The hyporheic zone as an invertebrate refuge: a review of variability in space, time, taxa and behaviour

The hyporheic zone is a potential refuge that can promote persistence of benthic invertebrates during adverse conditions in surface streams. For decades, changes in invertebrate depth distribution have been investigated in relation to flood, low flow and drying events, but evidence for use of the hyporheic refuge remains equivocal. This review examines the evidence for the hyporheic zone's refugial role during adverse hydrological conditions. Refuge potential is influenced by determinants in four categories. First, refuge use varies spatially in relation to physical habitat parameters, including sediment porosity and hydrologic exchange. Second, refuge use is temporally variable and reflects disturbance characteristics including rate of onset. Third, refuge use is taxon-specific, depending on a range of morphological, behavioural and physiological traits. Fourth, the behaviours governing refuge use vary, with both active migrations and passive habitat use playing important roles in community persistence. These four determinants interact to influence refuge use; for example, the physical habitat providing an adequate refuge will vary between taxa. Despite this variability, the hyporheic zone is an important component in the suite of refuges that facilitate community resilience to disturbance events. As such, its ecological integrity should be safeguarded through sensitive management and effective rehabilitation schemes.