Legacy effects of nitrogen and phosphorus additions on vegetation and carbon stocks of upland heaths.

Soil carbon (C) pools and plant community composition are regulated by nitrogen (N) and phosphorus (P) availability. Atmospheric N deposition impacts ecosystem C storage, but the direction of response varies between systems. Phosphorus limitation may constrain C storage response to N, hence P application to increase plant productivity and thus C sequestration has been suggested. We revisited a 23-yr-old field experiment where N and P had been applied to upland heath, a widespread habitat supporting large soil C stocks. At 10 yr after the last nutrient application we quantified long-term changes in vegetation composition and in soil and vegetation C and P stocks. Nitrogen addition, particularly when combined with P, strongly influenced vegetation composition, favouring grasses over Calluna vulgaris, and led to a reduction in vegetation C stocks. However, soil C stocks did not respond to nutrient treatments. We found 40% of the added P had accumulated in the soil. This study showed persistent effects of N and N + P on vegetation composition, whereas effects of P alone were small and showed recovery. We found no indication that P application could mitigate the effects of N on vegetation or increase C sequestration in this system.

Rhizosphere allocation by canopy-forming species dominates soil CO2 efflux in a subarctic landscape.

In arctic ecosystems, climate change has increased plant productivity. As arctic carbon (C) stocks predominantly are located belowground, the effects of greater plant productivity on soil C storage will significantly determine the net sink/source potential of these ecosystems, but vegetation controls on soil CO2 efflux remain poorly resolved. In order to identify the role of canopy-forming species in belowground C dynamics, we conducted a girdling experiment with plots distributed across 1 km2 of treeline birch (Betula pubescens) forest and willow (Salix lapponum) patches in northern Sweden and quantified the contribution of canopy vegetation to soil CO2 fluxes and belowground productivity. Girdling birches reduced total soil CO2 efflux in the peak growing season by 53%, which is double the expected amount, given that trees contribute only half of the total leaf area in the forest. Root and mycorrhizal mycelial production also decreased substantially. At peak season, willow shrubs contributed 38% to soil CO2 efflux in their patches. Our findings indicate that C, recently fixed by trees and tall shrubs, makes a substantial contribution to soil respiration. It is critically important that these processes are taken into consideration in the context of a greening arctic because productivity and ecosystem C sequestration are not synonymous.

Correction to: Long-Term Recovery of Microbial Communities in the Boreal Bryosphere Following Fire Disturbance.

The original version of this article contained an error in the Molecular Analysis subsection of the Methods.

Exploring drivers of litter decomposition in a greening Arctic: results from a transplant experiment across a treeline.

Decomposition of plant litter is a key control over carbon (C) storage in the soil. The biochemistry of the litter being produced, the environment in which the decomposition is taking place, and the community composition and metabolism of the decomposer organisms exert a combined influence over decomposition rates. As deciduous shrubs and trees are expanding into tundra ecosystems as a result of regional climate warming, this change in vegetation represents a change in litter input to tundra soils and a change in the environment in which litter decomposes. To test the importance of litter biochemistry and environment in determining litter mass loss, we reciprocally transplanted litter between heath (Empetrum nigrum), shrub (Betula nana), and forest (Betula pubescens) at a sub-Arctic treeline in Sweden. As expansion of shrubs and trees promotes deeper snow, we also used a snow fence experiment in a tundra heath environment to understand the importance of snow depth, relative to other factors, in the decomposition of litter. Our results show that B. pubescens and B. nana leaf litter decomposed at faster rates than E. nigrum litter across all environments, while all litter species decomposed at faster rates in the forest and shrub environments than in the tundra heath. The effect of increased snow on decomposition was minimal, leading us to conclude that microbial activity over summer in the productive forest and shrub vegetation is driving increased mass loss compared to the heath. Using B. pubescens and E. nigrum litter, we demonstrate that degradation of carbohydrate-C is a significant driver of mass loss in the forest. This pathway was less prominent in the heath, which is consistent with observations that tundra soils typically have high concentrations of "labile" C. This experiment suggests that further expansion of shrubs and trees may stimulate the loss of undecomposed carbohydrate C in the tundra.

Using stable isotopes to estimate travel times in a data-sparse Arctic catchment: Challenges and possible solutions.

