 10, P10008 (2008).
Chen, X. et al. Changes in soil total, microbial and enzymatic C-N-P contents and stoichiometry with depth and latitude in forest ecosystems. Sci. Total Environ. 816, 151583 (2022).
Delgado-Baquerizo, M. et al. A global atlas of the dominant bacteria found in soil. Sci. 359, 320–325 (2018).
Marczylo, E. L., Macchiarulo, S. & Gant, T. W. Metabarcoding of soil fungi from different urban greenspaces around Bournemouth in the UK. Eco. Health 18, 315–330 (2021).
Ao, L. et al. Different urban forest tree species affect the assembly of the soil bacterial and fungal community. Microb. Ecol. 83, 447–458 (2022).
Qu, Y. et al. Characters and environmental driving factors of bacterial community in soil of Beijing urban parks. Environ. Res. 215, 114178 (2022).
Kielak, A. M. et al. The Ecology of Acidobacteria: Moving beyond Genes and Genomes. Front. Microbiol. 7, 744 (2016).
Hug, L. A. et al. Critical biogeochemical functions in the subsurface are associated with bacteria from new phyla and little studied lineages. Environ. Microbiol. 18, 159–173 (2016).
Miyauchi, S. et al. Large-scale genome sequencing of mycorrhizal fungi provides insights into the early evolution of symbiotic traits. Nat. Commun. 11, 5125 (2020).
Riley, R. et al. Extensive sampling of basidiomycete genomes demonstrates inadequacy of the white-rot/brown-rot paradigm for wood decay fungi. Proc. Natl. Acad. Sci. USA. 111, 9923–9928 (2014).
Piano, E. et al. Urbanization drives cross-taxon declines in abundance and diversity at multiple spatial scales. Glob. Change Biol. 26, 1196–1211 (2020).
Yu, R. et al. Anticipating global terrestrial ecosystem state change using FLUXNET. Glob. Change Biol. 25, 2352–2367 (2019).
Li, Q. et al. Impact of water level fluctuations on the development of phytoplankton in a large subtropical reservoir: implications for the management of cyanobacteria. Environ. Sci. Pollut. Res. 25, 1306–1318 (2018).
Zhao, C. et al. Analysis of changes in the spatiotemporal characteristics of impervious surfaces and their influencing factors in the Central Plains Urban Agglomeration of China from 2000 to 2018. Heliyon 9, e18849 (2023).
Yan, Y. et al. Impacts of impervious surface expansion on soil organic carbon-a spatially explicit study. Sci. Rep. 5, 17905 (2015).
Qiu, L. et al. Substantial terrestrial carbon emissions from global expansion of impervious surface area. Nat. Commun. 15, 6456 (2024).
Nega, W. & Balew, A. The relationship between land use land cover and land surface temperature using remote sensing: systematic reviews of studies globally over the past 5 years. Environ. Sci. Pollut. Res. 29, 42493–42508 (2022).
Hu, Y. et al. Impervious surfaces alter soil bacterial communities in urban areas: a case study in Beijing, China. Front. Microbiol. 9, 226 (2018).
Liu, R. et al. Novel Chloroflexi genomes from the deepest ocean reveal metabolic strategies for the adaptation to deep-sea habitats. Microbiome 10, 75 (2022).
Klaus, J. R. et al. Secondary metabolites from the Burkholderia pseudomallei complex: structure, ecology, and evolution. J. Ind. Microbiol. Biot. 47, 877–887 (2020).
Jaramillo-Madrid, A. C. et al. Phytosterol biosynthesis and production by diatoms (Bacillariophyceae). Phytochemistry 163, 46–57 (2019).
Nemcova, Y. et al. Nanopatterns on silica scales of Mallomonas (Chrysophyceae, Stramenopiles): Unraveling UV resistance potential and diverse response to UVA and UVB radiation. J. phycol. 60, 1256–1272 (2024).
Jiang, R., Wang, M. & Chen, W. Heavy metal pollution triggers a shift from bacteria-based to fungi-based soil micro-food web: Evidence from an abandoned mining-smelting area. J. Hazard. Mater. 459, 132164 (2023).
Dubey, M. et al. Environmental connectivity controls diversity in soil microbial communities. Commun. Biol. 4, 492 (2021).
