Symbiotic relationships between N 2 -fixing prokaryotes and their autotrophic hosts are essential in nitrogen (N)-limited ecosystems, yet the importance of this association in pristine boreal peatlands, which store 25 % of the world's soil (C), has been overlooked. External inputs of N to bogs are predominantly atmospheric, and given that regions of boreal Canada anchor some of the lowest rates found globally (*1 kg N ha -1 year -1), biomass production is thought to be limited primarily by N. Despite historically low N deposition, we show that boreal bogs have accumulated approximately 12-25 times more N than can be explained by atmospheric inputs.Here we demonstrate high rates of biological N 2 -fixation in prokaryotes associated with Sphagnum mosses that can fully account for the missing input of N needed to sustain high rates of C sequestration. Additionally, N amendment experiments in the field did not increase Sphagnum production, indicating that mosses are not limited by N. Lastly, by examining the composition and abundance of N 2 -fixing prokaryotes by quantifying gene expression of 16S rRNA and nitrogenase-encoding nifH, we show that rates of N 2 -fixation are driven by the substantial contribution from methanotrophs, and not from cyanobacteria. We conclude biological N 2 -fixation drives high sequestration of C in pristine peatlands, and may play an important role in moderating fluxes of methane, one of the most important greenhouse gases produced in peatlands. Understanding the mechanistic controls on biological N 2 -fixation is crucial for assessing the fate Responsible Editor: Matthew Wallenstein.Electronic supplementary material The online version of this article (doi:10.1007/s10533-014-0019-6) contains supplementary material, which is available to authorized users. Biogeochemistry (2014) 121:317-328 DOI 10.1007 of peatland carbon stocks under scenarios of climate change and enhanced anthropogenic N deposition.

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We examined concentrations of nitrogen (N) and phosphorus (P) and δ 15 N value in Sphagnum sections Acutifolia and Cuspidata inhabiting hummocks and hollows from 8 bogs along a transect from ~45 to ~55 °N in Ontario/Quebec. The N concentration in Sphagnum declined from South to North, correlating with a decrease in atmospheric N deposition. Although the overall N concentration was larger in hollows than hummocks, the pattern was inconsistent across the sites.There was a proportionally larger decline in P concentration from South to North and an overall larger P concentration in hollows than hummocks, but there were inconsistent differences across the sites. The N:P ratio ranged from 12:1 to 29:1, driven primarily by the variation in P concentration. Ratios of N and P concentration in Sphagnum capitulum:stem averaged 1.2:1, suggesting nutrient resorption from stem to capitulum during growth; the ratio rose with increasing N and P concentration in the capitulum. The δ 15 N value of Sphagnum rose from ~-6‰in the South to ~-1‰ in the North, correlated with the decrease in Sphagnum N concentration and with a rise in water table. We interpret this to indicate a greater dependence on N 2 -fixation for N acquisition in the northern and wetter sites.

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