Principal determinants of species and functional diversity of carabid beetle assemblages during succession at post-industrial sites.

Although ecological succession is one of the principal focuses of recent restoration ecology research, it is still unclear which factors drive this process and positively influence species richness and functional diversity. In this study we sought to elucidate how species traits and functional diversity change during forest succession, and to identify important factors that determine the species in the observed assemblages. We analyzed species richness and functional diversity of ground beetle assemblages in relation to succession on post-industrial localities after habitat deterioration caused by spoil deposition. We selected ground beetles as they are known to be sensitive to landscape changes (with a large range of responses), and their taxonomy and ecology are generally well-known. Ground beetles were sampled on the spoil heaps during the last 30 years when spontaneous succession occurred. To calculate functional diversity, we used traits related to habitat and trophic niche, i.e. food specialization, wing morphology, trophic level, and bio-indication value. Ground beetle species were found to be distributed non-randomly in the assemblages in the late phase of succession. Ordination analyses revealed that the ground beetle assemblage was significantly associated with the proportion of forested area. Environmental heterogeneity generated assemblages that contained over-dispersed species traits. Our findings indicated that environmental conditions at late successional stages supported less mobile carnivorous species. Overall, we conclude that the decline in species richness and functional diversity in the middle of the studied succession gradient indicated that the assemblages of open habitats had been replaced by species typical of forest ecosystems.