Anthony, M. A., Bender, S. F. & van der Heijden, M. G. A. Enumerating soil biodiversity. Proc. Natl Acad. Sci. USA 120, e2304663120 (2023).
Gessner, M. O. et al. Diversity meets decomposition. Trends Ecol. Evol. 25, 372–380 (2010).
Bardgett, R. D. & Wardle, D. A. Aboveground-Belowground Linkages: Biotic Interactions, Ecosystem Processes, and Global Change (Oxford Univ. Press, 2010).
Schmitz, O. J. & Leroux, S. J. Food webs and ecosystems: linking species interactions to the carbon cycle. Annu. Rev. Ecol. Evol. Syst. 51, 271–295 (2020).
de Vries, F. T. et al. Soil food web properties explain ecosystem services across European land use systems. Proc. Natl Acad. Sci. USA 110, 14296–14301 (2013).
Scheu, S. Plants and generalist predators as links between the below-ground and above-ground system. Basic Appl. Ecol. 2, 3–13 (2001).
Wardle, D. A. et al. Ecological linkages between aboveground and belowground biota. Science 304, 1629–1633 (2004).
Bardgett, R. D. & van der Putten, W. H. Belowground biodiversity and ecosystem functioning. Nature 515, 505–511 (2014).
Guerra, C. A. et al. Tracking, targeting, and conserving soil biodiversity. Science 371, 239–241 (2021).
Handa, I. T. et al. Consequences of biodiversity loss for litter decomposition across biomes. Nature 509, 218–221 (2014).
Heemsbergen, D. A. et al. Biodiversity effects on soil processes explained by interspecific functional dissimilarity. Science 306, 1019–1020 (2004).
Levine, J. M. & HilleRisLambers, J. The importance of niches for the maintenance of species diversity. Nature 461, 254–257 (2009).
Thébault, E. & Loreau, M. Trophic interactions and the relationship between species diversity and ecosystem stability. Am. Nat. 166, E95–E114 (2005).
Hedde, M. et al. A common framework for developing robust soil fauna classifications. Geoderma 426, 116073 (2022).
Potapov, A. M. et al. Feeding habits and multifunctional classification of soil-associated consumers from protists to vertebrates. Biol. Rev. 97, 1057–1117 (2022).
Blouin, M. et al. A review of earthworm impact on soil function and ecosystem services: earthworm impact on ecosystem services. Eur. J. Soil Sci. 64, 161–182 (2013).
Brussaard, L., Pulleman, M. M., Ouédraogo, É, Mando, A. & Six, J. Soil fauna and soil function in the fabric of the food web. Pedobiologia 50, 447–462 (2007).
Wu, D., Du, E., Eisenhauer, N., Mathieu, J. & Chu, C. Global engineering effects of soil invertebrates on ecosystem functions. Nature 640, 120–129 (2025).
Filser, J. et al. Soil fauna: key to new carbon models. Soil 2, 565–582 (2016).
Lussenhop, J. Mechanisms of microarthropod-microbial interactions in soil. Adv. Ecol. Res. 23, 1–33 (1992).
Scheu, S., Ruess, L. & Bonkowski, M. in Microorganisms in Soils: Roles in Genesis and Functions (eds Varma, A. & Buscot, F.) 253–275 (Springer-Verlag, 2005).
Majdi, N., Boiché, A., Traunspurger, W. & Lecerf, A. Predator effects on a detritus-based food web are primarily mediated by non-trophic interactions. J. Anim. Ecol. 83, 953–962 (2014).
Jiang, Y. et al. Unraveling the importance of top-down predation on bacterial diversity at the soil aggregate level. Geoderma 439, 116658 (2023).
Bommarco, R., Kleijn, D. & Potts, S. G. Ecological intensification: harnessing ecosystem services for food security. Trends Ecol. Evol. 28, 230–238 (2013).
Matson, P. A. Agricultural intensification and ecosystem properties. Science 277, 504–509 (1997).
