Nutrient limitation of primary producers and their consumers can have a large influence on ecosystem productivity. The nature and strength of nutrient limitation is driven both by external factors (e.g., nutrient loading) and internal processes (e.g., consumer-driven nutrient regeneration). Here we present results from a field study in 10 low productive headwater lakes in northern subarctic Sweden, where nitrogen (N) deposition is low and phytoplankton is primarily N-limited. We assessed the carbon:nitrogen:phosphorus (C:N:P) stoichiometry of seston and zooplankton and estimated the N:P ratio of consumer-driven nutrient regeneration. Based on stoichiometric models, the estimated elemental imbalances between seston and zooplankton suggest that zooplankton were mainly N-limited and regenerated nutrients with low N:P ratios (median 11.9, atomic ratio). The predicted N:P regeneration ratios were consistent with results from phytoplankton nutrient limitation bioassays in mid-summer, i.e., the N:P regeneration was predicted to be low when phytoplankton were N-limited, and high when phytoplankton were P-limited. During other seasons, when water discharge was high, nutrient loading from the surrounding catchments apparently had the strongest effect on phytoplankton nutrient limitation. We propose that lakes with higher N:P ratios than the open ocean is an effect of N deposition, that N-limitation of consumers and phytoplankton is further enhanced by low nutrient regeneration N:P ratios, and that in the absence of N deposition, lake and ocean N:P stoichiometry are similar.

Long-term observational studies have detected greening and shrub encroachment in the subarctic attributed to current climate change, while global change simulations have showed that community composition and productivity may shift drastically in arctic, subarctic, and alpine tundra plant communities in the future. However, responses to global change can be highly species- and context-dependent. We examined community-level and species-specific responses to a six-year factorial temperature and nutrient (nitrogen and phosphorus) amendment experiment in two alpine plant communities in northern Sweden: a species-poor dwarf shrub heath, and a more species-rich meadow. We hypothesized that abundance responses to global change would be variable within commonly defined vascular plant functional groups (e.g., forbs, evergreen shrubs, deciduous shrubs) and that new species would appear in experimental plots over time due to the ameliorated growing conditions. We found that within most functional groups, species were highly individualistic with respect to the global change simulation, particularly within the forbs, whereas within the shrubs, responses were neutral to negative and widely variable in magnitude. In the heath community the response of the graminoid functional group was driven almost entirely by the grass Calamagrostis lapponica, which increased in abundance by an order of magnitude in the combined temperature and nutrient treatment. Furthermore, community context was important for species’ responses to the simulations. Abundance of the pan-arctic species Carex bigelowii and Vaccinium vitis-idaea responded primarily to the temperature treatment in the meadow community whereas the nutrient treatment had stronger effects in the heath community. Over six growing seasons, more new species appeared in the experimental plots than in control plots in the meadow community, whereas in the heath community only one new species appeared. Our results from two closely situated but different plant communities show that functional groups do not predict individual species responses to simulated global change, and that these responses depend to a large extent on pre-existing physical conditions as well as biotic interactions such as competition and facilitation. It may be difficult to apply general trends of global change responses to specific local communities.

Arctic tundra consists of diverse habitats that differ in dominant vegetation, soil moisture regimes, and relative importance of organic vs. inorganic nutrient cycling. The Arctic is also the most rapidly warming global area, with winter warming dominating. This warming is expected to have dramatic effects on tundra carbon and nutrient dynamics. We completed a meta-analysis of 166 experimental warming study papers to evaluate the hypotheses that warming changes tundra biogeochemical cycles in a habitat- and seasonally specific manner and that the carbon (C), nitrogen (N), and phosphorus (P) cycles will be differentially accelerated, leading to decoupling of elemental cycles. We found that nutrient availability and plant leaf stoichiometry responses to experimental warming were variable and overall weak, but that both gross primary productivity and the plant C pool tended to increase with growing season warming. The effects of winter warming on C fluxes did not extend into the growing season. Overall, although warming led to more consistent increases in C fluxes compared to N or P fluxes, evidence for decoupling of biogeochemical cycles is weak and any effect appears limited to heath habitats. However, data on many habitats are too sparse to be able to generalize how warming might decouple biogeochemical cycles, and too few year-round warming studies exist to ascertain whether the season under which warming occurs alters how ecosystems respond to warming. Coordinated field campaigns are necessary to more robustly document tundra habitat-specific responses to realistic climate warming scenarios in order to better understand the mechanisms driving this heterogeneity and identify the tundra habitats, communities, and soil pools most susceptible to warming.

