Strong influence of long-distance edge effect on herb-layer vegetation in forest fragments in an agricultural landscape

In agricultural landscapes in central Europe, species richness of the herbaceous plant community may be compromised by processes associated with forest fragmentation, habitat loss, and management practices. We examined variability in species richness and composition of the herbaceous layer in 229 plots located in 23 forest fragments (0.1 to 255 ha), in a representative upland agricultural landscape in central Bohemia, in relation to the most important site environmental factors, edge effects, and site history. The influence of environmental factors on the composition of vegetation in the herb layer was evaluated using generalized additive models, which enabled us to analyze highly non-linear and non-monotonic relationships. Total species richness and number of red-listed and ancient forest species were significantly influenced by type of forest vegetation, light quality, soil pH, slope aspect, and distance from the forest edge. Implications of the significant explanatory variables corresponded well to previous findings, with the exception of distance from the forest edge, for which we found a positive relationship with species richness for distances up to 200 m toward the forest interior. Plant species with low colonization ability occupied plots with increasing frequency from edge to forest interior, while fast-colonizing species showed the opposite trend. Apart from the edge effect, forest continuity should be considered for its important contribution to the richness of ancient forest and red-listed species, whereas the effect of forest fragment size appeared to be generally weak. These results do not negate the importance of large forest fragments for the maintenance of herb layer species richness, but specifically emphasize the essential contribution of the core habitats of these forests. In summary, we showed that the negative effects of habitat fragmentation on the richness of ancient forest and red-listed species and on herb layer species in total can be largely attributed to either the edge effect itself or to aggregate effects of forest edge and forest continuity.