Human activities have increased the availability of reactive nitrogen in many ecosystems, leading to negative impacts on human health, biodiversity, and water quality. Freshwater ecosystems, including lakes, streams, and wetlands, are a large global sink for reactive nitrogen, but factors that determine the efficacy of freshwater nitrogen removal rates are poorly known. Using a global lake data set, we show that the availability of phosphorus, a limiting nutrient, affects both annual nitrogen removal rate and efficiency. This result indicates that increased phosphorus inputs from human activities have stimulated nitrogen removal processes in many lakes. Recent management-driven reductions in phosphorus availability promote water column accumulation and export of nitrogen from large lakes, an unintended consequence of single-element management that argues for greater control of nitrogen as well as phosphorus sources.

A central tenet of ecological stoichiometry is that consumer elemental composition is relatively independent of food resource nutrient content. Although the P content of some invertebrate consumer taxa can increase as a consequence of P-enriched food resources, little is known about how ecosystem nutrient loading can affect the elemental composition of entire consumer assemblages. Here we examine the potential for P enrichment across invertebrate consumer assemblages in response to chronic high P loading. We measured elemental ratios in invertebrate consumers and basal food resources in a series of streams in lowland Costa Rica that range widely in P levels (2-135 microg l(-1) soluble reactive P). Streams with high P levels receive natural long-term (over millennia) inputs of solute-rich groundwater while low-P streams do not receive these solute-rich groundwater inputs. P content of leaf litter and epilithon increased fourfold across the natural P gradient, exceeding basal resource P content values reported in the literature from other nutrient-rich streams. Invertebrate consumers from the high-P study stream were elevated twofold in P content across multiple taxonomic and functional feeding groups, including predators. Our results strongly support the hypothesis that elevated P content in consumers feeding on P-enriched food resources is a consequence of deviation from strict homeostasis. In contrast to prior studies, we found that between-stream variation in P content of a given taxon greatly exceeded within-stream variation among different taxa, suggesting that environment may be as important as phylogeny in controlling consumer stoichiometry. Relaxing the assumption of strict homeostasis presents challenges and opportunities for advancing our understanding of how nutrient limitation affects consumer growth. Moreover, our findings may provide a window into the future of how chronic anthropogenic nutrient loading can alter stoichiometric relationships in food webs.

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.

Recent discoveries have altered prevailing paradigms concerning the conditions under which nitrification takes place and the organisms responsible for nitrification in aquatic ecosystems. In Lake Superior, nitrate (NO is generated by nitrification within the lake, important questions remain concerning the magnitude and controls of nitrification, and which microbial groups are primarily responsible for this process. We measured water-column nitrification rates in the western basin of Lake Superior during five research cruises from November 2009 to March 2011. Using in situ bottle incubations at 10 depths, we quantified nitrification rates using both the oxidation of 15 N-labeled ammonium (NH z 4 ) and the uptake of 14 C associated with nitrification. Average rates of NH z 4 oxidation ranged from 18-34 nmol N L 21 d 21 across the five cruises, similar to values reported for the coastal ocean, and two orders of magnitude lower than values reported from other lakes. Low nitrification rates observed in the epilimnion corresponded to the absence of ammonium-oxidizing archaea and nitrite-oxidizing bacteria. The measured rates of nitrification are . 50-fold greater than the long-term NO { 3 rise in the lake, indicating that N is actively cycling and that long-term change in this ecosystem is mediated by internal dynamics.

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