Snails have stronger indirect positive effects on submerged macrophyte growth attributes than zooplankton

Phytoplankton and epiphyton often compete with submerged macrophytes. Grazing by zooplankton and/or epiphyton grazers should promote an indirect positive effect on submerged macrophyte growth rate. Hence, we mimicked shallow lakes conditions in mesocosms using a factorial design to evaluate the indirect effects of no grazers, zooplankton, snails or both grazers on macrophyte growth attributes. After 16 weeks, both snails and zooplankton had positive effects on macrophyte stem length and biomass. However, only snails had positive effects on macrophyte number of sprouts and root biomass. In addition, the positive effect size of snails on the submerged macrophytes was twice as large as the effect size of the zooplankton. Our study suggests that benthic food chains might be more capable of increasing resilience and affecting the stability of the clear-water state in shallow lakes than pelagic food chains. However, long-term experiments with varying relative proportions of herbivores and different macrophyte species, as well as in situ experiments, will be necessary to test the generality of our findings. Understanding the relative effects of benthic versus pelagic grazers on submerged macrophytes may increase the success of shallow lake restoration and should be taken into account when designing management and restoration efforts for shallow lake systems