Effects of urbanization on soil fauna community structure and diversity

doi:10.17520/biods.2022547

Abstract

Abstract:

Background & Aims: Urbanization threatens biodiversity on local to global scales, and soil fauna is an essential component of soil biodiversity and play a vital role in the maintenance of soil health and ecosystem functions in cities. In recent years, many studies previously focused on the structure and diversity of urban soil fauna communities that have made significant progress to understand their complex relationships. However, systematic summaries and discussions are still limited.
Progress: This review advances our knowledge in these areas by drawing upon current urban soil biodiversity studies with relevant broader research on soil fauna. From these works, we summarize the major pathways by which urbanization affects the community composition and diversity of soil fauna and highlight ecological processes that potentially threaten the persistence of soil fauna in cities.
Prospects: We recommend that future studies should focus on (1) the impact of urbanization on gut microbiomes of soil fauna and how it relates to changes in soil fauna functions, (2) the environmental determinants to the structures and functions of soil food webs in urban ecosystems, and (3) the role of soil fauna in the regulation of human pathogens. Ultimately, our review provides a relevant scientific basis for biodiversity protection and maintenance of soil health and ecosystem stability in cities.

Key words: urbanization, soil fauna diversity, soil food web, soil health, land use, soil properties

Haifeng Yao, Saichao Zhang, Huayuan Shangguan, Zhipeng Li, Xin Sun. Effects of urbanization on soil fauna community structure and diversity[J]. Biodiv Sci, 2022, 30(12): 22547.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.biodiversity-science.net/EN/10.17520/biods.2022547

https://www.biodiversity-science.net/EN/Y2022/V30/I12/22547

Figures/Tables 1

References 115

[1]	Adhikari K, Hartemink AE (2016) Linking soils to ecosystem services—A global review. Geoderma, 262, 101-111. DOI URL
[2]	Agamennone V, Jakupović D, Weedon JT, Suring WJ, van Straalen NM, Roelofs D, Röling WFM (2015) The microbiome of Folsomia candida: An assessment of bacterial diversity in a Wolbachia-containing animal. FEMS Microbiology Ecology, 91, fiv128.
[3]	Amin NM (2012) Techniques for assessment of heavy metal toxicity using Acanthamoeba sp., a small, naked and free-living amoeba. The Functioning of Ecosystems, 199-212.
[4]	Amossé J, Dózsa-Farkas K, Boros G, Rochat G, Sandoz G, Fournier B, Mitchell EAD,Le Bayon RC (2016) Patterns of earthworm, enchytraeid and nematode diversity and community structure in urban soils of different ages. European Journal of Soil Biology, 73, 46-58. DOI URL
[5]	Andriuzzi WS, Pulleman MM, Schmidt O, Faber JH, Brussaard L (2015) Anecic earthworms (Lumbricus terrestris) alleviate negative effects of extreme rainfall events on soil and plants in field mesocosms. Plant and Soil, 397, 103-113. DOI URL
[6]	Antrop M (2004) Landscape change and the urbanization process in Europe. Landscape and Urban Planning, 67, 9-26. DOI URL
[7]	Asabere SB, Zeppenfeld T, Nketia KA, Sauer D (2018) Urbanization leads to increases in pH, carbonate, and soil organic matter stocks of arable soils of Kumasi, Ghana (West Africa). Frontiers in Environmental Science, 6, 119. DOI URL
[8]	Bach EM, Ramirez KS, Fraser TD, Wall DH (2020) Soil biodiversity integrates solutions for a sustainable future. Sustainability, 12, 2662. DOI URL
[9]	Beaumelle L, Thouvenot L, Hines J, Jochum M, Eisenhauer N, Phillips HRP (2021) Soil fauna diversity and chemical stressors: A review of knowledge gaps and roadmap for future research. Ecography, 44, 845-859. DOI URL
[10]	Berg M, Stenuit B, Ho J, Wang A, Parke C, Knight M, Alvarez-Cohen L, Shapira M (2016) Assembly of the Caenorhabditis elegans gut microbiota from diverse soil microbial environments. The ISME Journal, 10, 1998-2009. DOI URL
[11]	Berger H, Foissner W, Adam H (1985) Morphological variation and comparative analysis of morphogenesis in Parakahliella macrostoma (Foissner, 1982) nov. gen. and Histriculus muscorum (Kahl, 1932), (Ciliophora, Hypotrichida). Protistologica, 21, 295-311.
