Soil organisms drive major ecosystem functions by mineralising carbon and releasing nutrients during decomposition processes, which supports plant growth, aboveground biodiversity and, ultimately, human nutrition. Soil ecologists often operate with functional groups to infer the effects of individual taxa on ecosystem functions and services. Simultaneous assessment of the functional roles of multiple taxa is possible using food‐web reconstructions, but our knowledge of the feeding habits of many taxa is insufficient and often based on limited evidence. Over the last two decades, molecular, biochemical and isotopic tools have improved our understanding of the feeding habits of various soil organisms, yet this knowledge is still to be synthesised into a common functional framework. Here, we provide a comprehensive review of the feeding habits of consumers in soil, including protists, micro‐, meso‐ and macrofauna (invertebrates), and soil‐associated vertebrates. We have integrated existing functional group classifications with findings gained with novel methods and compiled an overarching classification across taxa focusing on key universal traits such as food resource preferences, body masses, microhabitat specialisation, protection and hunting mechanisms. Our summary highlights various strands of evidence that many functional groups commonly used in soil ecology and food‐web models are feeding on multiple types of food resources. In many cases, omnivory is observed down to the species level of taxonomic resolution, challenging realism of traditional soil food‐web models based on distinct resource‐based energy channels. Novel methods, such as stable isotope, fatty acid and DNA gut content analyses, have revealed previously hidden facets of trophic relationships of soil consumers, such as food assimilation, multichannel feeding across trophic levels, hidden trophic niche differentiation and the importance of alternative food/prey, as well as energy transfers across ecosystem compartments. Wider adoption of such tools and the development of open interoperable platforms that assemble morphological, ecological and trophic data as traits of soil taxa will enable the refinement and expansion of the multifunctional classification of consumers in soil. The compiled multifunctional classification of soil‐associated consumers will serve as a reference for ecologists working with biodiversity changes and biodiversity–ecosystem functioning relationships, making soil food‐web research more accessible and reproducible.

Land-use change is threatening biodiversity worldwide, affecting above and below ground animal communities by altering their trophic niches. However, shifts in trophic niches with changes in land use are little studied and this applies in particular to belowground animals. Oribatid mites are among the most abundant soil animals, involved in decomposition processes and nutrient cycling. We analyzed shifts in trophic niches of six soil-living oribatid mite species with the conversion of lowland secondary rainforest into plantation systems of different land-use intensity (jungle rubber, rubber and oil palm monoculture plantation) in two regions of southwest Sumatra, Indonesia. We measured stable isotope ratios (13C/12C and 15N/14N) of single oribatid mite individuals and calculated shifts in stable isotope niches with changes in land use. Significant changes in stable isotope ratios in three of the six studied oribatid mite species indicated that these species shift their trophic niches with changes in land use. The trophic shift was either due to changes in trophic level (δ15N values), to changes in the use of basal resources (δ13C values) or to changes in both. The trophic shift generally was most pronounced between more natural systems (rainforest and jungle rubber) on one side and monoculture plantations systems (rubber and oil palm plantations) on the other, reflecting that the shifts were related to land-use intensity. Although trophic niches of the other three studied species did not differ significantly between land-use systems they followed a similar trend. Overall, the results suggest that colonization of very different ecosystems such as rainforest and intensively managed monoculture plantations by oribatid mite species likely is related to their ability to shift their trophic niches, i.e. to trophic plasticity.

Background Ectomycorrhizal fungi (ECM) play a central role in nutrient cycling in boreal and temperate forests, but their role in the soil food web remains little understood. One of the groups assumed to live as specialised mycorrhizal feeders are Protura, but experimental and field evidence is lacking. We used a combination of three methods to test if Protura are specialized mycorrhizal feeders and compared their trophic niche with other soil invertebrates. Using pulse labelling of young beech and ash seedlings we analysed the incorporation of 13 C and 15 N into Acerentomon gallicum. In addition, individuals of Protura from temperate forests were collected for the analysis of neutral lipid fatty acids and natural variations in stable isotope ratios. Results Pulse labelling showed rapid incorporation of root-derived 13 C, but no incorporation of root-derived 15 N into A. gallicum . The transfer of 13 C from lateral roots to ectomycorrhizal root tips was high, while it was low for 15 N. Neutral lipid fatty acid (NLFA) analysis showed high amounts of bacterial marker (16:1ω7) and plant marker (16:0 and 18:1ω9) fatty acids but not of the fungal membrane lipid 18:2ω6,9 in A. gallicum . Natural variations in stable isotope ratios in Protura from a number of temperate forests were distinct from those of the great majority of other soil invertebrates, but remarkably similar to those of sporocarps of ECM fungi. Conclusions Using three in situ methods, stable isotope labelling, neutral lipid fatty acid analysis and natural variations of stable isotope ratios, we showed that Protura predominantly feed on mycorrhizal hyphae via sucking up hyphal cytoplasm. Predominant feeding on ectomycorrhizal mycelia by Protura is an exception; the limited consumption of ECM by other soil invertebrates may contribute to carbon sequestration in temperate and boreal forests. Electronic supplementary material The online version of this article (10.1186/s12898-019-0227-y) contains supplementary material, which is available to authorized users.

