Indirect effects of predators control herbivore richness and abundance in a benthic eelgrass (Zostera marina) mesograzer community

1. Herbivore communities can be sensitive to changes in predator pressure (top-down effects) and resource availability (bottom-up effects) in a wide range of systems. However, it remains unclear whether such top-down and bottom-up effects reflect direct impacts of predators and/or resources on herbivores, or are indirect, reflecting altered interactions among herbivore species. 2. We quantified direct and indirect effects of bottom-up and top-down processes on an eelgrass (Zostera marina) herbivore assemblage. In a field experiment, we factorially manipulated water column nutrients (with Osmocote™ slow-release fertilizer) and predation pressure (with predator exclusion cages) and measured the effects on herbivore abundance, richness and beta diversity. We examined likely mechanisms of community responses by statistically exploring the response of individual herbivore species to trophic manipulations. 3. Predators increased herbivore richness and total abundance, in both cases through in...