Use of isotopes to quantify the temporal dynamics of the transformation of precipitation into run-off has revealed fundamental new insights into catchment flow paths and mixing processes that influence biogeochemical transport. However, catchments underlain by permafrost have received little attention in isotope-based studies, despite their global importance in terms of rapid environmental change. These high-latitude regions offer limited access for data collection during critical periods (e.g., early phases of snowmelt). Additionally, spatio-temporal variable freeze-thaw cycles, together with the development of an active layer, have a time variant influence on catchment hydrology. All of these characteristics make the application of traditional transit time estimation approaches challenging. We describe an isotope-based study undertaken to provide a preliminary assessment of travel times at Siksik Creek in the western Canadian Arctic. We adopted a model-data fusion approach to estimate the volumes and isotopic characteristics of snowpack and meltwater. Using samples collected in the spring/summer, we characterize the isotopic composition of summer rainfall, melt from snow, soil water, and stream water. In addition, soil moisture dynamics and the temporal evolution of the active layer profile were monitored. First approximations of transit times were estimated for soil and streamwater compositions using lumped convolution integral models and temporally variable inputs including snowmelt, ice thaw, and summer rainfall. Comparing transit time estimates using a variety of inputs revealed that transit time was best estimated using all available inflows (i.e., snowmelt, soil ice thaw, and rainfall). Early spring transit times were short, dominated by snowmelt and soil ice thaw and limited catchment storage when soils are predominantly frozen. However, significant and increasing mixing with water in the active layer during the summer resulted in more damped steam water variation and longer mean travel times (~1.5 years). The study has also highlighted key data needs to better constrain travel time estimates in permafrost catchments.

Draft Genome Sequence of Methylocella silvestris TVC, a Facultative Methanotroph Isolated from Permafrost.

Permafrost environments play a crucial role in global carbon and methane cycling. We report here the draft genome sequence of Methylocella silvestris TVC, a new facultative methanotroph strain, isolated from the Siksik Creek catchment in the continuous permafrost zone of Inuvik (Northwest Territories, Canada).

Long-Term Recovery of Microbial Communities in the Boreal Bryosphere Following Fire Disturbance.

Our study used a ∼360-year fire chronosequence in northern Sweden to investigate post-fire microbial community dynamics in the boreal bryosphere (the living and dead parts of the feather moss layer on the forest floor, along with the associated biota). We anticipated systematic changes in microbial community structure and growth strategy with increasing time since fire (TSF) and used amplicon pyrosequencing to establish microbial community structure. We also recorded edaphic factors (relating to pH, C and N accumulation) and the physical characteristics of the feather moss layer. The molecular analyses revealed an unexpectedly diverse microbial community. The structure of the community could be largely explained by just two factors, TSF and pH, although the importance of TSF diminished as the forest recovered from disturbance. The microbial communities on the youngest site (TSF = 14 years) were clearly different from older locations (>100 years), suggesting relatively rapid post-fire recovery. A shift towards Proteobacterial taxa on older sites, coupled with a decline in the relative abundance of Acidobacteria, suggested an increase in resource availability with TSF. Saprotrophs dominated the fungal community. Mycorrhizal fungi appeared to decline in abundance with TSF, possibly due to changing N status. Our study provided evidence for the decadal-scale legacy of burning, with implications for boreal forests that are expected to experience more frequent burns over the course of the next century.

Biogeochemistry of "pristine" freshwater stream and lake systems in the western Canadian Arctic.

Climate change poses a substantial threat to the stability of the Arctic terrestrial carbon (C) pool as warmer air temperatures thaw permafrost and deepen the seasonally-thawed active layer of soils and sediments. Enhanced water flow through this layer may accelerate the transport of C and major cations and anions to streams and lakes. These act as important conduits and reactors for dissolved C within the terrestrial C cycle. It is important for studies to consider these processes in small headwater catchments, which have been identified as hotspots of rapid mineralisation of C sourced from ancient permafrost thaw. In order to better understand the role of inland waters in terrestrial C cycling we characterised the biogeochemistry of the freshwater systems in a c. 14 km2 study area in the western Canadian Arctic. Sampling took place during the snow-free seasons of 2013 and 2014 for major inorganic solutes, dissolved organic and inorganic C (DOC and DIC, respectively), carbon dioxide (CO2) and methane (CH4) concentrations from three water type groups: lakes, polygonal pools and streams. These groups displayed differing biogeochemical signatures, indicative of contrasting biogeochemical controls. However, none of the groups showed strong signals of enhanced permafrost thaw during the study seasons. The mean annual air temperature in the region has increased by more than 2.5 °C since 1970, and continued warming will likely affect the aquatic biogeochemistry. This study provides important baseline data for comparison with future studies in a warming Arctic.

