Effects of herbivorous fish on shallow lake ecosystems increase at moderate nutrient conditions.

Submerged macrophytes are vital in shallow lakes, as they provide critical ecosystem functions and services and can stabilize the clear-water conditions by various mechanisms. Nutrient enrichment reduces the resilience of macrophyte dominance in shallow lakes, thereby making them susceptible to shifts towards phytoplankton dominance following perturbations. Here, we conducted a mesocosm experiment to examine the individual and combined effects of nutrient enrichment and the addition of grass carp (Ctenopharyngodon idella) on the abundance of submerged macrophytes, epiphyton, and phytoplankton. We hypothesized that moderate nutrient enrichment facilitates macrophyte abundance, but also phytoplankton abundance after macrophyte removal by herbivorous fish. Our data showed that herbivory by grass carp could trigger a shift from macrophytes to algal dominance in mesocosms with moderate nutrient concentrations, but not in those with low nutrient concentrations. Moderate nutrient enrichment alone promoted submerged macrophyte growth, whereas the introduction of grass carp induced a collapse of submerged macrophytes regardless of nutrient conditions. Moreover, the introduction of grass carp showed more negative effects on light conditions of the water column in mesocosms with moderate nutrient concentrations compared to those with low nutrient concentrations. A recovery of submerged macrophytes might thus be limited by low light availability in lakes with moderate nutrient conditions suffering grass carp perturbation. Our results suggest that submerged macrophyte-dominated shallow lakes with moderate nutrient conditions are vulnerable to perturbation by herbivorous fish such as grass carp. In turn, managing the abundance of herbivores in these lakes can support the dominance of macrophytes and associated clear water conditions.