Modeling the refuge effect of submerged macrophytes in ecological dynamics of shallow lakes: A new model of fish functional response

Submerged macrophytes often provide refuge for zooplankton from fish predation in temperate and subtropical shallow lakes. However, since the relationship between submerged macrophyte abundance and its refuge effect has not been well established, the refuge effect is difficult to be simulated. In this paper, we constructed mathematical models to describe the refuge effect of submerged macrophytes on fish foraging activities and ecological dynamics of shallow lakes based on the previous studies. We clarified the underlying behavioral mechanisms of the observed functional responses through analyses of the fish foraging behavior, extracted the affected variables related to the refuge effect, formulized the relationship between the affected variables and submerged vegetation density, and determined parameter values with a compensative procedure. Calibration and validation results indicated that the new functional response model was successful to simulate the refuge effect on interfering with fish foraging behavior. Moreover, the model was cooperated into a minimal ecological model for shallow lakes. Modeling results showed that the model was able to simulate the refuge effect in ecological dynamics, and made the ecological model produce significantly different results from those with the existing functional response models.