[12]	Bongers T, van der Meulen H, Korthals G (1997) Inverse relationship between the nematode maturity index and plant parasite index under enriched nutrient conditions. Applied Soil Ecology, 6, 195-199. DOI URL
[13]	Bottinelli N, Jouquet P, Capowiez Y, Podwojewski P, Grimaldi M, Peng X (2015) Why is the influence of soil macrofauna on soil structure only considered by soil ecologists? Soil and Tillage Research, 146, 118-124.
[14]	Chang CH, Johnston MR, Görres JH, Dávalos A, McHugh D, Szlavecz K (2018) Co-invasion of three Asian earthworms, Metaphire hilgendorfi, Amynthas agrestis and Amynthas tokioensis in the USA. Biological Invasions, 20, 843-848.
[15]	Chen XD, Dunfield KE, Fraser TD, Wakelin SA, Richardson AE, Condron LM (2020) Soil biodiversity and biogeochemical function in managed ecosystems. Soil Research, 58, 1-20. DOI URL
[16]	Creamer RE, Rimmer DL, Black HIJ (2008) Do elevated soil concentrations of metals affect the diversity and activity of soil invertebrates in the long-term? Soil Use and Management, 24, 37-46.
[17]	David JF (2014) The role of litter-feeding macroarthropods in decomposition processes: A reappraisal of common views. Soil Biology and Biochemistry, 76, 109-118. DOI URL
[18]	Deckmyn G, Flores O, Mayer M, Domene X, Schnepf A, Kuka K, van Looy K, Rasse DP, Briones MJI, Barot S, Berg M, Vanguelova E, Ostonen I, Vereecken H, Suz LM, Frey B, Frossard A, Tiunov A, Frouz J, Grebenc T, Öpik M, Javaux M, Uvarov A, Vindušková O, Krogh PH, Franklin O, Jiménez J, Yuste JC (2020) KEYLINK: Towards a more integrative soil representation for inclusion in ecosystem scale models. I. Review and model concept. PeerJ, 8, e9750.
[19]	Dehlin H, Peltzer DA, Allison VJ, Yeates GW, Nilsson MC, Wardle DA (2008) Tree seedling performance and below-ground properties in stands of invasive and native tree species. New Zealand Journal of Ecology, 32, 67-79.
[20]	Delgado-Baquerizo M, Eldridge DJ, Liu YR, Sokoya B, Wang JT, Hu HW, He JZ, Bastida F, Moreno JL, Bamigboye AR, Blanco-Pastor JL, Cano-Díaz C, Illán JG, Makhalanyane TP, Siebe C, Trivedi P, Zaady E, Verma JP, Wang L, Wang JY, Grebenc T, Peñaloza-Bojacá GF, Nahberger TU, Teixido AL, Zhou XQ, Berdugo M, Duran J, Rodríguez A, Zhou XB, Alfaro F, Abades S, Plaza C, Rey A, Singh BK, Tedersoo L, Fierer N (2021) Global homogenization of the structure and function in the soil microbiome of urban greenspaces. Science Advances, 7, eabg5809.
[21]	Delgado-Baquerizo M, Reich PB, Trivedi C, Eldridge DJ, Abades S, Alfaro FD, Bastida F, Berhe AA, Cutler NA, Gallardo A, García-Velázquez L, Hart SC, Hayes PE, He JZ, Hseu ZY, Hu HW, Kirchmair M, Neuhauser S, Pérez CA, Reed SC, Santos F, Sullivan BW, Trivedi P, Wang JT, Weber-Grullon L, Williams MA, Singh BK (2020) Multiple elements of soil biodiversity drive ecosystem functions across biomes. Nature Ecology and Evolution, 4, 210-220. DOI PMID
[22]	Dequiedt S, Saby NPA, Lelievre M, Jolivet C, Thioulouse J, Toutain B, Arrouays D, Arrouays D, Bispo A, Lemanceau P, Ranjard L (2011) Biogeographical patterns of soil molecular microbial biomass as influenced by soil characteristics and management. Global Ecology and Biogeography, 20, 641-652. DOI URL
[23]	Deshmukh VA, Shrikhande JG (1974) Effect of herbicides on protozoa in the soil. Indian Journal of Microbiology, 14, 273.
[24]	Faeth SH, Bang C, Saari S (2011) Urban biodiversity:Patterns and mechanisms. Annals of the New York Academy of Sciences, 1223, 69-81.
[25]	FAO, ITPS, GSBI, SCBD, EC,Marin C (2020) State of Knowledge of Soil Biodiversity—Status, Challenges and Potentialities, Report 2020. FAO, Rome.
