Integrating earthworm movement and life history through dynamic energy budgets.

Earthworms are considered ecosystem engineers and, as such, they are an integral part of the soil ecosystem. The movement of earthworms is significantly influenced by environmental factors such as temperature and soil properties. As movement may directly be linked to food ingestion, especially of endogeic species like , changes in those environmental factors also affect life history traits such as growth and reproduction. In our laboratory studies, earthworms showed a decrease in burrowing activity with decreasing moisture levels and, to some extent, the organic matter content. The burrowing activity of earthworms was also affected by temperature, for which the casts produced per earthworm was used as a proxy in laboratory experiments. We integrated changes in earthworm movement and life histories in response to temperature, soil organic matter content and the moisture level, as observed in our experiment and reported in the literature, through dynamic energy budget (DEB) modelling. The joint parametrization of a DEB model for based on movement and life history data revealed that food ingestion via movement is an integral part of the earthworms' energy budgets. Our findings highlight the importance of soil properties to be considered in the model development for earthworms. Furthermore, by understanding and incorporating the effect of environmental factors on the physiology, this mechanistic approach can help assess the impact of environmental changes such as temperature rise or drought.