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Implementation of the EU Water Framework Directive (WFD) has drawn much attention to hydromorphological alterations of surface waters. The Lake Habitat Survey (LHS) protocol provides a method for characterising and assessing the physical habitats of lakes and reservoirs. Two metrics were developed based on this method: the Lake Habitat Modification Score (LHMS) and the Lake Habitat Quality Assessment (LHQA), as measures of lake modification and habitat value, respectively. However, the use of these metrics to predict measures of ecological quality remains largely untested. Thus, we assessed the relationships between LHS metrics and the littoral macroinvertebrate community in 42 lakes across Europe. A significant relationship was found between littoral macrophyte descriptors and riparian natural land cover variables of the LHQA score and macroinvertebrate community composition in 2 out of 4 European regions. No relationship was found between macroinvertebrate community composition and the LHMS. Some significant correlations were found between selected macroinvertebrate metrics and the LHS scores, but this pattern was not consistent across regions, and no relationship was found with the overall LHMS or LHQA scores. This demonstrates that the LHS metrics do not consistently predict the quality of littoral macroinvertebrate communities across Europe, and a region specific approach may be necessary. However, we could demonstrate a relationship between the site specific LHS variables and the macroinvertebrate community at the site level, and in some cases at the regional level. Therefore, although the LHS metrics do not appear to be a useful for relating habitat quality and pressure to littoral macroinvertebrate communities, selected LHS variables may exhibit stronger relationships with the biota.

Summary: Beta diversity is the compositional heterogeneity of biotic assemblages among sites, and biotic homogenization is the decrease in beta diversity, facilitated by an increase in similarity of biotic communities over time. Environmental harshness decreases the importance of stochastic processes in structuring assemblages, resulting in a homogenization of the biota. We investigated if increasing nutrient enrichment, land-use pressure, and within-lake habitat alteration would decrease the beta diversity of macroinvertebrates in 46 lakes across Europe. Beta diversity was calculated using global multivariate dispersion. We utilized a structural equation modelling approach to account for hierarchical interdependence between potential impacts, that is the direct effects and correlations among the different impacts. We found clear indications that European macroinvertebrate communities are being homogenized by ongoing lake shore development. Increasing land-use pressure in the form of residential and commercial development had a direct negative effect on beta diversity (standardized coefficient = -0·40), as did roadways, albeit indirectly through an increase in engineering structures (standardized coefficient = -0·31). Increasing within-lake silt levels also homogenized macroinvertebrate communities (standardized coefficient = -0·18), independent of near shore land use. Our results indicate the negative effect of both the near shore land-use pressure and the within-lake habitat alteration on macroinvertebrate beta diversity, with significant interactions between these pressures. Habitat protection should take a more holistic approach to assessing lake development pressure, over a range of scales, as a solely site specific approach is not always biologically meaningful. Thus, future management plans should carefully control and mitigate ongoing development pressure if lake ecosystem health and resilience is to be maintained. Synthesis and applications. This study is the first of its kind to demonstrate European-wide homogenization of littoral macroinvertebrate lake communities with increasing habitat alteration and land-use pressure. Significant interactions occur between different habitat scales, with no one scale entirely accounting for the homogenization effect. To avoid further biotic homogenization, development pressure must be carefully managed at multiple scales, and where possible, minimized. This presents a challenge, as globally there is an increasing expansion of the human population and a consequent increase in anthropogenic pressure across all habitats. This study is the first of its kind to demonstrate European-wide homogenization of littoral macroinvertebrate lake communities with increasing habitat alteration and land-use pressure. Significant interactions occur between different habitat scales, with no one scale entirely accounting for the homogenization effect. To avoid further biotic homogenization, development pressure must be carefully managed at multiple scales, and where possible, minimized. This presents a challenge, as globally there is an increasing expansion of the human population and a consequent increase in anthropogenic pressure across all habitats.

Besides pollution, lakes are affected by human alterations of lake-shore morphology. However, ecological effects of such alterations have rarely been studied systematically. Hence, we developed tools to assess the ecological effects of anthropogenic morphological alterations on European lake-shores based on pressure-specific response patterns of littoral macroinvertebrate community composition. Littoral invertebrates were sampled from 51 lakes in seven European countries. Sampling covered a range of natural to heavily morphologically degraded sites including natural shorelines, recreational beaches, ripraps and retaining walls. Biological data were supplemented by standardized morphological data that were collected via a Lake Habitat Survey (LHS) protocol and subsequently used to develop a morphological stressor index. Two biotic multimetric indices were developed based on habitat-specific samples (Littoral Invertebrate Multimetric based on HAbitat samples, LIMHA) and composite samples (Littoral Invertebrate Multimetric based on COmposite samples, LIMCO) through correlations with the morphological stressor index. Similarity analyses showed strong spatial differences in macroinvertebrate community composition between four main geographical regions, i.e. Western, Northern, Central and Southern Europe. The morphological stressor index as well as LIMCO and LIMHA have been developed for each geographical region specifically, thereby optimizing correlations of LIMCO and LIMHA with the respective morphological stressor index. The metric composition of LIMCO and LIMHA and their correlation coefficients with the morphological stressor index are comparable to existing national and regional methods that assess morphological lakeshore degradation via macroinvertebrate communities. Hence, LIMCO and LIMHA indices constitute a new stressor-specific assessment tool that enables comparable lake morphology assessment across Europe, as it has been developed involving a uniform methodology followed by regionalized optimization. These tools fulfil the standards of the EU Water Framework Directive and thus may complement existing assessment approaches used in lake monitoring focusing solely on lake eutrophication so far.