Species co-occurrence networks of ground beetles in managed grasslands

Grassland biodiversity, including traditional rural biotopes maintained by traditional agricultural practices, has become threatened worldwide. Road verges have been suggested to be complementary or compensatory habitats for species inhabiting grasslands. Species co-occurrence patterns linked with species traits can be used to separate between the different mechanisms (stochasticity, environmental filtering, biotic interactions) behind community structure. Here, we study species co-occurrence networks and underlying mechanisms of ground beetle species (Carabidae) in three different managed grassland types (meadows, pastures, road verges, n = 12 in each type) in Central Finland. We aimed to find out whether road verges can be considered as compensatory to traditional rural biotopes (meadows and pastures). We found that stochasticity explained over 90% of the pairwise co-occurrences, and the non-random co-occurrences were best explained by environmental filtering, regardless of the grassland type. However, the identities and traits of the species showing non-random co-occurrences differed among the habitat types. Thus, environmental factors behind environmental filtering differ among the habitat types and are related to the site-specific characteristics and variation therein. This poses challenges to habitat management since the species' response to management action may depend on the site-specific characteristics. Although road verges are not fully compensatory to meadows and pastures, the high similarity of species richness and the high level of shared species suggest that for carabids road verges may be corridors connecting the sparse network of the remaining traditional rural biotopes.