An experimental test of the effects of food resources and hydraulic refuge on patch colonization by stream macroinvertebrates during spates.

1. The passive or active movement of organisms between habitat patches plays important roles in achieving ecosystem resilience to disturbance and long-term control of population levels. However, causal mechanisms of disturbance-induced movements of mobile biota across heterogeneous habitat patches at a relatively short time-scale are little understood. 2. We experimentally tested the effects of food resource values on macroinvertebrate colonization of hydraulic refugia from spates in a second-order creek. Experimental cages were colonized by macroinvertebrates with combinations of resource types (natural or polyester leaves), and extent of exposure to stream flow (exposed to or sheltered from current); one half of each set was collected before and after a spate. This experiment was repeated over three spates of varying magnitude and seasonal contexts. 3. Pre-spate colonization was consistently greater for the cages with natural leaves relative to artificial leaves regardless of the extent of flow exposure. Two autumn spates with relatively low and stable antecedent flow conditions caused large movements of organic matter and macroinvertebrates across the stream, showing community-level accumulations into hydraulically sheltered patches independent of food treatment. The smallest spate with high and variable antecedent flows during winter resulted in negligible responses, which we interpret to be a result of depletion of easily transportable organic matter and organisms. 4. Two detritivorous taxa, the mayfly Paraleptophlebia spp. and stonefly Despaxia augusta (Banks) were the most responsive to autumn spates, and had disproportionately higher colonization rates of cages when provided with natural leaves during the largest autumn spate. Preferential settlement in food-enriched hydraulic refugia was attributable to taxon-specific mobility related to efficient acquisition of detritus resource, whose availability varies spatially and temporally. 5. Our findings suggest (1) detritivorous macroinvertebrate colonization of hydraulic refugia can be influenced by hydraulic controls as well as food resource value, and (2) pre-spate environmental conditions in terms of resource distribution and availability may pre-condition organisms' susceptibility to spates and also affect refugium usage, at least in food-limited, detritus-based stream systems.