A landscape-scale field experiment reveals the importance of dispersal in a resource-limited metacommunity

Dispersal may play a strong role in driving species diversity across landscapes. Theoretically, dispersal permits species to remain extant within a metacommunity, even if they are periodically excluded from some local communities. Field tests of dispersal effects are difficult, and most non-experimental data suggest that environmental conditions play the predominant role in setting species diversity. However, most such studies cannot differentiate between patterns caused primarily by dispersal constraints vs. abiotic factors vs. biotic constraints (e.g., priority effects). In 22 km of a sand-bed stream in southeastern Australia, strong longitudinal gradients in the abiotic environment and detritus densities (resources) mean that downstream locations have abiotic conditions that may be physiologically stressful, low resource densities and low species diversities. We experimentally increased the retention of detritus over 40 m stream lengths along the gradient, with other sites acting as controls. If dispersal is constrained, or abiotic or biotic factors primarily control community structure, then we predicted that increasing resources would result in no change in species composition. Alternatively, if dispersal is common, we predicted that species diversity would increase at treatment sites through colonization by species able to tolerate abiotic conditions downstream and able to invade established communities. Invertebrates were sampled prior to manipulation and then four times (1, 4, 9 and 12 months) following manipulation. Detrital standing stocks increased by an order of magnitude at treatment sites. Over 1 yr, invertebrate densities and species richness also increased in treatment sites. Effect sizes were strong in middle and downstream areas, which were colonized by upstream species. Thus, faunal composition of the downstream treatment sites became more similar to upstream locales, and β-diversity across treatment sites declined as α-diversity increased. Out of 54 common taxa, roughly half responded to the experiment; responders and non-responders had similar proportions of upstream specialists and of different functional feeding groups. Contrary to many non-experimental studies, our results demonstrate that extensive dispersal can be very important and, for many species, potentially more important than the abiotic environment or biotic constraints in affecting community structure when adequate resources are available.