[26]	Fenoglio MS, Rossetti MR, Videla M (2020) Negative effects of urbanization on terrestrial arthropod communities: A meta-analysis. Global Ecology and Biogeography, 29, 1412-1429. DOI URL
[27]	Ferris H, Tuomisto H (2015) Unearthing the role of biological diversity in soil health. Soil Biology and Biochemistry, 85, 101-109. DOI URL
[28]	Filser J, Faber JH, Tiunov AV, Brussaard L, Frouz J, De Deyn G, Uvarov AV, Berg MP, Lavelle P, Loreau M, Wall DH, Querner P, Eijsackers H, Jiménez JJ (2016) Soil fauna: Key to new carbon models. Soil, 2, 565-582. DOI URL
[29]	Foissner W (1987) Soil protozoa: Fundamental problems, ecological significance, adaptation in ciliates and testaceans, bioindicators, and guide to the literature. Prog Protistol, 2, 69-212.
[30]	Frouz J (2018) Effects of soil macro- and mesofauna on litter decomposition and soil organic matter stabilization. Geoderma, 332, 161-172. DOI URL
[31]	Ge BM, Li ZX, Zhang DZ, Zhang HB, Liu ZT, Zhou CL, Tang BP (2012) Biodiversity of soil macrofauna communities in five kinds of green-land in spring at Yancheng City, Jiangsu. Chinese Journal of Zoology, 47, 1-7. (in Chinese with English abstract)
	[ 葛宝明, 李振兴, 张代臻, 张华彬, 刘总堂, 周春霖, 唐伯平 (2012) 盐城5种绿地春季大型土壤动物群落的生物多样性. 动物学杂志, 47, 1-7.]
[32]	Geisen S, Mitchell EAD, Adl S, Bonkowski M, Dunthorn M, Ekelund F, Fernández LD, Jousset A, Krashevska V, Singer D, Spiegel FW, Walochnik J, Lara E (2018) Soil protists: A fertile frontier in soil biology research. FEMS Microbiology Reviews, 42, 293-323. DOI PMID
[33]	George PBL, Keith AM, Creer S, Barrett GL, Lebron I, Emmett BA, Robinson DA, Jones DL (2017) Evaluation of mesofauna communities as soil quality indicators in a national-level monitoring programme. Soil Biology and Biochemistry, 115, 537-546. DOI URL
[34]	Gongalsky KB (2021) Soil macrofauna: Study problems and perspectives. Soil Biology and Biochemistry, 159, 108281. DOI URL
[35]	Graham SI, Kinnaird MF, O’Brien TG, Vågen TG, Winowiecki LA, Young TP, Young HS (2019) Effects of land-use change on community diversity and composition are highly variable among functional groups. Ecological Applications, 29, e01973.
[36]	Grewal SS, Cheng Z, Masih S, Wolboldt M, Huda N, Knight A, Grewal PS (2011) An assessment of soil nematode food webs and nutrient pools in community gardens and vacant lots in two post-industrial American cities. Urban Ecosystems, 14, 181-194. DOI URL
[37]	Guilland C, Maron PA, Damas O, Ranjard L (2018) Biodiversity of urban soils for sustainable cities. Environmental Chemistry Letters, 16, 1267-1282. DOI URL
[38]	Harada H, Kurauchi M, Hayashi R, Eki T (2007) Shortened lifespan of nematode Caenorhabditis elegans after prolonged exposure to heavy metals and detergents. Ecotoxicology and Environmental Safety, 66, 378-383. DOI URL
[39]	Hartley DJ, Koivula MJ, Spence JR, Pelletier R, Ball GE (2007) Effects of urbanization on ground beetle assemblages (Coleoptera, Carabidae) of grassland habitats in western Canada. Ecography, 30, 673-684. DOI URL
[40]	Heringer G, Del Bianco Faria L, Villa PM, Araújo AU, Botan ALM, Zenni RD (2022) Urbanization affects the richness of invasive alien trees but has limited influence on species composition. Urban Ecosystems, 25, 753-763. DOI URL
[41]	Heringer G, Thiele J, Alves Meira-Neto JA, Neri AV (2019) Biological invasion threatens the sandy-savanna Mussununga ecosystem in the Brazilian Atlantic Forest. Biological Invasions, 21, 2045-2057. DOI
[42]	Hruska K (2006) Notes on the evolution and organization of the urban ecosystem. Urban Ecosystems, 9, 291-298. DOI URL
[43]	Hu MM, Tian L, Wu YN, Yang JY, Lü XC, Huang XR (2019) Influences of thinning and mixed transformation of Larix principis-rupprechtii plantations on the community structure of soil macro faunal in Saihanba Area. Scientia Silvae Sinicae, 55, 153-162. (in Chinese with English abstract)
	[ 扈梦梅, 田龙, 吴亚楠, 杨晋宇, 吕小翠, 黄选瑞 (2019) 塞罕坝华北落叶松人工林间伐和混交改造对大型土壤动物群落结构的影响. 林业科学, 55, 153-162.]
