Divergence of Arctic shrub growth associated with sea ice decline.

Arctic sea ice extent (SIE) is declining at an accelerating rate with a wide range of ecological consequences. However, determining sea ice effects on tundra vegetation remains a challenge. In this study, we examined the universality or lack thereof in tundra shrub growth responses to changes in SIE and summer climate across the Pan-Arctic, taking advantage of 23 tundra shrub-ring chronologies from 19 widely distributed sites (56°N to 83°N). We show a clear divergence in shrub growth responses to SIE that began in the mid-1990s, with 39% of the chronologies showing declines and 57% showing increases in radial growth (decreasers and increasers, respectively). Structural equation models revealed that declining SIE was associated with rising air temperature and precipitation for increasers and with increasingly dry conditions for decreasers. Decreasers tended to be from areas of the Arctic with lower summer precipitation and their growth decline was related to decreases in the standardized precipitation evapotranspiration index. Our findings suggest that moisture limitation, associated with declining SIE, might inhibit the positive effects of warming on shrub growth over a considerable part of the terrestrial Arctic, thereby complicating predictions of vegetation change and future tundra productivity.

Resistance and change in a High Arctic ecosystem, NW Greenland: Differential sensitivity of ecosystem metrics to 15 years of experimental warming and wetting.

Dramatic increases in air temperature and precipitation are occurring in the High Arctic (>70°N), yet few studies have characterized the long-term responses of High Arctic ecosystems to the interactive effects of experimental warming and increased rain. Beginning in 2003, we applied a factorial summer warming and wetting experiment to a polar semidesert in northwest Greenland. In summer 2018, we assessed several metrics of ecosystem structure and function, including plant cover, greenness, ecosystem CO2 exchange, aboveground (leaf, stem) and belowground (litter, root, soil) carbon (C) and nitrogen (N) concentrations (%) and pools, as well as leaf and soil stable isotopes (δ13 C and δ15 N). Wetting induced the most pronounced changes in ecosystem structure, accelerating the expansion of Salix arctica cover by 370% and increasing aboveground C, N, and biomass pools by 94%-101% and root C, N, and biomass pools by 60%-122%, increases which coincided with enhanced net ecosystem CO2 uptake. Further, wetting combined with warming enhanced plot-level greenness, whereas in isolation neither wetting nor warming had an effect. At the plant level, the effects of warming and wetting differed among species and included warming-linked decreases in leaf N and δ15 N in S. arctica, whereas leaf N and δ15 N in Dryas integrifolia did not respond to the climate treatments. Finally, neither plant- nor plot-level C and N allocation patterns nor soil C, N, δ13 C, or δ15 N concentrations changed in response to our manipulations, indicating that these ecosystem metrics may resist climate change, even in the longer term. In sum, our results highlight the importance of summer precipitation in regulating ecosystem structure and function in arid parts of the High Arctic, but they do not completely refute previous findings of resistance in some High Arctic ecosystem properties to climate change.

Winter snow and spring temperature have differential effects on vegetation phenology and productivity across Arctic plant communities.

Tundra dominates two-thirds of the unglaciated, terrestrial Arctic. Although this region has experienced rapid and widespread changes in vegetation phenology and productivity over the last several decades, the specific climatic drivers responsible for this change remain poorly understood. Here we quantified the effect of winter snowpack and early spring temperature conditions on growing season vegetation phenology (timing of the start, peak, and end of the growing season) and productivity of the dominant tundra vegetation communities of Arctic Alaska. We used daily remotely sensed normalized difference vegetation index (NDVI), and daily snowpack and temperature variables produced by SnowModel and MicroMet, coupled physically based snow and meteorological modeling tools, to (1) determine the most important snowpack and thermal controls on tundra vegetation phenology and productivity and (2) describe the direction of these relationships within each vegetation community. Our results show that soil temperature under the snowpack, snowmelt timing, and air temperature following snowmelt are the most important drivers of growing season timing and productivity among Arctic vegetation communities. Air temperature after snowmelt was the most important control on timing of season start and end, with warmer conditions contributing to earlier phenology in all vegetation communities. In contrast, the controls on the timing of peak season and productivity also included snowmelt timing and soil temperature under the snowpack, dictated in part by the snow insulating capacity. The results of this novel analysis suggest that while future warming effects on phenology may be consistent across communities of the tundra biome, warming may result in divergent, community-specific productivity responses if coupled with reduced snow insulating capacity lowers winter soil temperature and potential nutrient cycling in the soil.

