Combined effects of area, connectivity, history and structural heterogeneity of woodlands on the species richness of hoverflies (Diptera: Syrphidae)

Hoverflies are often used as bio-indicators for ecosystem conservation, but only few studies have actually investigated the key factors explaining their richness in woodlands. In a fragmented landscape in southwest France, we investigated the joint effects of woodland area, structural heterogeneity, connectivity and history on the species richness of forest-specialist hoverflies, and whether there was a time lag in the response of hoverflies to habitat changes, and tested the effect of spatiotemporal changes. Current species richness was sampled in 48 woodlands using 99 Malaise traps. Structural variables were derived from a rapid habitat assessment protocol. Old maps and aerial photographs were used to extract past and present spatial patterns of the woodlands since 1850. Relationships between species richness and explanatory variables were explored using generalized linear models. We show that current habitat area, connectivity, historical continuity and the average density of tree-microhabitats explained 35 % of variation in species richness. Species richness was affected differently by changes in patch area between 1979 and 2010, depending on woodland connectivity. In isolated woodlands, extinction debt and colonization credit were revealed, showing that even several decades are not sufficient for hoverflies to adapt to landscape-scale habitat conditions. These findings emphasise the importance of maintaining connectedness between woodlands, which facilitates the dispersion in a changing landscape. Our results also highlight the benefits of using a change-oriented approach to explain the current distribution patterns of species, especially when several spatial processes act jointly.