Colonization of central European abandoned fields by dry grassland species depends on the species richness of the source habitats: a new approach for measuring habitat isolation

Abandoned fields are perceived as potential habitats for species of threatened semi-natural dry grasslands. However, information is lacking regarding how the spontaneous colonization of abandoned fields depends on the broader spatial context. We recorded the occurrence of 87 target species in 46 abandoned fields and 339 dry grasslands. We tested the effect of the isolation of abandoned fields from source grasslands on the number of dry grassland species occurring in abandoned fields either with or without habitat characteristics being used as covariates. The isolation of the fields was calculated using the distance and area (I (A) ) or distance and species richness (I (S) ) of source habitats. I (S) always explained the number of grassland species in the abandoned fields better than I (A) . The effect of isolation became smaller or even non-significant with the inclusion of covariates; it also changed with the method used for measuring distance (edge-to-edge or center-to-center), and it was lower when other abandoned fields were considered as additional source habitats. The different performance of the two isolation measures can be explained by the weak species-area relationship in the grasslands, indicating differences in their habitat quality. Species richness is a better proxy of habitat importance in terms of propagule source than habitat area, and the new isolation measure is therefore suitable for studying the effects of landscape structure on species richness in landscapes presenting a weak species-area relationship, such as areas exhibiting pronounced effects of land-use history. Inclusion of habitat characteristics as covariates may considerably alter conclusions regarding the effect of isolation, which might actually be overestimated when assessed separately.