Perspectives in modelling earthworm dynamics and their feedbacks with abiotic soil properties

Effects of earthworms on soil abiotic properties are well documented from several decades of laboratory and mesocosm experiments, and they are supposed to affect large-scale soil ecosystem functioning. The prediction of the spatiotemporal occurrence of earthworms and the related functional effects in the field or at larger scales, however, is constrained by adequate modelling approaches. Correlative, phenomenological methods, such as species distribution models, facilitate the identification of factors that drive species' distributions. However, these methods ignore the ability of earthworms to select and modify their own habitat and therefore may lead to unreliable predictions. Understanding these feedbacks between earthworms and abiotic soil properties is a key requisite to better understand their spatiotemporal distribution as well as to quantify the various functional effects of earthworms in soil ecosystems. Process-based models that investigate either effects or responses of earthworms on soil environmental conditions are mostly applied in ecotoxicological and bioturbation studies. Process-based models that describe feedbacks between earthworms and soil abiotic properties explicitly are rare. In this review, we analysed 18 process-based earthworm dynamic modelling studies pointing out the current gaps and future challenges in feedback modelling. We identify three main challenges: (i) adequate and reliable process identification in model development at and across relevant spatiotemporal scales (individual behaviour and population dynamics of earthworms), (ii) use of information from different data sources in one model (laboratory or field experiments, earthworm species or functional type) and (iii) quantification of uncertainties in data (e.g. spatiotemporal variability of earthworm abundances and soil hydraulic properties) and derived parameters (e.g. population growth rate and hydraulic conductivity) that are used in the model.