Fluid-mediated dispersal in streams: models of settlement from the drift.
I propose proximal mechanisms that help explain, unify, and expand the predictions of widely accepted empirical models of settlement in streams. I separated the process that leads to settlement of a drifting particle into three stages: (1) initial contact with a substrate, (2) attachment, and (3) settlement sensu stricto. I used physical principles (height above the bed, sinking rate, current speed profile) to predict time until contact (stage 1). I compared these predictions with empirical measurements of settlement of individual black fly larvae (Simulium vittatum) in a laboratory flume. I developed models from empirical data for stages 2 and 3. Each of these models is individual-based and predicts the fate of a single individual. To obtain a population level prediction, models for the three stages were combined and used to simulate the settlement of a group of black fly larvae. The predictions of this simulation were qualitatively similar to population level data from the literature particularly after the incorporation of channel-wide spatial heterogeneity in current speed. The effect of flow heterogeneity on the model agrees with previous work on the lateral transport of stream invertebrates during drift events showing that many organisms settle preferentially in slower areas. By using proximal principles, the approach used in this study brings into focus basic parameters and processes that influence settlement at the scale of the organisms. It also provides a null hypothesis against which to study the effect of local flow heterogeneity on the settlement of stream invertebrates and the capacity of organisms to actively influence settlement. Water currents in streams and rivers commonly transport large numbers of organisms. Consequently, hydrodynamic factors that favor or hamper the settlement of these organisms can potentially influence distributions and abundance. Moreover, if settlement probabilities vary with flow characteristics, this can in turn influence foraging strategies that rely on fluid-mediated dispersal.