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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Oil extraction and development activities in the Athabasca Oil Sands Region of northern Alberta, Canada, release NO, SO, and NH to the atmosphere, ultimately resulting in increasing N and S inputs to surrounding ecosystems through atmospheric deposition. Peatlands are a major feature of the northern Alberta landscape, with bogs covering 6-10% of the land area, and fens covering 21-53%. Bulk deposition of NH-N, NO-N, dissolved inorganic N (DIN), and SO-S, was quantified using ion-exchange resin collectors deployed at 23 locations, over 1-6 years. The results reveal maximum N and S deposition of 9.3 and 12.0 kg ha yr, respectively, near the oil sands industrial center (the midpoint between the Syncrude and Suncor upgrader stacks), decreasing with distance to a background deposition of 0.9 and 1.1 kg ha yr, respectively. To assess potential influences of high N and S deposition on bogs, we quantified N and S concentrations in tissues of two Sphagnum species, two lichen species, and four vascular plant species, as well as surface porewater concentrations of H, NH-N, NO-N, SO-S and dissolved organic N in 19 ombrotrophic bogs, distributed across a 3255 km sampling area surrounding the oil sands industrial center. The two lichen species (Evernia mesomorpha and Cladonia mitis), two vascular plant species (Rhododendron groenlandicum and Picea mariana), and to a lesser extent one moss (Sphagnum fuscum), showed patterns of tissue N and S concentrations that were (1) highest near the oil sands industrial center and (2) positively correlated with bulk deposition of N or S. Concentrations of porewater H and SO-S, but not of NH-N, NO-N, DIN, or dissolved inorganic N, also were higher near the oil sands industrial center than at more distant locations. The oil sands region of northern Alberta is remote, with few roads, posing challenges to the monitoring of oil sands-related N and S deposition. Quantification of N and S concentrations in bog plant/lichen tissues and porewaters may serve as a monitoring tool to assess both the local intensity and the spatial extent of bulk N and S deposition, and as harbingers of potential shifts in ecosystem structure and function.

Increasing gaseous emissions of nitrogen (N) and sulfur (S) associated with oil sands development in northern Alberta (Canada) has led to changing regional wet and dry N and S deposition regimes. We assessed the potential for using bog plant/lichen tissue chemistry (N and S concentrations, C:N and C:S ratios, in 10 plant/lichen species) to monitor changing atmospheric N and S deposition through sampling at five bog sites, 3–6 times per growing season from 2009 to 2016. During this 8-year period, oil sands N emissions steadily increased, while S emissions steadily decreased. We examined the following: (1) whether each species showed changes in tissue chemistry with increasing distance from the Syncrude and Suncor upgrader stacks (the two largest point sources of N and S emissions); (2) whether tissue chemistry changed over the 8 year period in ways that were consistent with increasing N and decreasing S emissions from oil sands facilities; and (3) whether tissue chemistry was correlated with growing season wet deposition of NH4+-N, NO3−-N, or SO42−-S. Based on these criteria, the best biomonitors of a changing N deposition regime were Evernia mesomorpha, Sphagnum fuscum, and Vaccinium oxycoccos. The best biomonitors of a changing S deposition regime were Evernia mesomorpha, Cladonia mitis, Sphagnum fuscum, Sphagnum capillifolium, Vaccinium oxycoccos, and Picea mariana. Changing N and S deposition regimes in the oil sands region appear to be influencing N and S cycling in what once were pristine ombrotrophic bogs, to the extent that these bogs may effectively monitor future spatial and temporal patterns of deposition.

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