Disturbance and patch-specific responses: the interactive effects of woody debris and floods on lotic invertebrates.

Disturbance may play an important role in generating patterns of abundance and distribution of biotic assemblages, particularly if its impact differs among habitat patches. Despite much speculation concerning the probable importance of spatial variation in the response of stream fauna to flooding, empirical work on patch-specific responses to spates is largely lacking. Floods typically reduce the abundance of lotic invertebrates dramatically in open-channel areas. We conducted a set of experiments to determine if faunal abundances are less affected in patches more sheltered due to the presence of woody debris dams. Specifically, we tested two hypotheses using chironomids and copepods living in a warmwater, 4th order stream: (1) the effect of flooding on the fauna varies between patches associated with debris dams versus the open channel, and (2) the absence of woody debris in a stream impedes faunal recovery throughout the channel following floods. We tested the first hypothesis by quantifying faunal abundances prior to, during, and following two floods in four patch types: mid-channel sandy patches distant from dams, coarse sediments associated with dams, fine sediments associated with dams, and leafy debris in dams. The second hypothesis was tested by removing all of the woody debris from two stretches of the stream and comparing the impact of a flood on fauna in debris-removed versus control stretches. Across all of the eight study dams, there were patchspecific faunal responses to two floods. Removal of woody debris from the stream did not prevent faunal recovery throughout the channel; however, the presence of woody debris dams did confer greater resistance of fauna to floods (as measured by no decrease in abundance during flooding) in two patch types. Abundances of chironomids and, to a lesser extent, copepods in the leafy debris of dams and in fine sediment patches associated with some dams either did not change or increased during floods, despite the fact that abundances in the dominant patch type of the stream (the sandy mid-channel) were reduced by 75-95%. All instances of faunal increase were limited to fine sediment patches associated with dams, thus entire dams cannot be labeled as flow refugia per se. Statistically, we distinguished fine patches which accumulated animals during floods from the other fine patches based on two physical attributes. Patches accumulating animals were all characterized by low water flux and nearbed flow, which likely contributed to the retention and/or passive deposition of animals. Whole dam attributes (e.g. dam size or complexity) were not useful in predicting which of the dams would accumulate animals in their fine sediments during flooding. Although structural complexity - here in the form of wood and leafy debris - is clearly important in generating biotic pattern in many ecosystems, our work underscores the need to understand what processes are responsible for the link between physical structure and biotic pattern.