We determined the proportion of butterfly species that occur as metapopulations with grasslands as the only habitat. We counted all butterflies in 19 semi-natural grassland patches in a forest-dominated landscape in southern Sweden over a 5-y period. Seven of the 44 butterfly species observed exhibited a metapopulation structure. The other species either use grassland mainly for adult feeding but not for breeding (8 species), breed both in grassland and in surrounding habitat types (19 species), or are grassland specialists but their colonization-extinction dynamics are probably not significant, since they were present in nearly all (> 80%) patches (10 species). Occupancy was generally higher in larger patches, and tended to increase with patch connectivity. Among grassland specialists and habitat generalists, the connectivity measure tended to explain more of the variation in occupancy if the shortest dispersal paths avoiding routes over water were considered rather than a measure based on the Euclidian distance between patches. This indicates that lakes, even when they are just a few hundred metres wide, can act as barriers to dispersal for butterflies. We conclude that for many butterflies that occur in semi-natural grasslands in forest-dominated landscapes, intervening habitats are important as breeding sites or as dispersal barriers.
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