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.

A N Severtsov Institute of Ecology and Evolution Russian Academy of Sciences Leninsky Prospect 33 119071 Moscow Russia

Biology Centre CAS Institute of Entomology Branisovska 1160 31 370 05 Ceske Budejovice Czech Republic

Biology Centre of the Czech Academy of Sciences Institute of Soil Biology Na Sádkách 702 7 37005 České Budějovice Czech Republic

Canadian National Collection of Insects Arachnids and Nematodes Agriculture and Agri Food Canada Ottawa ON K1A 0C6 Canada

Centre of Biodiversity and Sustainable Land Use Büsgenweg 1 37077 Göttingen Germany

Colecciones Entomológicas Torres Sala Servei de Patrimoni Històric Ajuntament de València València Spain

Departament de Didàctica de les Cièncias Experimentals i Socials Facultat de Magisteri Universitat de València València Spain

Department of Crop and Soil Science Oregon State University Corvallis OR 97331 U S A

Department of Ecology Brandenburg University of Technology Karl Wachsmann Allee 6 03046 Cottbus Germany

Department of Nematology Wageningen University and Research 6700ES Wageningen The Netherlands

Institute of Systematics and Evolution of Animals PAS Slawkowska 17 Pl 31 016 Krakow Poland

J F Blumenbach Institute of Zoology and Anthropology University of Göttingen Untere Karspüle 2 37073 Göttingen Germany

Joint Russian Vietnamese Tropical Center №3 Street 3 Thang 2 Q10 Ho Chi Minh City Vietnam

State Museum Natural History of NAS of Ukraine Teatralna 18 79008 Lviv Ukraine

UCD School of Agriculture and Food Science University College Dublin Belfield Dublin 4 Ireland

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