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Seminatural grasslands in Europe are susceptible to habitat destruction and fragmentation that result in negative effects on biodiversity because of increased isolation and area effects on extinction rate. However even small habitatpatches of seminatural grasslands might be of value for conservation and restoration of species richness in a landscape with a long history of management, which has been argued to lead to high species richness. We tested whether ant communities have been negatively affected by habitat loss and increased isolation of seminatural grasslands during the twentieth century. We examined species richness and community composition in seminatural grasslands of different size in a mosaic landscape in Central Sweden. Grasslands managed continuously over centuries harbored species-rich and ecologically diverse ant communities. Grassland remnant size had no effect on ant species richness. Small grassland remnants did not harbor a nested subset of the ant species of larger habitats. Community composition of ants was mainly affected by habitat conditions. Our results suggest that the abandonment of traditional land use and the encroachment of trees, rather than the effects of fragmentation, are important for species composition in seminatural grasslands. Our results highlight the importance of considering land-use continuity and dispersal ability of thefocal organisms when examining the effects of habitat loss and fragmentation on biodiversity. Landscape history should be considered in conservation programs focusing on effects of land-use change.

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Understanding and disentangling different processes underlying the assembly and diversity of communities remains a key challenge in ecology. Species can assemble into communities either randomly or due to deterministic processes. Deterministic assembly leads to species being more similar (underdispersed) or more different (overdispersed) in certain traits than would be expected by chance. However, the relative importance of those processes is not well understood for many organisms, including terrestrial invertebrates. Based on knowledge of a broad range of species traits, we tested for the presence of trait underdispersion (indicating dispersal or environmental filtering) and trait overdispersion (indicating niche partitioning) and their relative importance in explaining land snail community composition on lake islands. The analysis of community assembly was performed using a functional diversity index (Rao's quadratic entropy) in combination with a null model approach. Regression analysis with the effect sizes of the assembly tests and environmental variables gave information on the strength of under- and overdispersion along environmental gradients. Additionally, we examined the link between community weighted mean trait values and environmental variables using a CWM-RDA. We found both trait underdispersion and trait overdispersion, but underdispersion (eight traits) was more frequently detected than overdispersion (two traits). Underdispersion was related to four environmental variables (tree cover, habitat diversity, productivity of ground vegetation, and location on an esker ridge). Our results show clear evidence for underdispersion in traits driven by environmental filtering, but no clear evidence for dispersal filtering. We did not find evidence for overdispersion of traits due to diet or body size, but overdispersion in shell shape may indicate niche differentiation between snail species driven by small-scale habitat heterogeneity. The use of species traits enabled us to identify key traits involved in snail community assembly and to detect the simultaneous occurrence of trait underdispersion and overdispersion.