Community structure or function: effects of environmental stress on benthic macroinvertebrates at different spatial scales

Summary 1. This study investigated the relation of benthic macroinvertebrates to environmental gradients in Central European lowland rivers. Taxonomic structure (taxa) and functional composition (metrics) were related to gradients at four different spatial scales (ecoregion, catchment, reach and site). The environmental variables at the catchment‐, reach‐ and site scales reflected the intensity of human impact: catchment and floodplain land use, riparian and floodplain degradation, flow regulation and river bank and bed modification. 2. Field surveys and GIS yielded 130 parameters characterising the hydromorphology and land use of 75 river sections in Sweden, the Netherlands, Germany and Poland. Two hundred and forty‐four macroinvertebrate taxa and 84 derived community metrics and biotic indices such as functional guilds, diversity and composition measures were included in the analysis. 3. Canonical Correspondence Analysis (CCA) and Redundancy Analysis (RDA) showed that hydromorphological and land use variables explained 11.4%, 22.1% and 15.8% of the taxa variance at the catchment (‘macro’), reach (‘meso’) and site (‘micro’) scales, respectively, compared with 14.9%, 33.2% and 21.5% of the variance associated with the derived metrics. Ecoregion and season accounted for 10.9% and 20.5% of the variance of the taxonomic structure and functional composition, respectively. 4. Partial CCA (pCCA) and RDA (pRDA) showed that the unique variance explained was slightly higher for taxa than for metrics. By contrast, the joint variance explained for metrics was much higher at all spatial scales and largest at the reach scale. Environmental variables explained 46.8% of metric variance and 32.4% of taxonomic structure. 5. Canonical Correspondence Analysis and RDA identified clear environmental gradients along the two main ordination axes, namely, land use and hydromorphological degradation. The impact of catchment land use on benthic macroinvertebrates was mainly revealed by the proportion of urban areas. At the reach scale, riparian and floodplain attributes (bank fixation, riparian wooded vegetation, shading) and the proportion of large woody debris were strong predictors of the taxonomic structure and functional composition of benthic macroinvertebrates. At the site scale, artificial substrata indicated human impact, particularly the proportion of macro‐ and mesolithal used for bank enforcement (rip–rap). 6. Our study revealed the importance of benthic macroinvertebra