Grazers and Nutrients Simultaneously Limit Lotic Primary Productivity

Nitrate and phosphate were added to two of four flow through channels in a second-order stream in eastern Tennessee to raise nutrient concentrations to 3–4 times background, while two of four sets of colonized ceramic tiles in each channel were raised above the substratum to exclude grazing snails (Elimia clavaeformis). Snail grazing maintained a thin layer of periphyton dominated by Stigeoclonium basal cells, regardless of nutrient regime. Although nutrient effects on periphyton ash-free dry mass were statistically insignificant, nutrient additions significantly increased chlorophyll a, especially where snails were excluded. Snail densities were 89% higher in nutrient-enriched channels. Photosynthesis–irradiance data suggested that nutrient enrichment increased self-shading in the periphyton. Areal-specific productivity was simultaneously limited by grazing and low nutrient concentrations: snail exclusion and nutrient enrichment both increased productivity > 2 times. The negative effect of snails on areal-specific productivity was due to (1) reduction in biomass by cropping and (2) depression of chlorophyll-specific productivity. The means by which Elimia depresses chlorophyll-specific productivity is unclear, but the depression is clearly disadvantageous to food-limited grazers. Because Elimia was the dominant invertebrate, our results indicate that low nutrient concentrations limit secondary as well as primary production in autumn.