[44]	Huang YM, Zhang K, Sun LX, Zhang J, Li SY, Hu ZY, Wang RR, Li X, Xiong X, Xie WF (2018) Effects of Trifolium repens invasion on soil animals in an urban turf ecosystem. Acta Ecologica Sinica, 38, 8489-8499. (in Chinese with English abstract)
	[ 黄玉梅, 张凯, 孙凌霞, 张健, 李三月, 胡贞艳, 王若然, 李向, 熊茜, 解文峰 (2018) 白三叶(Trifolium repens)入侵对城市草坪生态系统土壤动物的影响. 生态学报, 38, 8489-8499.]
[45]	Joimel S, Capiaux H, Schwartz C, Hedde M, Lebeau T, Le Guern C, Nahmani J, Pernin C, Salmon S, Santorufo L, Béchet B, Cortet J (2018a) Effect of geogenic lead on fungal and collembolan communities in garden topsoil. Pedosphere, 28, 215-226. DOI URL
[46]	Joimel S, Grard B, Auclerc A, Hedde M, Le Doaré N, Salmon S, Chenu C (2018b) Are Collembola “flying” onto green roofs? Ecological Engineering, 111, 117-124.
[47]	Joimel S, Schwartz C, Maurel N, Magnus B, Machon N, Bel J, Cortet J (2019) Contrasting homogenization patterns of plant and collembolan communities in urban vegetable gardens. Urban Ecosystems, 22, 553-566. DOI URL
[48]	Kaiser D, Lepage M, Konaté S, Linsenmair KE (2017) Ecosystem services of termites (Blattoidea: Termitoidae) in the traditional soil restoration and cropping system Zaï in northern Burkina Faso (West Africa). Agriculture, Ecosystems and Environment, 236, 198-211. DOI URL
[49]	Knapp S, Dinsmore L, Fissore C, Hobbie SE, Jakobsdottir I, Kattge J, King JY, Klotz S, McFadden JP, Cavender-Bares J (2012) Phylogenetic and functional characteristics of household yard floras and their changes along an urbanization gradient. Ecology, 93, S83-S98.
[50]	Kooch Y, Noghre N (2020) The effect of shrubland and grassland vegetation types on soil fauna and flora activities in a mountainous semi-arid landscape of Iran. Science of the Total Environment, 703, 135497. DOI URL
[51]	Kozlov MV, Zverev V, Gusarov VI, Korobushkin DI, Krivosheina NP, Mattila J, Mutanen M, Popova A, Prosvirov AS, Punttila P, Söderman G, Stańska M, Taylor A, Vahtera V, Zubrii NA, Zvereva EL (2022) Changes in biomass and diversity of soil macrofauna along a climatic gradient in European boreal forests. Insects, 13, 94. DOI URL
[52]	Lazzaro L, Mazza G, d’Errico G, Fabiani A, Giuliani C, Inghilesi AF, Lagomarsino A, Landi S, Lastrucci L, Pastorelli R, Roversi PF, Torrini G, Tricarico E, Foggi B (2018) How ecosystems change following invasion by Robinia pseudoacacia: Insights from soil chemical properties and soil microbial, nematode, microarthropod and plant communities. Science of the Total Environment, 622/623, 1509-1518.
