Abstract
Ecological theory suggests that the impact of predation can be strongly modified by the existence of regions of the environment in which prey are less accessible to predators, which underscores the need for empirical studies examining the factors influencing the availability and importance of such prey refuges. Our study tested whether benthic microhabitats with high flows provide suspension-feeding larval black flies (Simulium␣vittatum) with a spatial refuge in which the negative impact of predatory flatworms (Dugesia dorotocephala) is reduced. We conducted a short-term field experiment in Chester Creek (southeastern Pennsylvania, United States) to examine how the number of black fly larvae inhabiting tile substrates responded to manipulated variations in flatworm abundance and current speed. The abundance of flatworms declined with increasing current speed, thereby creating the potential for sites with high flows to provide larvae with a refuge from these predators. Multiple regression analysis revealed that the final abundance of larvae exhibited a significant negative relationship to flatworm abundance and a significant positive relationship to current speed. After adjusting for variations in elapsed time and initial larval abundance, flow and predators explained 38% of the variation in the rate of change in larval abundance. The positive correlation between larval abundance and flow had two components: a positive, direct effect of flow on larvae, which arises because these food-limited consumers prefer to reside within sites with faster flows where they can feed at higher rates; and a negative effect of flow on predators, and of predators on larvae, which combine to yield a positive indirect effect of flow on larvae. This indirect effect demonstrates the existence of flow-mediated refuges (i.e., microhabitats in which the impact of predation is reduced due to high flows), although the effect accounts for a small proportion of total variation in larval abundance. A consideration of biomechanical relationships suggests that microhabitats with high flows are likely to create prey refuges in a wide range of freshwater and marine benthic environments. In particular, predators will often experience greater dislodgement forces than prey because of their larger size and because they project farther above the bed where current speeds are faster. Moreover, the ability to resist a given dislodgement force may be greater for many prey, especially those that are sessile or semi- sessile.
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