Newbold, T. et al. Global effects of land use on local terrestrial biodiversity. Nature 520, 45–50 (2015).
Potapov, A. M. et al. Rainforest transformation reallocates energy from green to brown food webs. Nature 627, 116–122 (2024).
de Vries, F. T. et al. Land use alters the resistance and resilience of soil food webs to drought. Nat. Clim. Change 2, 276–280 (2012).
Zhou, Z., Krashevska, V., Widyastuti, R., Scheu, S. & Potapov, A. Tropical land use alters functional diversity of soil food webs and leads to monopolization of the detrital energy channel. eLife 11, e75428 (2022).
Barnes, A. D. et al. Consequences of tropical land use for multitrophic biodiversity and ecosystem functioning. Nat. Commun. 5, 5351 (2014).
Rooney, N., McCann, K., Gellner, G. & Moore, J. C. Structural asymmetry and the stability of diverse food webs. Nature 442, 265–269 (2006).
Pagani-Núñez, E. et al. Niches in the Anthropocene: passerine assemblages show niche expansion from natural to urban habitats. Ecography 42, 1360–1369 (2019).
Magioli, M. et al. Human-modified landscapes alter mammal resource and habitat use and trophic structure. Proc. Natl Acad. Sci. USA 116, 18466–18472 (2019).
Parreira De Castro, D. M. et al. Land use influences niche size and the assimilation of resources by benthic macroinvertebrates in tropical headwater streams. PLoS ONE 11, e0150527 (2016).
Price, E. L., Sertić Perić, M., Romero, G. Q. & Kratina, P. Land use alters trophic redundancy and resource flow through stream food webs. J. Anim. Ecol. 88, 677–689 (2019).
Wang, Y. et al. Trophic structure in response to land use in subtropical streams. Ecol. Indic. 127, 107746 (2021).
van den Hoogen, J. et al. Soil nematode abundance and functional group composition at a global scale. Nature 572, 194–198 (2019).
Phillips, H. R. P. et al. Global distribution of earthworm diversity. Science 366, 480–485 (2019).
Potapov, A. M. et al. Globally invariant metabolism but density–diversity mismatch in springtails. Nat. Commun. 14, 674 (2023).
Tsiafouli, M. A. et al. Intensive agriculture reduces soil biodiversity across Europe. Glob. Change Biol. 21, 973–985 (2015).
Postma-Blaauw, M. B., De Goede, R. G. M., Bloem, J., Faber, J. H. & Brussaard, L. Soil biota community structure and abundance under agricultural intensification and extensification. Ecology 91, 460–473 (2010).
Burdon, F. J., McIntosh, A. R. & Harding, J. S. Mechanisms of trophic niche compression: evidence from landscape disturbance. J. Anim. Ecol. 89, 730–744 (2020).
Potapov, A., Tiunov, A. V. & Scheu, S. Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition. Biol. Rev. 94, 37–59 (2019).
Post, D. M. Using stable isotopes to estimate trophic position: models, methods, and assumptions. Ecology 83, 703–718 (2002).
Jackson, A. L., Inger, R., Parnell, A. C. & Bearhop, S. Comparing isotopic niche widths among and within communities: SIBER—Stable Isotope Bayesian Ellipses in R. J. Anim. Ecol. 80, 595–602 (2011).
Layman, C. A., Arrington, D. A., Montana, C. G. & Post, D. M. Can stable isotope ratios provide for community-wide measures of trophic structure?. Ecology 88, 42–48 (2007).
Marshall, H. H. et al. Stable isotopes are quantitative indicators of trophic niche. Ecol. Lett. 22, 1990–1992 (2019).
Burton, V. J. et al. Effects of land use and soil properties on taxon richness and abundance of soil assemblages. Eur. J. Soil Sci. 74, e13430 (2023).
Nash, L. N. et al. Tropical and temperate differences in the trophic structure and aquatic prey use of riparian predators. Ecol. Lett. 26, 2122–2134 (2023).
