Ecological effects of perturbation by drought in flowing waters

SUMMARY Knowledge of the ecology of droughts in flowing waters is scattered and fragmentary, with much of the available information being gathered opportunistically. Studies on intermittent and arid‐zone streams have provided most of the information. Drought in streams may be viewed as a disturbance in which water inflow, river flow and water availability fall to extremely low levels for extended periods of time. As an ecological perturbation, there is the disturbance of drought and the responses of the biota to the drought. Droughts can either be periodic, seasonal or supra‐seasonal events. The types of disturbance for seasonal droughts are presses and for supra‐seasonal droughts, ramps. In droughts, hydrological connectivity is disrupted. Such disruption range from flow reduction to complete loss of surface water and connectivity. The longitudinal patterns along streams as to where flow ceases and drying up occurs differs between streams. Three patterns are outlined: ‘downstream drying’, ‘headwater drying’ and ‘mid‐reach drying’. There are both direct and indirect effects of drought on stream ecosystems. Marked direct effects include loss of water, loss of habitat for aquatic organisms and loss of stream connectivity. Indirect effects include the deterioration of water quality, alteration of food resources, and changes in the strength and structure of interspecific interactions. Droughts have marked effects on the densities and size‐ or age‐structure of populations, on community composition and diversity, and on ecosystem processes. Organisms can resist the effects of drought by the use of refugia. Survival in refugia may strongly influence the capacity of the biota to recover from droughts once they break. Recovery by biota varies markedly between seasonal and supra‐seasonal droughts. Faunal recovery from seasonal droughts follows predictable sequences, whilst recovery from supra‐seasonal droughts varies from one case to another and may be marked by dense populations of transient species and the depletion of biota that normally occur in the streams. The restoration of streams must include the provision of drought refugia and the inclusion of drought in the long‐term flow regime.