Ectomycorrhizal fungi (EMF) form symbioses with a few plant species that comprise a large fraction of the arctic vegetation. Despite their importance, the identity, abundance and distribution of EMF in the Arctic, as well as the key drivers controlling their community composition are poorly understood. In this study, we investigated the diversity and structure of EMF communities across a bioclimatic gradient spanning much of the North American Arctic. We collected roots from two principal arctic ectomycorrhizal host plants, Salix arctica and Dryas integrifolia, typically growing intermingled, at 23 locations stratified across the five bioclimatic subzones of the Arctic. DNA was extracted from ectomycorrhizal root tips and the ITS region was sequenced and phylogenetically analyzed. A total of 242 fungal Operational Taxonomic Units (OTUs) were documented, with 203 OTUs belonging to the Basidiomycota and 39 to the Ascomycota, exceeding the number of previously morphologically described EMF in the Arctic. EMF communities were dominated by a few common and species-rich families such as Thelephoraceae, Inocybaceae, Sebacinaceae, Cortinariaceae, and Pyronemataceae. Both host plants showed similar species richness, with 176 OTUs on Salix arctica and 154 OTUs on Dryas integrifolia. Host plant identity did not affect EMF community composition. The ten most abundant OTUs had a wide geographic distribution throughout the Arctic, and were also found in boreal, temperate and Mediterranean regions, where they were associated with a variety of hosts. Species richness did not decline with increasing latitude. However, EMF community structure changed gradually across the bioclimatic gradient with the greatest similarity between neighboring bioclimatic subzones and locations. EMF community structure was correlated with environmental factors at a regional scale, corresponding to a complex of glaciation history, geology, soil properties, plant productivity and climate. This is the first large-scale study of EMF communities across all five bioclimatic subzones of the North American Arctic, accompanied by an extensive set of environmental factors analyzed to date. While our study provides baseline data to assess shifts of plant and fungi distribution in response to climate change, it also suggests that with ongoing climate warming, EMF community composition may be affected by northward shifts of some taxa.

Shrubs commonly form islands of fertility and are expanding their distribution and dominance in the arctic due to climate change, yet how soil properties may be influenced when different species of shrubs expand under warmer climates remains less explored. Important plant traits, such as their associated root community, are linked to functionally different and dominant shrub species in the arctic and these traits likely shape biogeochemical cycling in areas of shrub expansion. Using an elevational gradient as a proxy for warming, we explored how biochemical processes beneath two important arctic shrubs varied under warmer (low elevation) and cooler (high elevation) climates. Interestingly, the influence of elevation on biogeochemistry varied between the two shrubs. At the low elevation, Betula nana L., an ectomycorrhizal shrub, had high carbon (C) degrading enzyme activities, and relatively low potential net nitrogen (N) mineralization rates. Conversely, Empetrum nigrum ssp. hermaphroditum Hagerup, an ericoid mycorrhizal dwarf-shrub, had higher enzyme activities and net N immobilization rates at the higher elevation. Further, E. nigrum ssp. hermpahroditum appeared to have a more closed C and nutrient cycle than B. nana?enzymes degrading C, N, and phosphorus were tightly correlated with each other and with total C and ammonium concentrations in the humus beneath E. nigrum ssp. hermaphroditum, but not beneath B. nana. Our results suggest differences in the warming responses of C and N cycling beneath shrub species across an arctic tundra landscape.