Plant and soil nitrogen in oligotrophic boreal forest habitats with varying moss depths: does exclusion of large grazers matter?

The boreal forest consists of drier sunlit and moister-shaded habitats with varying moss abundance. Mosses control vascular plant-soil interactions, yet they all can also be altered by grazers. We determined how 2 decades of reindeer (Rangifer tarandus) exclusion affect feather moss (Pleurozium schreberi) depth, and the accompanying soil N dynamics (total and dissolvable inorganic N, δ15N), plant foliar N, and stable isotopes (δ15N, δ13C) in two contrasting habitats of an oligotrophic Scots pine forest. The study species were pine seedling (Pinus sylvestris L.), bilberry (Vaccinium myrtillus L.), lingonberry (V. vitis-idaea L.), and feather moss. Moss carpet was deeper in shaded than sunlit habitats and increased with grazer exclusion. Humus N content increased in the shade as did humus δ15N, which also increased due to exclusion in the sunlit habitats. Exclusion increased inorganic N concentration in the mineral soil. These soil responses were correlated with moss depth. Foliar chemistry varied due to habitat depending on species identity. Pine seedlings showed higher foliar N content and lower foliar δ15N in the shaded than in the sunlit habitats, while bilberry had both higher foliar N and δ15N in the shade. Thus, foliar δ15N values of co-existing species diverged in the shade indicating enhanced N partitioning. We conclude that despite strong grazing-induced shifts in mosses and subtler shifts in soil N, the N dynamics of vascular vegetation remain unchanged. These indicate that plant-soil interactions are resistant to shifts in grazing intensity, a pattern that appears to be common across boreal oligotrophic forests.

Two Decades of Mercury Concentrations in Barents Sea Polar Bears (Ursus maritimus) in Relation to Dietary Carbon, Sulfur, and Nitrogen.

Temporal trends of total mercury (THg) were examined in female polar bear (Ursus maritimus) hair (n = 199) from the Barents Sea in 1995-2016. In addition, hair values of stable isotopes (n = 190-197) of carbon (δ13C), sulfur (δ34S), and nitrogen (δ15N) and information on breeding status, body condition, and age were obtained. Stable isotope values of carbon and sulfur reflect dietary source (e.g., marine vs terrestrial) and the nitrogen trophic level. Values for δ13C and δ34S declined by -1.62 and -1.18‰ over the time of the study period, respectively, while values for δ15N showed no trend. Total Hg concentrations were positively related to both δ13C and δ34S. Yearly median THg concentrations ranged from 1.61 to 2.75 µg/g and increased nonlinearly by 0.86 µg/g in total over the study. Correcting THg concentrations for stable isotope values of carbon and sulfur and additionally breeding status and age slightly accelerated the increase in THg concentrations; however, confidence intervals of the raw THg trend and the corrected THg trend had substantial overlap. The rise in THg concentrations in the polar bear food web was possibly related to climate-related re-emissions of previously stored Hg from thawing sea-ice, glaciers, and permafrost.

Pacific North American circulation pattern links external forcing and North American hydroclimatic change over the past millennium.