Erktan, A., Or, D. & Scheu, S. The physical structure of soil: determinant and consequence of trophic interactions. Soil Biol. Biochem. 148, 107876 (2020).
Steffan, S. A. et al. Microbes are trophic analogs of animals. Proc. Natl Acad. Sci. USA 112, 15119–15124 (2015).
Maraun, M. et al. New perspectives on soil animal trophic ecology through the lens of C and N stable isotope ratios of oribatid mites. Soil Biol. Biochem. 177, 108890 (2023).
Potapov, A. A., Semenina, E. E., Korotkevich, A. Y., Kuznetsova, N. A. & Tiunov, A. V. Connecting taxonomy and ecology: trophic niches of collembolans as related to taxonomic identity and life forms. Soil Biol. Biochem. 101, 20–31 (2016).
Klarner, B. et al. Trophic shift of soil animal species with forest type as indicated by stable isotope analysis. Oikos 123, 1173–1181 (2014).
Korobushkin, D. I., Gongalsky, K. B. & Tiunov, A. V. Isotopic niche (δ13С and δ15N values) of soil macrofauna in temperate forests. Rapid Commun. Mass Spectrom. 28, 1303–1311 (2014).
Wolkovich, E. M. Reticulated channels in soil food webs. Soil Biol. Biochem. 102, 18–21 (2016).
Potapov, A. et al. Towards a global synthesis of Collembola knowledge—challenges and potential solutions. Soil Org. 92, 161–188 (2020).
Zuev, A. et al. Different groups of ground-dwelling spiders share similar trophic niches in temperate forests. Ecol. Entomol. 45, 1346–1356 (2020).
Hunt, H. W. et al. The detrital food web in a shortgrass prairie. Biol. Fertil. Soils 3, 57–68 (1987).
Behmer, S. T. Insect herbivore nutrient regulation. Annu. Rev. Entomol. 54, 165–187 (2009).
Bernays, E. A. & Chapman, R. E. Host–Plant Selection by Phytophagous Insects (Springer, 1994).
Schallhart, N., Tusch, M. J., Wallinger, C., Staudacher, K. & Traugott, M. Effects of plant identity and diversity on the dietary choice of a soil-living insect herbivore. Ecology 93, 2650–2657 (2012).
Yin, R., Siebert, J., Eisenhauer, N. & Schädler, M. Climate change and intensive land use reduce soil animal biomass via dissimilar pathways. eLife 9, e54749 (2020).
Zhou, Z. et al. Plant roots fuel tropical soil animal communities. Ecol. Lett. 26, 742–753 (2023).
Rembold, K., Mangopo, H., Tjitrosoedirdjo, S. S. & Kreft, H. Plant diversity, forest dependency, and alien plant invasions in tropical agricultural landscapes. Biol. Conserv. 213, 234–242 (2017).
Maraun, M., Martens, H., Migge, S., Theenhaus, A. & Scheu, S. Adding to ‘the enigma of soil animal diversity’: fungal feeders and saprophagous soil invertebrates prefer similar food substrates. Eur. J. Soil Biol. 39, 85–95 (2003).
Schneider, K. & Maraun, M. Feeding preferences among dark pigmented fungal taxa (‘Dematiacea’) indicate limited trophic niche differentiation of oribatid mites (Oribatida, Acari). Pedobiologia 49, 61–67 (2005).
Korotkevich, A. Y., Potapov, A. M., Tiunov, A. V. & Kuznetsova, N. A. Collapse of trophic-niche structure in belowground communities under anthropogenic disturbance. Ecosphere 9, e02528 (2018).
Betancur-Corredor, B., Lang, B. & Russell, D. J. Organic nitrogen fertilization benefits selected soil fauna in global agroecosystems. Biol. Fertil. Soils 59, 1–16 (2023).
Crowther, T. W. et al. The global soil community and its influence on biogeochemistry. Science 365, eaav0550 (2019).
