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Summary1. 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. 2. 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. 3. 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 = À0Á40), as did roadways, albeit indirectly through an increase in engineering structures (standardized coefficient = À0Á31). Increasing within-lake silt levels also homogenized macroinvertebrate communities (standardized coefficient = À0Á18), independent of near shore land use. Our results indicate the negative effect of both the near shore landuse pressure and the within-lake habitat alteration on macroinvertebrate beta diversity, with significant interactions between these pressures. 4. 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. 5. 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.

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ABSTRACT1. Lake Habitat Survey (LHS) provides a standard method for characterizing the physical habitat of lakes and reservoirs, but has not been tested for its relevance to the composition and abundance of macroinvertebrates. This study investigated the relationship between the metrics used in LHS and components of macroinvertebrate communities found in the littoral zone of a shallow calcareous lake in the west of Ireland.2. A scoring system, the Habitat Quality Assessment (HabQA), developed from the Lake Habitat Quality Assessment (LHQA) of the LHS, was used to assess the relationship between habitat quality based on physical structure within 10 LHS 'habplots' and metrics of the macroinvertebrate community.3. Macroinvertebrate taxon richness, both of adults found in the riparian zone and larvae found in the littoral zone, correlated positively with the HabQA score. Macrophytes within the littoral zone, and complexity of riparian vegetation within the riparian zone, were particularly important in driving the HabQA score. While overall abundance of macroinvertebrates did not vary with HabQA score, that of particular genera did.4. The HabQA score was a useful surrogate of taxon richness for adult and larval aquatic macroinvertebrates, suggesting that, in general, LHS provides a useful conservation assessment tool relevant for macroinvertebrates. However, in some circumstances, such as wave-washed stony substrates devoid of macrophytes, the HabQA score may not capture the quality of a site for macroinvertebrates, and the importance of natural but low diversity sites should not be neglected in conservation assessment of lakes. Similarly, while the LHS method notes the presence of alien species, further work on how these could be incorporated into the method would be useful.5. Reliance on a single, or overall combined, metric score across quality elements, whether based on biotic or structural assessment, has some potential limitations. It is clear that for conservation management a holistic assessment of naturalness, representativeness and species rarity needs to be made in conjunction with scoring systems.

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ABSTRACT1. Morphological degradation constitutes one of the most severe threats to the ecological integrity of lakes. The development of biotic assessment methods for human lake shore alterations using littoral macroinvertebrates requires quantification of the degree of degradation by a stressor index and is complicated through simultaneous physical pressures that alter natural habitat structure.2. The Lake Habitat Survey (LHS) method and macroinvertebrate sampling were used to produce a pan-European dataset of morphological lake shore degradation and macroinvertebrate densities covering 51 lakes in seven countries and across four geographical regions -northern, western, southern and central Europe.3. Lake Habitat Survey parameters that differed significantly among three categories of morphological pressure were combined to develop the stressor index components 'Number of habitats', 'Habitat diversity', 'Total percentage volume inhabited by macrophytes', 'Sum of macrophyte types', 'Sum of vegetation cover types', 'Sum of coarse woody debris/roots/overhanging vegetation', 'Pressure index' (number of human disturbance sources) and 'Natural/artificial dominant land cover type '. 4. Stressor index components were tested for cross-correlations and for differences among pressure levels. The final composition of the stressor index was optimized for the four studied geographical regions in Europe. The resulting stressor index correlated more strongly with macroinvertebrate metrics than simpler site-specific LHS parameters or the HabQA index developed previously in one lake in north-western Europe.5. The stressor index developed provides deeper insight into the morphological pressures that affect littoral invertebrate communities. The results also support the use of LHS to quantify morphological stressors at sampling site level, which can ease developing other multimetric bioassessment methods.6. The stressor index offers the possibility for wide and regional specific application to assess hydromorphological pressures on lakes to assist conservation planning and management and further global efforts to develop and test biotic assessment methods for lakes.

Morphological alteration of shorelines and eutrophication both affect the biological integrity of European lakes. These pressures, often acting simultaneously, are difficult to tease apart. In this study, we related the number of taxa with specific habitat preference to habitat complexity across lakes of varying nutrient state. Habitat complexity at morphologically altered shorelines was significantly lower than at unaltered sites across trophic categories. A generalised linear mixed‐effects model showed decreased number of taxa with specific mesohabitat preference at morphologically simplified sites in oligotrophic and mesotrophic, but not eutrophic lakes. These results suggest: (1) an antagonistic interaction between the effect of nutrient enrichment and morphological alterations on lake littoral communities and (2) the number of macroinvertebrate habitat specialists could potentially be used to assess the effects of structural simplifications of shorelines in lakes of low to medium nutrient status. We conclude that the use of functional traits approach in aquatic ecology should foster better understanding of stressor–response relationships for combined effect of multiple stressors.