[53]	Ledeganck P, Nijs I, Beyens L (2003) Plant functional group diversity promotes soil protist diversity. Protist, 154, 239-249. PMID
[54]	Lei YB, Xiao HF, Feng YL (2010) Impacts of alien plant invasions on biodiversity and evolutionary responses of native species. Biodiversity Science, 18, 622-630. (in Chinese with English abstract) DOI
	[ 类延宝, 肖海峰, 冯玉龙 (2010) 外来植物入侵对生物多样性的影响及本地生物的进化响应. 生物多样性, 18, 622-630.] DOI
[55]	Lescano MN, Elizalde L, Werenkraut V, Pirk GI, Flores GE (2017) Ant and tenebrionid beetle assemblages in arid lands: Their associations with vegetation types in the Patagonian steppe. Journal of Arid Environments, 138, 51-57. DOI URL
[56]	Li J, Peng P, Zhao J (2020) Assessment of soil nematode diversity based on different taxonomic levels and functional groups. Soil Ecology Letters, 2, 33-39. DOI
[57]	Li X, Liu T, Li H, Geisen S, Hu F, Liu M (2022) Management effects on soil nematode abundance differ among functional groups and land‐use types at a global scale. Journal of Animal Ecology, 91, 1770-1780. DOI URL
[58]	Liu A, Zhao XK, Zhou X, Lu HC (2022) Investigation on the diversity of soil macrofauna in different vegetation in Century Park in spring. Special Economic Animals and Plants, 25(1), 6-9. (in Chinese)
	[ 刘傲, 赵新康, 周鑫, 卢慧翠 (2022) 世纪公园周边春季不同植被大型土壤动物多样性研究. 特种经济动植物, 25(1), 6-9.]
[59]	Lopez BE, Urban D, White PS (2018) Testing the effects of four urbanization filters on forest plant taxonomic, functional, and phylogenetic diversity. Ecological Applications, 28, 2197-2205. DOI URL
[60]	Lu YY, Wu BQ, Xu YJ, Zeng L (2012) Effects of red imported fire ants (Solenopsis invicta) on the species structure of ant communities in South China. Sociobiology, 59, 275-286. DOI URL
[61]	Malcicka M, Berg MP, Ellers J (2017) Ecomorphological adaptations in Collembola in relation to feeding strategies and microhabitat. European Journal of Soil Biology, 78, 82-91. DOI URL
[62]	Mao Q, Huang G, Buyantuev A, Wu J, Luo S, Ma K (2014) Spatial heterogeneity of urban soils: The case of the Beijing Metropolitan region, China. Ecological Processes, 3, 1-11. DOI URL
[63]	Mathieu J, Grimaldi M, Jouquet P, Rouland C, Lavelle P, Desjardins T, Rossi JP (2009) Spatial patterns of grasses influence soil macrofauna biodiversity in Amazonian pastures. Soil Biology and Biochemistry, 41, 586-593. DOI URL
[64]	McCay TS, Scull P (2019) Invasive lumbricid earthworms in northeastern North American forests and consequences for leaf-litter fauna. Biological Invasions, 21, 2081-2093.
[65]	McDonald RI, Marcotullio PJ, Güneralp B (eds 2013) Urbanization and global trends in biodiversity and ecosystem services. In: Urbanization, Biodiversity and Ecosystem Services: Challenges and Opportunities (Elmqvist T, Fragkias M, Goodness J, Güneralp B, Marcotullio PJ, McDonald RI, Parnell S, Schewenius M, Sendstad M, Seto KC, Wilkinson C), pp.31-52. Springer, Dordrecht.
[66]	Mealor BA, Hild AL (2007) Post-invasion evolution of native plant populations: A test of biological resilience. Oikos, 116, 1493-1500. DOI URL
[67]	Milano V, Cortet J, Baldantoni D, Bellino A, Dubs F, Nahmani J, Strumia S, Maisto G (2017) Collembolan biodiversity in Mediterranean urban parks: Impact of history, urbanization, management and soil characteristics. Applied Soil Ecology, 119, 428-437. DOI URL
[68]	Milano V, Maisto G, Baldantoni D, Bellino A, Bernard C, Croce A, Dubs F, Strumia S, Cortet J (2018) The effect of urban park landscapes on soil Collembola diversity: A Mediterranean case study. Landscape and Urban Planning, 180, 135-147. DOI URL
[69]	Miles LS, Breitbart ST, Wagner HH, Johnson MT (2019) Urbanization shapes the ecology and evolution of plant-arthropod herbivore interactions. Frontiers in Ecology and Evolution, 7, 310. DOI URL
[70]	Minor MA, Babenko AB, Ermilov SG, Khaustov AA, Makarova OL (2016) Effects of cushion plants on high-altitude soil microarthropod communities: Cushions increase abundance and diversity of mites (Acari), but not springtails (Collembola). Arctic, Antarctic, and Alpine Research, 48, 485-500.
