Do resources or natural enemies drive bee population dynamics in fragmented habitats?

The relative importance of bottom-up or top-down forces has been mainly studied for herbivores but rarely for pollinators. Habitat fragmentation might change driving forces of population dynamics by reducing the area of resource-providing habitats, disrupting habitat connectivity, and affecting natural enemies more than their host species. We studied spatial and temporal population dynamics of the solitary bee Osmia rufa (Hymenoptera: Megachilidae) in 30 fragmented orchard meadows ranging in size from 0.08 to 5.8 ha in an agricultural landscape in central Germany. From 1998 to 2003, we monitored local bee population size, rate of parasitism, and rate of larval and pupal mortality in reed trap nests as an accessible and standardized nesting resource. Experimentally enhanced nest site availability resulted in a steady increase of mean local population size from 80 to 2740 brood cells between 1998 and 2002. Population size and species richness of natural enemies increased with habitat area, whereas rate of parasitism and mortality only varied among years. Inverse density-dependent parasitism in three study years with highest population size suggests rather destabilizing instead of regulating effects of top-down forces. Accordingly, an analysis of independent time series showed on average a negative impact of population size on population growth rates but provides no support for top-down regulation by natural enemies. We conclude that population dynamics of O. rufa are mainly driven by bottom-up forces, primarily nest site availability.