Land and sea surface temperatures, precipitation, and storm tracks in North America and the North Pacific are controlled to a large degree by atmospheric variability associated with the Pacific North American (PNA) pattern. The modern instrumental record indicates a trend toward a positive PNA phase in recent decades, which has led to accelerated warming and snowpack decline in northwestern North America. The brevity of the instrumental record, however, limits our understanding of long-term PNA variability and its directional or cyclic patterns. Here we develop a 937-y-long reconstruction of the winter PNA based on a network of annually resolved tree-ring proxy records across North America. The reconstruction is consistent with previous regional records in suggesting that the recent persistent positive PNA pattern is unprecedented over the past millennium, but documents patterns of decadal-scale variability that contrast with previous reconstructions. Our reconstruction shows that PNA has been strongly and consistently correlated with sea surface temperature variation, solar irradiance, and volcanic forcing over the period of record, and played a significant role in translating these forcings into decadal-to-multidecadal hydroclimate variability over North America. Climate model ensembles show limited power to predict multidecadal variation in PNA over the period of our record, raising questions about their potential to project future hydroclimatic change modulated by this circulation pattern.

Delayed herbivory by migratory geese increases summer-long CO2 uptake in coastal western Alaska.

The advancement of spring and the differential ability of organisms to respond to changes in plant phenology may lead to "phenological mismatches" as a result of climate change. One potential for considerable mismatch is between migratory birds and food availability in northern breeding ranges, and these mismatches may have consequences for ecosystem function. We conducted a three-year experiment to examine the consequences for CO2 exchange of advanced spring green-up and altered timing of grazing by migratory Pacific black brant in a coastal wetland in western Alaska. Experimental treatments represent the variation in green-up and timing of peak grazing intensity that currently exists in the system. Delayed grazing resulted in greater net ecosystem exchange (NEE) and gross primary productivity (GPP), while early grazing reduced CO2 uptake with the potential of causing net ecosystem carbon (C) loss in late spring and early summer. Conversely, advancing the growing season only influenced ecosystem respiration (ER), resulting in a small increase in ER with no concomitant impact on GPP or NEE. The experimental treatment that represents the most likely future, with green-up advancing more rapidly than arrival of migratory geese, results in NEE changing by 1.2 µmol m-2  s-1 toward a greater CO2 sink in spring and summer. Increased sink strength, however, may be mitigated by early arrival of migratory geese, which would reduce CO2 uptake. Importantly, while the direct effect of climate warming on phenology of green-up has a minimal influence on NEE, the indirect effect of climate warming manifest through changes in the timing of peak grazing can have a significant impact on C balance in northern coastal wetlands. Furthermore, processes influencing the timing of goose migration in the winter range can significantly influence ecosystem function in summer habitats.

Temporal Trends of Persistent Organic Pollutants in Barents Sea Polar Bears ( Ursus maritimus) in Relation to Changes in Feeding Habits and Body Condition.

Temporal trends of persistent organic pollutants (POPs: PCBs, OH-PCBs, p, p'-DDE, HCB, ß-HCH, oxychlordane, BDE-47, and 153) in relation to changes in feeding habits and body condition in adult female polar bears ( Ursus maritimus) from the Barents Sea subpopulation were examined over 20 years (1997-2017). All 306 samples were collected in the spring (April). Both stable isotope values of nitrogen (δ15N) and carbon (δ13C) from red blood cells declined over time, with a steeper trend for δ13C between 2012 and 2017, indicating a decreasing intake of marine and high trophic level prey items. Body condition, based on morphometric measurements, had a nonsignificant decreasing tendency between 1997 and 2005, and increased significantly between 2005 and 2017. Plasma concentrations of BDE-153 and ß-HCH did not significantly change over time, whereas concentrations of Σ4PCB, Σ5OH-PCB, BDE-47, and oxychlordane declined linearly. Concentrations of p, p'-DDE and HCB, however, declined until 2012 and 2009, respectively, and increased thereafter. Changes in feeding habits and body condition did not significantly affect POP trends. The study indicates that changes in diet and body condition were not the primary driver of POPs in polar bears, but were controlled in large part by primary and/or secondary emissions of POPs.

Choose Your Poison-Space-Use Strategy Influences Pollutant Exposure in Barents Sea Polar Bears.

