Evidence for a direct negative effect of habitat fragmentation on forest herb functional diversity

The effects of habitat fragmentation on species richness and composition have been extensively studied. However, little is known about how fragmentation affects functional diversity patterns. Fragmentation can indeed affect functional diversity directly (e.g. by promoting traits associated to long-distance dispersal when fragment isolation increases) or indirectly (e.g. by decreasing species richness, hence trait diversity, when fragment area decreases). Here, we used structural equation modeling to determine whether factors associated to forest fragmentation, namely area, habitat heterogeneity, spatial isolation and age have a direct effect on forest herb functional diversity. Using occurrence data from 243 forest fragments located in northern France and six plant life-history traits, we estimated species richness and calculated functional diversity in each of these 243 forest fragments. We found that species richness was the primary driver of functional diversity in these fragments, with a strong positive and direct relationship between species richness and functional diversity. Interestingly, both fragment isolation and age had a direct negative effect on functional diversity independent of their effects on species richness. Isolation selected life-history traits associated with long-distance dispersal, while age selected for life-history traits typical of forest habitat specialists. Isolated and/or older forest fragments are thus at greater risk of local species and functional extinctions, and hence making these forest fragments particularly vulnerable to future global changes.