[71]	Moore JC, Berlow EL, Coleman DC, De Suiter PC, Dong Q, Hastings A, Johnson NC, McCann KS, Melville K, Morin PJ, Nadelhoffer K, Rosemond AD, Post DM, Sabo JL, Scow KM, Vanni MJ, Wall DH (2004) Detritus, trophic dynamics and biodiversity. Ecology Letters, 7, 584-600. DOI URL
[72]	Neher DA (2001) Role of nematodes in soil health and their use as indicators. Journal of Nematology, 33, 161-168. PMID
[73]	Nielsen UN (2019) Soil Fauna Assemblages. Cambridge University Press, Cambridge.
[74]	Nielsen UN, Ayres E, Wall DH, Li G, Bardgett RD, Wu T, Garey JR (2014) Global-scale patterns of assemblage structure of soil nematodes in relation to climate and ecosystem properties. Global Ecology and Biogeography, 23, 968-978. DOI URL
[75]	Oliverio AM, Geisen S, Delgado-Baquerizo M, Maestre FT, Turner BL, Fierer N (2020) The global-scale distributions of soil protists and their contributions to belowground systems. Science Advances, 6, eaax8787.
[76]	Pothula SK, Grewal PS, Auge RM, Saxton AM, Bernard EC (2019) Agricultural intensification and urbanization negatively impact soil nematode richness and abundance: A meta-analysis. Journal of Nematology, 51, e2019-11.
[77]	Pouyat RV, Parmelee RW, Carreiro MM (1994) Environmental effects of forest soil-invertebrate and fungal densities in oak stands along an urban-rural land use gradient. Pedobiologia, 38, 385-399.
[78]	Pouyat RV, Szlavecz K, Yesilonis ID, Groffman PM, Schwarz K (2010) Chemical, physical, and biological characteristics of urban soils. Urban Ecosystem Ecology, 55, 119-152.
[79]	Qian S, Qin D, Wu X, Hu S, Hu L, Lin D, Zhao L, Shang K, Song K, Yang Y (2020) Urban growth and topographical factors shape patterns of spontaneous plant community diversity in a mountainous city in Southwest China. Urban Forestry and Urban Greening, 55, 126814. DOI URL
[80]	Qin J, Xu KF (2018) Research summary and prospect of urban green space soil quality in China. Ecological Science, 37, 200-210. (in Chinese with English abstract)
	[ 秦娟, 许克福 (2018) 我国城市绿地土壤质量研究综述与展望. 生态科学, 37, 200-210.]
[81]	Quan GM, Zhang JE, Xie JF, Mao DJ, Xu HQ, Jiang WB, Wen DJ (2011) Impact of Mikania micrantha invasion on soil meso- and micro-invertebrate community structure. Chinese Journal of Applied Ecology, 22, 1863-1870. (in Chinese with English abstract)
	[ 全国明, 章家恩, 谢俊芳, 毛丹鹃, 徐华勤, 姜万兵, 文杜娟 (2011) 薇甘菊入侵对中小型土壤动物群落结构特征的影响. 应用生态学报, 22, 1863-1870.]
[82]	Rzeszowski K, Zadrożny P, Nicia P (2017) The effect of soil nutrient gradients on Collembola communities inhabiting typical urban green spaces. Pedobiologia, 64, 15-24. DOI URL
[83]	Sadler JP, Small EC, Fiszpan H, Telfer MG, Niemelä J (2006) Investigating environmental variation and landscape characteristics of an urban-rural gradient using woodland carabid assemblages. Journal of Biogeography, 33, 1126-1138. DOI URL
[84]	Santorufo L, Cortet J, Arena C, Goudon R, Rakoto A, Morel JL, Maisto G (2014) An assessment of the influence of the urban environment on collembolan communities in soils using taxonomy- and trait-based approaches. Applied Soil Ecology, 78, 48-56. DOI URL
[85]	Santorufo L, Cortet J, Nahmani J, Pernin C, Salmon S, Pernot A, Morel JL, Maisto G (2015) Responses of functional and taxonomic collembolan community structure to site management in Mediterranean urban and surrounding areas. European Journal of Soil Biology, 70, 46-57. DOI URL
[86]	Scherber C, Eisenhauer N, Weisser WW, Schmid B, Voigt W, Fischer M, Schulze ED, Roscher C, Weigelt A, Allan E, Beßler H, Bonkowski M, Buchmann N, Buscot F, Clement LW, Ebeling A, Engels C, Halle S, Kertscher I, Klein AM, Koller R, König S, Kowalski E, Kummer V, Kuu A, Lange M, Lauterbach D, Middelhoff C, Migunova VD, Milcu A, Müller R, Partsch S, Petermann JS, Renker C, Rottstock T, Sabais A, Scheu S, Schumacher J, Temperton VM, Tscharntke T (2010) Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment. Nature, 468, 553-556. DOI URL
[87]	Seto KC, Güneralp B, Hutyra LR (2012) Global forecasts of urban expansion to 2030 and direct impacts on biodiversity and carbon pools. Proceedings of the National Academy of Sciences, USA, 109, 16083-16088.
