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As part of a long-term study on the effects of nitrogen (N) loading in a shallow temperate lagoon, we measured rates of N₂fixation associated with seagrass (Zostera marina) epiphytes during the summer from 2005 to 2019, at two sites along a gradient from where high N groundwater enters the system (denoted SH) to a more well-flushed outer harbor (OH). The data presented here are the first such long-term N₂fixation estimates for any seagrass system and one of the very few reported for the phyllosphere in a temperate system. Mean daily N₂fixation was estimated from light and dark measurements using the acetylene reduction assay intercalibrated using both incorporation of¹⁵N₂into biomass and a novel application of the N₂:Ar method. Surprisingly, despite a large inorganic N input from a N-contaminated groundwater plume, epiphytic N₂fixation rates were moderately to very high for a seagrass system (OH site 14-year mean of 0.94 mmol N m⁻² d⁻¹), with the highest rates (2.6 mmol N m⁻² d⁻¹) measured at the more N-loaded eutrophic site (SH) where dissolved inorganic N was higher and soluble reactive phosphorus was lower than in the better-flushed OH. Over 95% of the total N₂fixation measured was in the light, suggesting the importance of cyanobacteria in the epiphyte assemblages. We observed large inter-annual variation both within and across the two study sites (range from 0.1 to 2.6 mmol N fixed m⁻²d⁻¹), which we suggest is in part related to climatic variation. We estimate that input from phyllosphere N₂fixation over the study period contributes on average an additional 20% to the total daily N load per area within the seagrass meadow.

 

Abstract Whether net primary productivity in an aquatic ecosystem is limited by nitrogen (N), limited by phosphorus (P), or co-limited by N & P is determined by the relative supply of N and P to phytoplankton compared to their elemental requirements for primary production, often characterized by the “Redfield” ratio. The supply of these essential nutrients is affected by both external inputs and biogeochemical processes within the ecosystem. In this paper, we examine external sources of nutrients to aquatic systems and how the balance of N to P inputs influences nutrient limitation. For ocean subtropical gyres, a relatively balanced input of N and P relative to the Redfield ratio from deep ocean sources often leads to near co-limitation by N and P. For lakes, the external nutrient inputs come largely from watershed sources, and we demonstrate that on average the N:P ratio for these inputs across the United States is well above that needed by phytoplankton, which may contribute to P limitation in those lake that experience this average nutrient loading. Watershed inputs are also important for estuaries and coastal marine ecosystems, but ocean sources of nutrients are also significant contributors to overall nutrient loads. The ocean-nutrient sources of N and P are very often at or below the Redfield ratio of 16:1 molar, and can be substantially so, particularly in areas where the continental shelf is wide. This large input of coastal ocean nutrients with a low N:P ratio is one factor that may make N limitation more likely in many coastal marine ecosystems than in lakes.