Asaf, S. et al. Sphingomonas: from diversity and genomics to functional role in environmental remediation and plant growth. Crit. Rev. Biotechnol. 40, 138–152 (2020).
Jahan, I. et al. Comprehensive Analysis of Penicillium Sclerotiorum: Biology, Secondary Metabolites, and Bioactive Compound Potential─A Review. J. Agric. Food Chem. 72, 9555–9566 (2024).
Delgado-Baquerizo, M. et al. Multiple elements of soil biodiversity drive ecosystem functions across biomes. Nat. Ecol. Evol. 4, 210–220 (2020).
Yang, C. X. et al. Plant exudates-driven microbiome recruitment and assembly facilitates plant health management. FEMS Microbiol. Rev. 49, fuaf008 (2025).
Aponte, C., Marañón, T. & García, L. V. Microbial C, N and P in soils of Mediterranean oak forests: influence of season, canopy cover and soil depth. Biogeochemistry 101, 77–92 (2010).
Wang, B. et al. Trophic relationships between protists and bacteria and fungi drive the biogeography of rhizosphere soil microbial community and impact plant physiological and ecological functions. Microbiol. Res. 280, 127603 (2024).
Ren, B. et al. Soil pH and plant diversity shape soil bacterial community structure in the active layer across the latitudinal gradients in continuous permafrost region of Northeastern China. Sci. Rep. 8, 5619 (2018).
Yan, B. et al. Urban-development-induced changes in the diversity and composition of the soil bacterial community in Beijing. Sci. Rep. 6, 38811 (2016).
Wang, H. et al. Soil bacterial diversity is associated with human population density in urban greenspaces. Environ. Sci. Technol. 52, 5115–5124 (2018).
Moriyama, M. & Numata, H. Urban soil compaction reduces cicada diversity. Zool. Lett. 1, 19 (2015).
Wei, Z. et al. Density and stability of soil organic carbon beneath impervious surfaces in urban areas. PloS one 9, e109380 (2014).
Chowdhury, S. et al. Improving stormwater infiltration and retention in compacted urban soils at impervious/pervious surface disconnections with biochar. J. Environ. Manag. 360, 121032 (2024).
Li, J. et al. Thermal adaptation of microbial respiration persists throughout long-term soil carbon decomposition. Ecol. Lett. 26, 1803–1814 (2023).
Munir, R. et al. Detrimental effects of Cd and temperature on rice and functions of microbial community in paddy soils. Environ. Pollut. 324, 121371 (2023).
Chen, Y. et al. Environmental determinants and demographic influences on global urban microbiomes, antimicrobial resistance and pathogenicity. NPJ Biofilms Microb. 9, 94 (2023).
Delgado-Baquerizo, M. et al. Global homogenization of the structure and function in the soil microbiome of urban greenspaces. Sci. Adv. 7, eabg5809 (2021).
Fu, S. et al. Enhancing urban ecological resilience through integrated green technology progress: evidence from Chinese cities. Environ. Sci. Pollut. Res. 31, 36349–36366 (2024).
Audebert, P. et al. Ecological zoning for climate policy and global change studies. Nat. Sustain. 7, 1294–1303 (2024).
Liu, X. et al. Data and code for Urbanization increases the complexity of soil microbial interaction networks [Data set]. Zenodo. https://doi.org/10.5281/zenodo.18819982 (2026).
Liu, X. et al. Data and code for Urbanization increases the complexity of soil microbial interaction networks [Data set]. Zenodo. https://doi.org/10.5281/zenodo.19104164 (2026).
Acknowledgements
We thank the National Key R & D Program of China (2023YFF1304600) for funding support.
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X.L.: Formal analysis, investigation, methodology, visualization, writing-original draft. G.L.: Investigation, methodology. M.H.: Investigation, methodology. P.L.: Conceptualization, data curation, supervision, visualization, validation, writing-review & editing. W.L.: Conceptualization, supervision, writing-review & editing.
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Liu, X., Li, G., Han, M. et al. Urbanization increases the complexity of soil microbial interaction networks.
Commun Biol (2026). https://doi.org/10.1038/s42003-026-09997-x
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DOI: https://doi.org/10.1038/s42003-026-09997-x
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