[88]	Shao YH, Zhang WX, Liu SJ, Wang XL, Fu SL (2015) Diversity and function of soil fauna. Acta Ecologica Sinica, 35, 6614-6625. (in Chinese with English abstract)
	[ 邵元虎, 张卫信, 刘胜杰, 王晓丽, 傅声雷 (2015) 土壤动物多样性及其生态功能. 生态学报, 35, 6614-6625.]
[89]	Stefan G, Cornelia B, Jörg R, Michael B (2014) Soil water availability strongly alters the community composition of soil protists. Pedobiologia, 57, 205-213. DOI URL
[90]	Sun JL, Chen ZX, Ni HG, Zeng H (2013) PBDEs as indicator chemicals of urbanization along an urban/rural gradient in South China. Chemosphere, 92, 471-476. DOI URL
[91]	Sutton CA, Wilkinson DM (2007) The effects of Rhododendron on testate amoebae communities in woodland soils in North West England. Acta Protozoologica, 46, 333-338.
[92]	Sylvain ZA, Wall DH (2011) Linking soil biodiversity and vegetation: Implications for a changing planet. American Journal of Botany, 98, 517-527. DOI PMID
[93]	Tibbett M, Fraser TD, Duddigan S (2020) Identifying potential threats to soil biodiversity. PeerJ, 8, e9271.
[94]	Tresch S, Frey D, Le Bayon RC, Zanetta A, Rasche F, Fliessbach A, Moretti M (2019) Litter decomposition driven by soil fauna, plant diversity and soil management in urban gardens. Science of the Total Environment, 658, 1614-1629. DOI
[95]	van den Hoogen J, Geisen S, Routh D, Ferris H, Traunspurger W, Wardle DA, de Goede RGM, Adams BJ, Ahmad W, Andriuzzi WS, Bardgett RD, Bonkowski M, Campos-Herrera R, Cares JE, Caruso T, de Brito Caixeta L, Chen XY, Costa SR, Creamer R,Mauro da Cunha Castro J, Dam M, Djigal D, Escuer M, Griffiths BS, Gutiérrez C, Hohberg K, Kalinkina D, Kardol P, Kergunteuil A, Korthals G, Krashevska V, Kudrin AA, Li Q, Liang W, Magilton M, Marais M, Martín JAR, Matveeva E, Mayad EH, Mulder C, Mullin P, Neilson R, Nguyen TAD, Nielsen UN, Okada H, Rius JEP, Pan K, Peneva V, Pellissier L, Carlos Pereira da Silva J, Pitteloud C, Powers TO, Powers K, Quist CW, Rasmann S, Moreno SS, Scheu S, Setälä H, Sushchuk A, Tiunov AV, Trap J, van der Putten W, Vestergård M, Villenave C, Waeyenberge L, Wall DH, Wilschut R, Wright DG, Yang JI, Crowther TW (2019) Soil nematode abundance and functional group composition at a global scale. Nature, 572, 194-198. DOI URL
[96]	Vohník M, Burdíková Z, Wilkinson DM (2012) Testate amoebae communities in the rhizosphere of Rhododendron ponticum (Ericaceae) in an evergreen broadleaf forest in southern Spain. Acta Protozoologica, 51, 259-269.