Camenzind, T., Hättenschwiler, S., Treseder, K. K., Lehmann, A. & Rillig, M. C. Nutrient limitation of soil microbial processes in tropical forests. Ecol. Monogr. 88, 4–21 (2018).
McGroddy, M. E., Daufresne, T. & Hedin, L. O. Scaling of C:N:P stoichiometry in forests worldwide: implications of terrestrial Redfield-type ratios. Ecology 85, 2390–2401 (2004).
Roslin, T. et al. Higher predation risk for insect prey at low latitudes and elevations. Science 356, 742–744 (2017).
Gauzens, B. et al. Flexible foraging behaviour increases predator vulnerability to climate change. Nat. Clim. Change 14, 387–392 (2024).
Gossner, M. M. et al. Land-use intensification causes multitrophic homogenization of grassland communities. Nature 540, 266–269 (2016).
McKinney, M. L. & Lockwood, J. L. Biotic homogenization: a few winners replacing many losers in the next mass extinction. Trends Ecol. Evol. 14, 450–453 (1999).
Bluhm, S. L. et al. Protura are unique: first evidence of specialized feeding on ectomycorrhizal fungi in soil invertebrates. BMC Ecol. 19, 10 (2019).
Crotty, F. V., Blackshaw, R. P., Adl, S. M., Inger, R. & Murray, P. J. Divergence of feeding channels within the soil food web determined by ecosystem type. Ecol. Evol. 4, 1–13 (2014).
Goncharov, A. A., Khramova, E., Yu & Tiunov, A. V. Spatial variations in the trophic structure of soil animal communities in boreal forests of Pechora-Ilych Nature Reserve. Eur. Soil Sci. 47, 441–448 (2014).
Goncharov, A. A. et al. Sex-related variation in δ15N values of ground beetles (Coleoptera, Carabidae): a case study. Pedobiologia 58, 147–151 (2015).
Goncharov, A. A., Tsurikov, S. M., Potapov, A. M. & Tiunov, A. V. Short-term incorporation of freshly fixed plant carbon into the soil animal food web: field study in a spruce forest. Ecol. Res. 31, 923–933 (2016).
Hyodo, F., Takematsu, Y., Matsumoto, T., Inui, Y. & Itioka, T. Feeding habits of Hymenoptera and Isoptera in a tropical rain forest as revealed by nitrogen and carbon isotope ratios. Insect. Soc. 58, 417–426 (2011).
Hyodo, F. et al. Stable isotope analysis reveals the importance of plant-based diets for tropical ant-mimicking spiders. Entomol. Sci. 21, 461–468 (2018).
Klarner, B. et al. Trophic niches, diversity and community composition of invertebrate top predators (Chilopoda) as affected by conversion of tropical lowland rainforest in Sumatra (Indonesia). PLoS ONE 12, e0180915 (2017).
Korobushkin, D. I. Role of allochthonous carbon in the energy of terrestrial invertebrate communities at different distances from the Black Sea and a freshwater lake (isotopic evidence). Russ. J. Ecol. 45, 223–230 (2014).
Korobushkin, D. I. et al. Consumption of aquatic subsidies by soil invertebrates in coastal ecosystems. Contemp. Probl. Ecol. 9, 396–406 (2016).
Korobushkin, D. I. et al. Are there different trophic niches of enchytraeids? A stable isotopic (δ13C, δ15N) evidence. Soil Biol. Biochem. 194, 109422 (2024).
Li, Z. et al. Incorporation of root-derived carbon into soil microarthropods varies between cropping systems. Biol. Fertil. Soils 56, 839–851 (2020).
Melody, C., Griffiths, B., Dyckmans, J. & Schmidt, O. Stable isotope analysis (δ13C and δ15N) of soil nematodes from four feeding groups. PeerJ 4, e2372 (2016).
Okuzaki, Y., Tayasu, I., Okuda, N. & Sota, T. Vertical heterogeneity of a forest floor invertebrate food web as indicated by stable-isotope analysis. Ecol. Res. 24, 1351–1359 (2009).
