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Summary1. In recent years, river restoration science has been searching for biological indicators of improvement in the physical habitats of streams. To date, research has mainly focused on the use of fish and macroinvertebrates as indicators. Despite their importance in aquatic ecosystems, the response of macrophytes to habitat restoration has been rarely studied. 2. We investigated the macrophyte communities of 40 restored river reaches in the lowland and lower mountainous areas of Germany. Each restored reach was compared to an upstream, unrestored reach using a space-for-time-substitution approach. At each reach, a 100 m stretch was surveyed for submerged and emergent macrophytes, recording the quantity, abundance and growth form of each species. Additionally, microhabitat patterns (substrate, depth, current velocity) and channel parameters (mean and bankfull width, number of channel elements) were recorded. 3. Restored reaches had a significantly higher macrophyte cover, richness, diversity and number of growth forms. Macrophyte diversity and richness were both positively correlated with depth, current and substrate. 4. The analysis of growth forms showed that Lemnids, Helodids, Parvopotamids, Elodids, Peplids and Juncids are all significant indicators of restoration. These species all responded directly to the restoration measures either by highly increasing in abundance or by being present in the restored reaches and absent in the unrestored reaches. While the restored reaches of the lowland rivers were characterized by a high abundance of Peplids and Parvopotamids, the restored reaches of the mountain rivers showed a significantly higher presence and abundance of Lemnids and Helodids. 5. Three macrophyte species (Lemna minor, Persicaria hydropiper, Potamogeton crispus) were regarded as significant indicators of restoration. No species were found to be indicators of unrestored reaches. 6. Synthesis and applications. Macrophyte communities benefit from river restoration by showing increased cover, abundance and diversity. The main drivers of this enhancement are more natural and diverse substrates and an increased floodplain area in the restored reaches, as well as a greater variability of current and depth patterns. Monitoring of macrophytes could thus be an easy and cost-effective means to gauge the success of river restoration measures.

1. Restoration of river hydromorphology often has limited detected effects on river biota. One frequently discussed reason is that the restored river length is insufficient to allow populations to develop and give the room for geomorphological processes to occur. 2. We investigated ten pairs of restored river sections of which one was a large project involving a long, intensively restored river section and one represented a smaller restoration effort. The restoration effect was quantified by comparing each restored river section to an upstream nonrestored section. We sampled the following response variables: habitat composition in the river and its floodplain, three aquatic organism groups (aquatic macrophytes, benthic invertebrates and fish), two floodplain-inhabiting organism groups (floodplain vegetation, ground beetles), as well as food web composition and land-water interactions reflected by stable isotopes. 3. For each response variable, we compared the difference in dissimilarity of the restored and nearby non-restored section between the larger and the smaller restoration projects. In a second step, we regrouped the pairs and compared restored sections with large changes in substrate composition to those with small changes. 4. When comparing all restored to all non-restored sections, ground beetles were most strongly responding to restoration, followed by fish, floodplain vegetation, benthic invertebrates and aquatic macrophytes. Aquatic habitats and stable isotope signatures responded less strongly. 5.When grouping the restored sections by project size, there was no difference in the response to restoration between the projects targeting long and short river sections with regard to any of the measured response variables except nitrogen isotopic composition. In contrast, when grouping the restored sections by substrate composition, the responses of fish, benthic invertebrates, aquatic macrophytes, floodplain vegetation and nitrogen isotopic composition were greater in sections with larger changes in substrate composition as compared to those with smaller changes. 6. Synthesis and applications. The effects of hydromorphological restoration measures on aquatic and floodplain biota strongly depend on the creation of habitat for aquatic organisms, which were limited or not present prior to restoration. These positive effects on habitats are not necessarily related to the restored river length. Therefore, we recommend a focus on habitat enhancement in river restoration projects.

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