Extrinsically and intrinsically generated spatial patterns of algal abundance in an experimental stream

Spatial heterogeneity in species abundance arises from both extrinsic (largely abiotic) and intrinsic (largely biotic) processes. The relative importance of these two types of processes can vary across ecological systems and across temporal and spatial scales. Numerous empirical studies have explored spatial patterns resulting from extrinsic and intrinsic processes, however the interaction of these two types of processes can result in complex patterns that are difficult to test. We used a unique model system consisting of periphytic algae grown on agar in an experimental stream to manipulate an extrinsic and an intrinsic process. We manipulated an extrinsic process by varying the spatial arrangement of nutrients creating both heterogeneous and homogeneous environments for the algae. We manipulated an intrinsic process by introducing a snail herbivore to the system. The resulting spatial algal patterns showed that both types of processes were important in producing spatial abundance patterns and that the patterns occurred at two distinct spatial scales in our system. At the scale of the imposed nutrient heterogeneity, algae “tracked” the differences in nutrient supply rates. The snail herbivores both reduced and promoted spatial patterns in algal abundance at different spatial scales reflecting their species-specific foraging behavior. An ability to detect differences in algal abundance allowed the snails to reduce the power of patterns at the scale of the imposed nutrient heterogeneity; however below a spatial scale of approximately 30 mm the snails could no longer detect differences in algal abundance and so foraged randomly. At this spatial scale the spatial heterogeneity in algal abundance increased and the resulting algal patterns were relatively spatially fixed through time. We suggest that this relative constancy may arise in part from a detected weak Allee effect in algal growth rates.