Variation in space-use is common within mammal populations. In polar bears, Ursus maritimus, some individuals follow the sea ice (offshore bears) whereas others remain nearshore yearlong (coastal bears). We studied pollutant exposure in relation to space-use patterns (offshore vs coastal) in adult female polar bears from the Barents Sea equipped with satellite collars (2000-2014, n = 152). First, we examined the differences in home range (HR) size and position, body condition, and diet proxies (nitrogen and carbon stable isotopes, n = 116) between offshore and coastal space-use. Second, we investigated how HR, space-use, body condition, and diet were related to plasma concentrations of polychlorinated biphenyls (PCBs), organochlorine pesticides (OCPs) ( n = 113), perfluoroalkyl substances (PFASs; n = 92), and hydroxylated-PCBs ( n = 109). Offshore females were in better condition and had a more specialized diet than did coastal females. PCBs, OCPs, and hydroxylated-PCB concentrations were not related to space-use strategy, yet PCB concentrations increased with increasing latitude, and hydroxylated-PCB concentrations were positively related to HR size. PFAS concentrations were 30-35% higher in offshore bears compared to coastal bears and also increased eastward. On the basis of the results we conclude that space-use of Barents Sea female polar bears influences their pollutant exposure, in particular plasma concentrations of PFAS.

Arctic plant ecophysiology and water source utilization in response to altered snow: isotopic (δ18O and δ2H) evidence for meltwater subsidies to deciduous shrubs.

Warming-linked woody shrub expansion in the Arctic has critical consequences for ecosystem processes and climate feedbacks. The snow-shrub interaction model has been widely implicated in observed Arctic shrub increases, yet equivocal experimental results regarding nutrient-related components of this model have highlighted the need for a consideration of the increased meltwater predicted in expanding shrub stands. We used a 22-year snow manipulation experiment to simultaneously address the unexplored role of snow meltwater in arctic plant ecophysiology and nutrient-related components of the snow-shrub hypothesis. We coupled measurements of leaf-level gas exchange and leaf tissue chemistry (%N and δ13C) with an analysis of stable isotopes (δ18O and δ2H) in soil water, precipitation, and stem water. In deeper snow areas photosynthesis, conductance, and leaf N increased and δ13C values decreased in the deciduous shrubs, Betula nana and Salix pulchra, and the graminoid, Eriophorum vaginatum, with the strongest treatment effects observed in deciduous shrubs, consistent with predictions of the snow-shrub hypothesis. We also found that deciduous shrubs, especially S. pulchra, obtained much of their water from snow melt early in the growing season (40-50%), more than either E. vaginatum or the evergreen shrub, Rhododendron tomentosum (Ledum palustre). This result provides the basis for adding a meltwater-focused feedback loop to the snow-shrub interaction model of shrub expansion in the Arctic. Our results highlight the critical role of winter snow in the ecophysiology of Arctic plants, particularly deciduous shrubs, and underline the importance of understanding how global warming will affect the Arctic winter snowpack.

Greater temperature sensitivity of plant phenology at colder sites: implications for convergence across northern latitudes.

Warmer temperatures are accelerating the phenology of organisms around the world. Temperature sensitivity of phenology might be greater in colder, higher latitude sites than in warmer regions, in part because small changes in temperature constitute greater relative changes in thermal balance at colder sites. To test this hypothesis, we examined up to 20 years of phenology data for 47 tundra plant species at 18 high-latitude sites along a climatic gradient. Across all species, the timing of leaf emergence and flowering was more sensitive to a given increase in summer temperature at colder than warmer high-latitude locations. A similar pattern was seen over time for the flowering phenology of a widespread species, Cassiope tetragona. These are among the first results highlighting differential phenological responses of plants across a climatic gradient and suggest the possibility of convergence in flowering times and therefore an increase in gene flow across latitudes as the climate warms.

Emission Changes Dwarf the Influence of Feeding Habits on Temporal Trends of Per- and Polyfluoroalkyl Substances in Two Arctic Top Predators.

