Stable isotopes revisited: Their use and limits for oribatid mite trophic ecology

In this review we summarize our knowledge of using stable isotopes (15N/14N, 13C/12C) to better understand the trophic ecology of oribatid mites. Our aims are (a) to recapitulate the history of stable isotope research in soil animals with a focus on oribatid mites, (b) to present new stable isotope data for oribatid mites and overview the current state of knowledge of oribatid mite trophic niche differentiation, (c) to compile problems and limitations of stable isotope based analyses of trophic relationships and (d) to suggest future challenges, questions and problems that may be solved using stable isotope analyses and other novel techniques for improving our understanding on the trophic ecology of soil invertebrates. We conclude that (1) in addition to 15N/14N ratios, 13C/12C ratios contribute to our understanding of the trophic ecology of oribatid mites, allowing, e.g. separation of lichen- and moss-feeding species, (2) there likely are many lichen but few moss feeding oribatid mite species, (3) oribatid mite species that are endophagous as juveniles are separated by their stable isotope signatures from all other oribatid mite species, (4) fungivorous oribatid mite species cannot be separated further, e.g. the fungal taxa they feed on cannot be delineated. A particular problem in using stable isotope data is the difficulty in determining signatures for basal food resources, since decomposing material, fungi and lichens comprise various components differing in stable isotope signatures; 13C/12C ratios and potentially other isotopes may help in identifying the role of these resources for decomposer animal nutrition.