The influence of resource subsidies on cave invertebrates: results from an ecosystem-level manipulation experiment

Spatial resource subsidies can greatly affect the composition and dynamics of recipient communities. Caves are especially tractable for studying spatial subsidies because primary productivity is absent. Here, we performed an ecosystem-level manipulation experiment to test the direct influence of detrital subsidies on community structure in terrestrial cave ecosystems. After performing baseline censuses of invertebrates, we removed all organic material from 12 caves and constructed exclusion boxes to prevent natural resource inputs. Next, we stocked each cave with standardized quantities of two major natural subsidies to caves: leaves (leaf packs) and carcasses (commercially supplied rodents), and measured the invertebrate colonization and utilization of these resources for 23 months. Over the course of the experiment, 102 morphospecies were observed. Diplopods and collembolans were most abundant on leaf packs, and dipteran larvae and collembolans were most abundant on the rats. On average, caves receiving either treatment did not differ in species richness, but abundance was significantly higher in rat caves over both the duration of the experiment and the temporal "life" of the individual resources, which were restocked upon exhaustion. Post-manipulation invertebrate communities differed predictably depending on the type of subsidy introduced. Over the course of the experiment, caves that received the same subsidy clustered together based on community composition. In addition, the invertebrate community utilizing the resource changed over the duration of the two-year experiment, and evidence of succession (i.e., directional change) was observed. Results from this study demonstrate how allochthonous resources can drive the community dynamics of terrestrial invertebrates in cave ecosystems and highlight the need for consideration of the surface environment when managing and protecting these unique habitats.