The input of plant leaf litter has been assumed to be the most important resource for soil organisms of forest ecosystems, but there is increasing evidence that root-derived resources may be more important. By trenching roots of trees in deciduous and coniferous forests, we cut-off the input of root-derived resources and investigated the response of microorganisms using substrate-induced respiration and phospholipid fatty acid (PLFA) analysis. After one and three years, root trenching strongly decreased microbial biomass and concentrations of PLFAs by about 20%, but the microbial community structure was little affected and the effects were similar in deciduous and coniferous forests. However, the reduction in microbial biomass varied between regions and was more pronounced in forests on limestone soils (Hainich) than in those on sandy soils (Schorfheide). Trenching also reduced microbial biomass in the litter layer but only in the Hainich after one year, whereas fungal and bacterial marker PLFAs as well as the fungal-to-plant marker ratio in litter were reduced in the Schorfheide both after one and three years. The pronounced differences between forests of the two regions suggest that root-derived resources are more important in fueling soil microorganisms of base-rich forests characterized by mull humus than in forests poor in base cations characterized by moder soils. The reduction in microbial biomass and changes in microbial community characteristics in the litter layer suggests that litter microorganisms do not exclusively rely on resources from decomposing litter but also from roots, i.e. from resources based on labile recently fixed carbon. Our results suggest that both bacteria and fungi heavily depend on root-derived resources with both suffering to a similar extent to deprivation of these resources. Further, the results indicate that the community structure of microorganisms is remarkably resistant to changes in resource supply and adapts quickly to new conditions irrespective of tree species composition and forest management.

Forest soil food webs have been assumed to be fueled substantially by root-derived resources. However, until today the flux of root-derived resources into soil animals has been investigated virtually exclusively using isotope labeling experiments, whereas studies on the consequences of disrupting the flux of root-derived resources into the soil animal food web are scarce. We here investigated the importance of root-derived resources for a wide range of soil animals by interrupting the resource flux into the soil of different forest types in Central Europe using a trenching experiment. We recorded the abundance of soil animal taxa varying in body size (micro-, meso-, and macrofauna) 1 and 3 years after root trenching, and quantified changes in biomass, species composition, and trophic shift using stable isotopes and NLFA analysis. Among the microfauna groups studied (trophic groups of Nematoda) only the abundance of plant feeding nematodes showed a trend in being decreased by -58% due to root trenching. Major soil mesofauna groups, including Collembola and Oribatida, suffered to a similar extent from root trenching with their abundance and biomass being reduced by about 30–40%. The soil macrofauna groups studied (Diplopoda, Isopoda, Chilopoda, Araneae, Coleoptera) generally were only little affected by root trenching suggesting that they rely less on root-derived resources than micro- and in particular mesofauna. Notably, the community structure of micro-, meso-, and macrofauna was not affected by root trenching. Further, we observed trophic shifts only in 2 out of 10 investigated species with the shifts generally being only minor. The results indicate that soil animal communities are markedly resilient to deprivation of root-derived resources suggesting that links to root-derived resources are non-specific. However, this resilience appears to vary with body size, with mesofauna including both decomposers as well as predators being more sensitive to the deprivation of root-derived resources than microfauna (except for root feeders) and macrofauna. Overall, this suggests that body size constrains the channeling of energy through soil food webs, with root-derived resources in temperate forests being channeled predominantly via soil taxa of intermediate size, i.e., mesofauna.

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