We monitored concentrations of per- and polyfluoroalkyl substances (PFASs) in relation to climate-associated changes in feeding habits and food availability in polar bears (Ursus maritimus) and arctic foxes (Vulpes lagopus) (192 plasma and 113 liver samples, respectively) sampled from Svalbard, Norway, during 1997-2014. PFASs concentrations became greater with increasing dietary trophic level, as bears and foxes consumed more marine as opposed to terrestrial food, and as the availability of sea ice habitat increased. Long-chained perfluoroalkyl carboxylates (PFCAs) in arctic foxes decreased with availability of reindeer carcasses. The ∼9-14% yearly decline of C6-8 perfluoroalkyl sulfonates (PFSAs) following the cease in C6-8 PFSA precursor production in 2001 indicates that the peak exposure was mainly a result of atmospheric transport of the volatile precursors. However, the stable PFSA concentrations since 2009-2010 suggest that Svalbard biota is still exposed to ocean-transported PFSAs. Long-chain ocean-transported PFCAs increased 2-4% per year and the increase in C12-14 PFCAs in polar bears tended to level off since ∼2009. Emerging short-chain PFASs showed no temporal changes. Climate-related changes in feeding habits and food availability moderately affected PFAS trends. Our results indicate that PFAS concentrations in polar bears and arctic foxes are mainly affected by emissions.

Winter precipitation and snow accumulation drive the methane sink or source strength of Arctic tussock tundra.

Arctic winter precipitation is projected to increase with global warming, but some areas will experience decreases in snow accumulation. Although Arctic CH4 emissions may represent a significant climate forcing feedback, long-term impacts of changes in snow accumulation on CH4 fluxes remain uncertain. We measured ecosystem CH4 fluxes and soil CH4 and CO2 concentrations and (13) C composition to investigate the metabolic pathways and transport mechanisms driving moist acidic tundra CH4 flux over the growing season (Jun-Aug) after 18 years of experimental snow depth increases and decreases. Deeper snow increased soil wetness and warming, reducing soil %O2 levels and increasing thaw depth. Soil moisture, through changes in soil %O2 saturation, determined predominance of methanotrophy or methanogenesis, with soil temperature regulating the ecosystem CH4 sink or source strength. Reduced snow (RS) increased the fraction of oxidized CH4 (Fox) by 75-120% compared to Ambient, switching the system from a small source to a net CH4 sink (21 ± 2 and -31 ± 1 mg CH4  m(-2)  season(-1) at Ambient and RS). Deeper snow reduced Fox by 35-40% and 90-100% in medium- (MS) and high- (HS) snow additions relative to Ambient, contributing to increasing the CH4 source strength of moist acidic tundra (464 ± 15 and 3561 ± 97 mg CH4  m(-2)  season(-1) at MS and HS). Decreases in Fox with deeper snow were partly due to increases in plant-mediated CH4 transport associated with the expansion of tall graminoids. Deeper snow enhanced CH4 production within newly thawed soils, responding mainly to soil warming rather than to increases in acetate fermentation expected from thaw-induced increases in SOC availability. Our results suggest that increased winter precipitation will increase the CH4 source strength of Arctic tundra, but the resulting positive feedback on climate change will depend on the balance between areas with more or less snow accumulation than they are currently facing.

Long-term increase in snow depth leads to compositional changes in arctic ectomycorrhizal fungal communities.

Many arctic ecological processes are regulated by soil temperature that is tightly interconnected with snow cover distribution and persistence. Recently, various climate-induced changes have been observed in arctic tundra ecosystems, e.g. shrub expansion, resulting in reduction in albedo and greater C fixation in aboveground vegetation as well as increased rates of soil C mobilization by microbes. Importantly, the net effects of these shifts are unknown, in part because our understanding of belowground processes is limited. Here, we focus on the effects of increased snow depth, and as a consequence, increased winter soil temperature on ectomycorrhizal (ECM) fungal communities in dry and moist tundra. We analyzed deep DNA sequence data from soil samples taken at a long-term snow fence experiment in Northern Alaska. Our results indicate that, in contrast with previously observed responses of plants to increased snow depth at the same experimental site, the ECM fungal community of the dry tundra was more affected by deeper snow than the moist tundra community. In the dry tundra, both community richness and composition were significantly altered while in the moist tundra, only community composition changed significantly while richness did not. We observed a decrease in richness of Tomentella, Inocybe and other taxa adapted to scavenge the soil for labile N forms. On the other hand, richness of Cortinarius, and species with the ability to scavenge the soil for recalcitrant N forms, did not change. We further link ECM fungal traits with C soil pools. If future warmer atmospheric conditions lead to greater winter snow fall, changes in the ECM fungal community will likely influence C emissions and C fixation through altering N plant availability, fungal biomass and soil-plant C-N dynamics, ultimately determining important future interactions between the tundra biosphere and atmosphere.

