The concept of habitat specificity revisited

Habitat specificity indices reflect richness (alpha) and/or distinctiveness (beta) components of diversity. The latter may be defined by alpha and gamma (landscape) diversity in two alternative ways: multiplicatively ( beta = gamma/alpha ) and additively ( beta = gamma - alpha). We demonstrate that the original habitat specificity concept of Wagner and Edwards (Landscape Ecol 16:121-131, 2001) consists of three independent components: core habitat specificity (uniqueness of the species composition), patch area and patch species richness. We describe habitat specificity as a family of indices that may include either area or richness components, or none or both, and open for use of different types of mean in calculation of core habitat specificity. Core habitat specificity is a beta diversity measure: the effective number of completely distinct communities in the landscape. Habitat specificity weighted by species number is a gamma diversity measure: the effective number of species that a patch contributes to landscape richness. We compared 12 habitat specificity indices by theoretical reasoning and by use of field data (vascular plant species in SE Norwegian agricultural landscapes). Habitat specificity indices are strongly influenced by weights for patch area and patch species richness, and the relative contribution of rare vs. common species (type of mean). The relevance of properties emphasized by each habitat specificity index for evaluation of patches in a biodiversity context is discussed. Core habitat specificity is emphasized as an ecologically interpretable measure that specifically addresses patch uniqueness while habitat specificity weighted by species number combines species richness and species composition in ways relevant for conservation biological assessment.