[97]	Walker JS, Grimm NB, Briggs JM, Gries C, Dugan L (2009) Effects of urbanization on plant species diversity in central Arizona. Frontiers in Ecology and the Environment, 7, 465-470. DOI URL
[98]	Wall DH, Nielsen UN, Six J (2015) Soil biodiversity and human health. Nature, 528, 69-76. DOI URL
[99]	Wang HF, López-Pujol J, Meyerson LA, Qiu JX, Wang XK, Ouyang ZY (2011) Biological invasions in rapidly urbanizing areas: A case study of Beijing, China. Biodiversity and Conservation, 20, 2483-2509. DOI URL
[100]	Wang L, Xu YJ, Zeng L, Lu YY (2019) Impact of the red imported fire ant Solenopsis invicta Buren on biodiversity in South China: A review. Journal of Integrative Agriculture, 18, 788-796. DOI URL
[101]	Wang M, Li JX, Kuang SJ, He YJ, Chen GJ, Huang Y, Song CH, Anderson P, Łowicki D (2020) Plant diversity along the urban-rural gradient and its relationship with urbanization degree in Shanghai, China. Forests, 11, 171-188. DOI URL
[102]	Wang P, Wang TY, Giesy JP, Lu YL (2013) Perfluorinated compounds in soils from Liaodong Bay with concentrated fluorine industry parks in China. Chemosphere, 91, 751-757. DOI PMID
[103]	Wang QL, Adams CA, Wang FY, Sun YH, Zhang SW (2022) Interactions between microplastics and soil fauna: A critical review. Critical Reviews in Environmental Science and Technology, 52, 3211-3243. DOI URL
[104]	Wang XK, Su YB, Ren YF, Zhang HX, Sun X, Ouyang ZY (2020) Urban ecosystem: Highly spatial heterogeneity. Acta Ecologica Sinica, 40, 5103-5112. (in Chinese with English abstract)
	[ 王效科, 苏跃波, 任玉芬, 张红星, 孙旭, 欧阳志云 (2020) 城市生态系统: 高度空间异质性. 生态学报, 40, 5103-5112.]
[105]	Wu J (2014) Urban ecology and sustainability: The state-of-the-science and future directions. Landscape and Urban Planning, 125, 209-221. DOI URL
[106]	Wu W, Xiu CL, Hu YM, Li CL (2018) A review of the effects of urban landscape pattern on soil fauna diversity. Chinese Journal of Ecology, 37, 2199-2204. (in Chinese with English abstract)
	[ 吴文, 修春亮, 胡远满, 李春林 (2018) 城市景观格局对土壤动物多样性的影响研究进展. 生态学杂志, 37, 2199-2204.]
[107]	Xie T, Hou Y, Chen WP, Wang ME, Lü SD, Li XZ (2019) Impact of urbanization on the soil ecological environment: A review. Acta Ecologica Sinica, 39, 1154-1164. (in Chinese with English abstract)
	[ 谢天, 侯鹰, 陈卫平, 王美娥, 吕斯丹, 李勖之 (2019) 城市化对土壤生态环境的影响研究进展. 生态学报, 39, 1154-1164.]
[108]	Yan ZG, Teng MJ, He W, Liu AQ, Li YR, Wang PC (2019) Impervious surface area is a key predictor for urban plant diversity in a city undergone rapid urbanization. Science of the Total Environment, 650, 335-342. DOI URL
[109]	Yang LH, Shi HY, You ZQ, Tang ZH, Liu H, Jia XD (2015) Effect of different land use types on soil macrofauna community structure. Journal of Sichuan Agricultural University, 33, 208-214. (in Chinese with English abstract)
	[ 杨丽红, 石红艳, 游章强, 唐中海, 刘昊, 贾小东 (2015) 不同土地利用方式对大型土壤动物群落结构的影响. 四川农业大学学报, 33, 208-214.]
[110]	Yeates GW, Bongers T (1999) Nematode diversity in agroecosystems. In: InvertebrateBiodiversity as Bioindicators of Sustainable Landscapes (ed. Paoletti MG), pp.113-135. Elsevier, Amsterdam.
[111]	Yeates GW, Bongers T, de Goede RGM, Freckman DW, Georgieva S (1993) Feeding habits in soil nematode families and genera—An outline for soil ecologists. Journal of Nematology, 25, 315-331. PMID
[112]	You Y (2016) The situation analysis and countermeasures on situation of transmission and damage control for giant Africa snail. Journal of Guangxi Agriculture, 31(1), 46-48. (in Chinese with English abstract)
	[ 游意 (2016) 非洲大蜗牛的分布、传播、为害及防治现状. 广西农学报, 31(1), 46-48.]
[113]	Zhang L, Liu JG, Hu JX, Liu C, Guo WG, Wang Q, Wang H (2012) The inventory of sources, environmental releases and risk assessment for perfluorooctane sulfonate in China. Environmental Pollution, 165, 193-198. DOI PMID
[114]	Zhang P, Li B, Wu J, Hu S (2019) Invasive plants differentially affect soil biota through litter and rhizosphere pathways: A meta-analysis. Ecology letters, 22, 200-210. DOI PMID
[115]	Zhu D, Chen QL, An XL, Yang XR, Christie P, Ke X, Wu LH, Zhu YG (2018) Exposure of soil collembolans to microplastics perturbs their gut microbiota and alters their isotopic composition. Soil Biology and Biochemistry, 116, 302-310. DOI URL