Tree-ring isotopes reveal drought sensitivity in trees killed by spruce beetle outbreaks in south-central Alaska.

Increasing temperatures have resulted in reduced growth and increased tree mortality across large areas of western North American forests. We use tree-ring isotope chronologies (δ13 C and δ18 O) from live and dead trees from four locations in south-central Alaska, USA, to test whether white spruce trees killed by recent spruce beetle (Dendroctonus rufipennis Kirby) outbreaks showed evidence of drought stress prior to death. Trees that were killed were more sensitive to spring/summer temperature and/or precipitation than trees that survived. At two of our sites, we found greater correlations between the δ13 C and δ18 O chronologies and spring/summer temperatures in dead trees than in live trees, suggesting that trees that are more sensitive to temperature-induced drought stress are more likely to be killed. At one site, the difference between δ13 C in live and dead trees was related to winter/spring precipitation, with dead trees showing stronger correlations between δ13 C and precipitation, again suggesting increased water stress in dead trees. At all sites where δ18 O was measured, δ18 O chronologies showed the greatest difference in climate response between live and dead groups, with δ18 O in live trees correlating more strongly with late winter precipitation than dead trees. Our results indicate that sites where trees are already sensitive to warm or dry early growing-season conditions experienced the most beetle-kill, which has important implications for forecasting future mortality events in Alaska.

Changes in composition and abundance of functional groups of arctic fungi in response to long-term summer warming.

We characterized fungal communities in dry and moist tundra and investigated the effect of long-term experimental summer warming on three aspects of functional groups of arctic fungi: richness, community composition and species abundance. Warming had profound effects on community composition, abundance, and, to a lesser extent, on richness of fungal functional groups. In addition, our data show that even within functional groups, the direction and extent of response to warming tend to be species-specific and we recommend that studies on fungal communities and their roles in nutrient cycling take into account species-level responses.

Coupled long-term summer warming and deeper snow alters species composition and stimulates gross primary productivity in tussock tundra.

Climate change is expected to increase summer temperature and winter precipitation throughout the Arctic. The long-term implications of these changes for plant species composition, plant function, and ecosystem processes are difficult to predict. We report on the influence of enhanced snow depth and warmer summer temperature following 20 years of an ITEX experimental manipulation at Toolik Lake, Alaska. Winter snow depth was increased using snow fences and warming was accomplished during summer using passive open-top chambers. One of the most important consequences of these experimental treatments was an increase in active layer depth and rate of thaw, which has led to deeper drainage and lower soil moisture content. Vegetation concomitantly shifted from a relatively wet system with high cover of the sedge Eriophorum vaginatum to a drier system, dominated by deciduous shrubs including Betula nana and Salix pulchra. At the individual plant level, we observed higher leaf nitrogen concentration associated with warmer temperatures and increased snow in S. pulchra and B. nana, but high leaf nitrogen concentration did not lead to higher rates of net photosynthesis. At the ecosystem level, we observed higher GPP and NEE in response to summer warming. Our results suggest that deeper snow has a cascading set of biophysical consequences that include a deeper active layer that leads to altered species composition, greater leaf nitrogen concentration, and higher ecosystem-level carbon uptake.

Long-term experimental warming alters community composition of ascomycetes in Alaskan moist and dry arctic tundra.

