Stream environments are spatially and temporally heterogeneous. Mainstem rivers are often wide, sunlit, warm and productive, while tributaries are shaded by riparian trees, unproductive, and remain cool in summer. Within mainstem rivers themselves there is substantial spatial heterogeneity in habitat structure and physical conditions, such as water temperature. River environments also change dramatically with season. Organisms that live in the riverine environment respond to and take advantage of such heterogeneous environments by moving between microhabitats or shifting their phenology. I studied the life cycle of a riverine mayfly, Ephemerella maculata (Ephemerellidae), in a northern California river system, its responses to spatial and temporal heterogeneity, and how its movements connect stream and riparian food webs in space and time.

I discovered that E. maculata migrates between the mainstem and tributaries of rivers during its life cycle, thereby linking food webs in these two habitats, and enhancing predator growth in unproductive tributaries. The resource subsidy from productive but warm rivers to cool, unproductive tributaries associated with the mayfly migration increase the growth of stenothermic predators like juvenile salmonids in otherwise food-limited, cool thermal refuges, and increase their resilience to future warming. Furthermore, I examined the resilience of E. maculata to changes in water temperature using field surveys and lab rearing experiments. I discovered that different life stages of E. maculata have different thermal responses, and they shift their phenology depending on the water temperature, allowing each life stage to occur in the most desirable thermal condition. Therefore, as long as the natural seasonal pattern of the water temperature is sustained, E. maculata can resist temperature changes by shifting their phenology. Finally, I have shown that thermal spatial heterogeneity of rivers desynchronizes mayfly emergence timing, prolonging the subsidy period to riparian predators, and changing the predators’ responses to this subsidy.