Slow recovery of High Arctic heath communities from nitrogen enrichment.

Arctic ecosystems are strongly nutrient limited and exhibit dramatic responses to nitrogen (N) enrichment, the reversibility of which is unknown. This study uniquely assesses the potential for tundra heath to recover from N deposition and the influence of phosphorus (P) availability on recovery. We revisited an experiment in Svalbard, established in 1991, in which N was applied at rates representing atmospheric N deposition in Europe (10 and 50 kg N ha(-1)  yr(-1) ; 'low' and 'high', respectively) for 3-8 yr. We investigated whether significant effects on vegetation composition and ecosystem nutrient status persisted up to 18 yr post-treatment. Although the tundra heath is no longer N saturated, N treatment effects persist and are strongly P-dependent. Vegetation was more resilient to N where no P was added, although shrub cover is still reduced in low-N plots. Where P was also added (5 kg P ha(-1)  yr(-1) ), there are still effects of low N on community composition and nutrient dynamics. High N, with and without P, has many lasting impacts. Importantly, N + P has caused dramatically increased moss abundance, which influences nutrient dynamics. Our key finding is that Arctic ecosystems are slow to recover from even small N inputs, particularly where P is not limiting.

The role of mosses in carbon uptake and partitioning in arctic vegetation.

The Arctic is already experiencing changes in plant community composition, so understanding the contribution of different vegetation components to carbon (C) cycling is essential in order to accurately quantify ecosystem C balance. Mosses contribute substantially to biomass, but their impact on carbon use efficiency (CUE) - the proportion of gross primary productivity (GPP) incorporated into growth - and aboveground versus belowground C partitioning is poorly known. We used (13) C pulse-labelling to trace assimilated C in mosses (Sphagnum sect. Acutifolia and Pleurozium schreberi) and in dwarf shrub-P. schreberi vegetation in sub-Arctic Finland. Based on (13) C pools and fluxes, we quantified the contribution of mosses to GPP, CUE and partitioning. Mosses incorporated 20 ± 9% of total ecosystem GPP into biomass. CUE of Sphagnum was 68-71%, that of P. schreberi was 62-81% and that of dwarf shrub-P. schreberi vegetation was 58-74%. Incorporation of C belowground was 10 ± 2% of GPP, while vascular plants alone incorporated 15 ± 4% of their fixed C belowground. We have demonstrated that mosses strongly influence C uptake and retention in Arctic dwarf shrub vegetation. They increase CUE, and the fraction of GPP partitioned aboveground. Arctic C models must include mosses to accurately represent ecosystem C dynamics.

Incident radiation and the allocation of nitrogen within Arctic plant canopies: implications for predicting gross primary productivity.

Arctic vegetation is characterized by high spatial variability in plant functional type (PFT) composition and gross primary productivity (P). Despite this variability, the two main drivers of P in sub-Arctic tundra are leaf area index (LT ) and total foliar nitrogen (NT ). LT and NT have been shown to be tightly coupled across PFTs in sub-Arctic tundra vegetation, which simplifies up-scaling by allowing quantification of the main drivers of P from remotely sensed LT . Our objective was to test the LT -NT relationship across multiple Arctic latitudes and to assess LT as a predictor of P for the pan-Arctic. Including PFT-specific parameters in models of LT -NT coupling provided only incremental improvements in model fit, but significant improvements were gained from including site-specific parameters. The degree of curvature in the LT -NT relationship, controlled by a fitted canopy nitrogen extinction co-efficient, was negatively related to average levels of diffuse radiation at a site. This is consistent with theoretical predictions of more uniform vertical canopy N distributions under diffuse light conditions. Higher latitude sites had higher average leaf N content by mass (NM ), and we show for the first time that LT -NT coupling is achieved across latitudes via canopy-scale trade-offs between NM and leaf mass per unit leaf area (LM ). Site-specific parameters provided small but significant improvements in models of P based on LT and moss cover. Our results suggest that differences in LT -NT coupling between sites could be used to improve pan-Arctic models of P and we provide unique evidence that prevailing radiation conditions can significantly affect N allocation over regional scales.