Relative importance of the area and shape of patches to the diversity of multiple taxa.

Although enhancing reserve shape has been suggested as an alternative to enlarging nature reserves, the importance of reserve shape relative to reserve area remains unclear. Here we examined the relative importance of area and shape of forest patches to species richness, species composition, and species abundance (abundance of each species) for 3 taxa (33 birds, 41 butterflies, and 91 forest-floor plants) in a fragmented landscape in central Hokkaido, northern Japan. We grouped the species according to their potential edge responses (interior-, neutral-, and edge-species groups for birds and forest-floor plants, woodland- and open-land-species groups for butterflies) and analyzed them separately. We used a shape index that was independent of area as an index of shape circularization. Hierarchical partitioning and variation partitioning revealed that patch area was generally more important than patch shape for species richness and species composition of birds and butterflies. For forest-floor plants, effects of patch area and shape were small, whereas effects of local forest structure were large. Patch area and circularization generally increased abundances of interior species of birds and forest-floor plants and woodland species of butterflies. Nevertheless, only patch circularization increased abundances of 1 woodland species of butterfly and 2 and 6 interior species of birds and forest-floor plants, respectively. We did not find any significant interaction effects between patch area and shape. Our results suggest that although reserves generally should be large and circular, there is a trade-off between patch area and shape, which should be taken into consideration when managing reserves.