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Recent large-scale seagrass declines have prompted experimental investigations of potential mechanisms. Although many studies have implicated eutrophication or reductions of epi- phyte grazers in these declines, few experiments have simultaneously manipulated both factors to assess their relative effects. This study used meta-analyses of 35 published seagrass studies to com- pare the relative strength of 'top-down' grazer effects and 'bottom-up' nutrient effects on epiphyte biomass and seagrass above-ground growth rate, above-ground biomass, below-ground biomass, and shoot density. A surprising result was that seagrass growth and biomass were limited in situ by sediment nutrients; light limitation has been emphasized in the literature to date. Water column enrichments, which were correlated with increased epiphyte biomass, had strong negative effects on seagrass biomass. Grazers overall had a positive effect on shoot density, but negligible effects on sea- grass biomass and growth rate. However, analyzing epiphyte grazers separately from other grazers revealed positive effects of grazing on seagrass response variables and corresponding negative impacts on epiphyte biomass. The positive effects of epiphyte grazers were comparable in magnitude to the negative impacts of water column nutrient enrichment, suggesting that the 2 factors should not be considered in isolation of each other. Until the determinants of epiphyte grazer populations are empirically examined, it will be difficult to address the contribution that overfishing and cascading trophic effects have had on seagrass decline. Because increases in water column nutrients are docu- mented in many regions, efforts to reduce coastal eutrophication are an appropriate and necessary focus for the management and conservation of seagrass ecosystems.

Relative effects of grazers and nutrients on seagrasses: a meta-analysis approach