Arctic tundra regions have been responding to global warming with visible changes in plant community composition, including expansion of shrubs and declines in lichens and bryophytes. Even though it is well known that the majority of arctic plants are associated with their symbiotic fungi, how fungal community composition will be different with climate warming remains largely unknown. In this study, we addressed the effects of long-term (18 years) experimental warming on the community composition and taxonomic richness of soil ascomycetes in dry and moist tundra types. Using deep Ion Torrent sequencing, we quantified how OTU assemblage and richness of different orders of Ascomycota changed in response to summer warming. Experimental warming significantly altered ascomycete communities with stronger responses observed in the moist tundra compared with dry tundra. The proportion of several lichenized and moss-associated fungi decreased with warming, while the proportion of several plant and insect pathogens and saprotrophic species was higher in the warming treatment. The observed alterations in both taxonomic and ecological groups of ascomycetes are discussed in relation to previously reported warming-induced shifts in arctic plant communities, including decline in lichens and bryophytes and increase in coverage and biomass of shrubs.

Summer temperature increase has distinct effects on the ectomycorrhizal fungal communities of moist tussock and dry tundra in Arctic Alaska.

Arctic regions are experiencing the greatest rates of climate warming on the planet and marked changes have already been observed in terrestrial arctic ecosystems. While most studies have focused on the effects of warming on arctic vegetation and nutrient cycling, little is known about how belowground communities, such as fungi root-associated, respond to warming. Here, we investigate how long-term summer warming affects ectomycorrhizal (ECM) fungal communities. We used Ion Torrent sequencing of the rDNA internal transcribed spacer 2 (ITS2) region to compare ECM fungal communities in plots with and without long-term experimental warming in both dry and moist tussock tundra. Cortinarius was the most OTU-rich genus in the moist tundra, while the most diverse genus in the dry tundra was Tomentella. On the diversity level, in the moist tundra we found significant differences in community composition, and a sharp decrease in the richness of ECM fungi due to warming. On the functional level, our results indicate that warming induces shifts in the extramatrical properties of the communities, where the species with medium-distance exploration type seem to be favored with potential implications for the mobilization of different nutrient pools in the soil. In the dry tundra, neither community richness nor community composition was significantly altered by warming, similar to what had been observed in ECM host plants. There was, however, a marginally significant increase in OTUs identified as ECM fungi with the medium-distance exploration type in the warmed plots. Linking our findings of decreasing richness with previous results of increasing ECM fungal biomass suggests that certain ECM species are favored by warming and may become more abundant, while many other species may go locally extinct due to direct or indirect effects of warming. Such compositional shifts in the community might affect nutrient cycling and soil organic C storage.

Differential ecophysiological response of deciduous shrubs and a graminoid to long-term experimental snow reductions and additions in moist acidic tundra, Northern Alaska.

Changes in winter precipitation that include both decreases and increases in winter snow are underway across the Arctic. In this study, we used a 14-year experiment that has increased and decreased winter snow in the moist acidic tussock tundra of northern Alaska to understand impacts of variation in winter snow depth on summer leaf-level ecophysiology of two deciduous shrubs and a graminoid species, including: instantaneous rates of leaf gas exchange, and δ(13)C, δ(15)N, and nitrogen (N) concentrations of Betula nana, Salix pulchra, and Eriophorum vaginatum. Leaf-level measurements were complemented by measurements of canopy leaf area index (LAI) and depth of thaw. Reductions in snow lowered summer leaf photosynthesis, conductance, and transpiration rates by up to 40% compared to ambient and deep snow conditions for Eriophorum vaginatum, and reduced Salix pulchra conductance and transpiration by up to 49%. In contrast, Betula nana exhibited no changes in leaf gas exchange in response to lower or deeper snow. Canopy LAI increased with added snow, while reduced winter snow resulted in lower growing season soil temperatures and reduced thaw depths. Our findings indicate that the spatial and temporal variability of future snow depth will have individualistic consequences for leaf-level C fixation and water flux by tundra species, and that these responses will be manifested over the longer term by changes in canopy traits, depth of thaw, soil C and N processes, and trace gas (CO2 and H2O) exchanges between the tundra and the atmosphere.