Pressures at larger spatial scales strongly influence the ecological status of heavily modified river water bodies in Germany.

River biota are influenced by anthropogenic pressures that operate at different spatial scales. Understanding which pressures at which spatial scales affect biota is essential to manage and restore degraded rivers. In Europe, many river reaches were designated as Heavily Modified Water Bodies (HMWB) according to the European Water Framework Directive (WFD), where the ecological potential might mainly be determined by pressures at larger spatial scales outside the HMWB (e.g. hydromorphological alterations at the river network and land use at the catchment scale). In Germany, hydromorphological alterations and diffuse pollution were the main pressures. Therefore, the three objectives of this study were to (i) identify the hydromorphological pressures at the site, reach, and river network scale, and land use categories at the catchment scale which significantly affect the ecological status of HMWB in Germany, (ii) quantify the relative importance of these pressures at different spatial scales, and (iii) analyse the differences in response between fish and macroinvertebrates. The results indicated that: (i) At the reach scale, fish were most strongly influenced by channel-bank conditions whilst the naturalness of channel-planform was the best proxy for the ecological status of macroinvertebrates. At the catchment scale, urbanization was the most detrimental land use. (ii) The pressures at larger spatial scales (catchment land use and hydromorphological alterations in the river network) generally were more important than hydromorphological alterations at the reach scale. (iii) Fish were affected equally by both, hydromorphological alterations at the reach scale and large-scale pressures whereas the latter were far more important for the ecological status of macroinvertebrates. In conclusion, these results indicated that large-scale pressures may often limit the efficiency of reach-scale restoration, especially for macroinvertebrates, even in the absence of saprobic pollution, and have to be considered for the management and restoration of HMWB in Germany and comparable degraded river reaches.