Multiyear resource enrichment creates persistently higher species diversity in a landscape-scale field experiment

Short-term resource enrichment can increase species diversity in communities, but prolonged resource enrichment may result in either a diversity collapse or persistent high species diversity if fluctuation-dependent mechanisms of species coexistence are triggered. We tested the effects of resource enrichment on stream invertebrates by boosting densities of benthic detritus. In a 22-km stream length, we used wooden stakes to enhance retention of detritus at 40-m-long sites; other sites acted as controls. Detritus and invertebrates were sampled prior to treatment and then 1, 2, and 5 yr later. Previously, we reported that detrital densities, species diversity, and densities increased at enrichment sites after 12 months. Here we report that similar increases occurred 2 and 5 yr after manipulation. Prolonged resource enrichment produced persistently higher species diversity without loss of any taxa from the species pool, despite strong shifts in faunal composition in response to environmental variation, including a 1-in-100-yr flood. Detritus densities set upper limits to the densities of common taxa. Positive relations between invertebrate and detritus densities (density-resource relationships) took a variety of forms and showed that detritus was an essential resource for some taxa and a substitutable resource for others. Species varied in the minimum amount of detritus required for presence at a site, and population densities increased strongly from low densities when detritus was increased. These outcomes suggest that fluctuation-dependent mechanisms of coexistence enabled new taxa to coexist at manipulation sites, with relative nonlinear averaging of competition and the storage effect most likely to be in play. Two characteristics of the study stream underpin diversity increases with resource enrichment: overall low background densities of detritus and species that are able to disperse successfully from upstream areas where detritus is more abundant. Thus, the effects of resource enrichment are context dependent.