Hierarchical response of littoral macroinvertebrates to altered hydromorphology and eutrophication

The composition of littoral macroinvertebrate communities in lakes is governed by multiple natural and anthropogenic environmental influences interacting at different spatial scales. Since ecological assessment methods using littoral macroinvertebrates should respond specifically to a single stressor, knowledge on the unique effects of a given stressor is necessary. To effectively disentangle the effects of hydromorphology and trophic state requires analysing macroinvertebrate communities at lake sites with the full range of both stressors. We used a dataset of 98 lakes encompassing the entire gradient of geographical locations, lake types, hydromorphological degradation and trophic states in Central European lakes. We studied the unique and joint effects of hydromorphology and trophic state on macroinvertebrate richness, community composition and the Littoral Invertebrate Multimetric Index based on Composite Sampling (LIMCO). Variation partitioning analyses were conducted to test the importance of hydromorphology relative to trophic state across and within hydromorphological states (natural shorelines, hard and soft shore modifications) and trophic states (oligotrophic to hypertrophic states). At natural, hard and soft modification sites, hydromorphology explained 10, 16 and 19%, respectively, of the average unique variation of diversity, community composition and the LIMCO index, whereas trophic state explained on average 2, 5 and 5%, respectively. Similarly, in low, medium and high trophic state lakes, hydromorphology explained 10, 15 and 7%, respectively, of the average unique variation of diversity, community composition and the LIMCO index, whereas trophic state explained on average 0.3, 3 and 6%, respectively. Our results demonstrate that littoral hydromorphology was a more important driver of macroinvertebrate diversity, community composition and LIMCO than trophic state across hydromorphological states and trophic states. This indicates that multiple stressors in lakes act hierarchically on littoral macroinvertebrate communities and that the hydromorphological degradation of littoral zones is the primary driver for altered communities.