The Influence of Matrix and Edges on Species Richness Patterns of Ground Beetles (Coleoptera: Carabidae) in Habitat Islands

Aim: The aim of this study was to analyse whether, and how, the inclusion of habitat specialists and edge-preferring species modifies the species-area relationship predictions of the island biogeography theory for an insect group (ground beetles, Coloptera: Carabidae) living in natural fragments. Species-habitat island area relationships applied to terrestrial habitat islands can be distorted by the indiscriminate inclusion of all species occurring in the fragments. Matrices surrounding terrestrial habitat fragments can provide colonists that do not necessarily distinguish the fragment from the matrix and can survive and reproduce there. Edge-preferring species can further distort the expected relationship, as smaller fragments have larger edge:core ratios. Location: Nineteen forest fragments were studied in the Bereg Plain, Hungary, and SW Ukraine. This area contains natural forest patches, mainly of oak and hornbeam, and supports a mountain entomofauna. Methods: Ground beetles (Carabidae) present in the 19 forest patches were categorized into generalists, forest specialists and edge-preferring species. We analysed the relationship between species richness and fragment area using species richness in the different categories. Results: The assemblages contained a high share of generalist species (species that occur also in the surrounding matrix). Forest patch size and the number of generalist species showed a marginally significant negative relationship, indicating that generalist species were more important in smaller patches. Forest specialist species richness was correlated positively with patch area. Edge-preferring species were shown to influence the species-area relationship: the number of edge-preferring species increased with the edge:area ratio. Main conclusions: Both generalist and edge-preferring species can considerably distort the species-area relationship. Island biogeography theory can be applied to habitat islands only if the habitat islands are defined correctly from the viewpoint of the target species.