Pollierer, M. M., Langel, R., Scheu, S. & Maraun, M. Compartmentalization of the soil animal food web as indicated by dual analysis of stable isotope ratios (15N/14N and 13C/12C). Soil Biol. Biochem. 41, 1221–1226 (2009).
Potapov, A. M., Semenyuk, I. I. & Tiunov, A. V. Seasonal and age-related changes in the stable isotope composition (15N/14N and 13C/12C) of millipedes and collembolans in a temperate forest soil. Pedobiologia 57, 215–222 (2014).
Salamon, J., Wissuwa, J., Frank, T., Scheu, S. & Potapov, A. M. Trophic level and basal resource use of soil animals are hardly affected by local plant associations in abandoned arable land. Ecol. Evol. 10, 8279–8288 (2020).
Scheunemann, N., Scheu, S. & Butenschoen, O. Incorporation of decade old soil carbon into the soil animal food web of an arable system. Appl. Soil Ecol. 46, 59–63 (2010).
Scheunemann, N., Digel, C., Scheu, S. & Butenschoen, O. Roots rather than shoot residues drive soil arthropod communities of arable fields. Oecologia 179, 1135–1145 (2015).
Seeber, J. et al. Abundance and trophic structure of macro-decomposers on alpine pastureland (Central Alps, Tyrol): effects of abandonment of pasturing. Pedobiologia 49, 221–228 (2005).
Seeber, J., Langel, R., Meyer, E. & Traugott, M. Dwarf shrub litter as a food source for macro-decomposers in alpine pastureland. Appl. Soil Ecol. 41, 178–184 (2009).
Steinwandter, M., Rief, A., Scheu, S., Traugott, M. & Seeber, J. Structural and functional characteristics of high alpine soil macro-invertebrate communities. Eur. J. Soil Biol. 86, 72–80 (2018).
Tanaka, H. O., Haraguchi, T. F., Tayasu, I. & Hyodo, F. Stable and radio-isotopic signatures reveal how the feeding habits of ants respond to natural secondary succession in a cool-temperate forest. Insect. Soc. 66, 37–46 (2019).
Kempson, D., Lloyd, M. & Ghelardi, R. A new extractor for woodland litter. Pedobiologia 3, 1–21 (1963).
Potapov, A., Scheu, S. & Tiunov, A. V. Trophic consistency of supraspecific taxa in below-ground invertebrate communities: comparison across lineages and taxonomic ranks. Funct. Ecol. 33, 1172–1183 (2019).
Tsurikov, S. M., Goncharov, A. A. & Tiunov, A. V. Intra-body variation and ontogenetic changes in the isotopic composition (13C/12C and 15N/14N) of beetles (Coleoptera). Entmol. Rev. 95, 326–333 (2015).
Langel, R. & Dyckmans, J. Combined 13C and 15N isotope analysis on small samples using a near-conventional elemental analyzer/isotope ratio mass spectrometer setup: combined 13C and 15N isotope analysis on small samples via μEA/IRMS. Rapid Commun. Mass Spectrom. 28, 1019–1022 (2014).
R Core Team. R: A Language and Environment for Statistical Computing (R Foundation for Statistical Computing, 2020).
Lenth, R. V. emmeans: Estimated marginal means, aka least-squares means. R version 4.1.0 (2021).
Breiman, L. Random forests. Mach. Learn. 45, 5–32 (2001).
Shipley, B. Confirmatory path analysis in a generalized multilevel context. Ecology 90, 363–368 (2009).
Bates, D., Mächler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).
Lefcheck, J. S. piecewiseSEM: piecewise structural equation modelling in R for ecology, evolution, and systematics. Methods Ecol. Evol 7, 573–579 (2016).
Wickham, H. Ggplot2: Elegant Graphics for Data Analysis (Springer-Verlag, 2016).
Greater trophic diversity of soil animal communities under land use and warmer climate. Figshare https://figshare.com/s/c4a378183d4d35e982d1 (2026).