Climate scenarios for high-latitude areas predict not only increased summer temperatures, but also larger variation in snowfall and winter temperatures. By using open-top chambers, we experimentally manipulated both summer temperatures and winter and spring snow accumulations and temperatures independently in a blanket bog in subarctic Sweden, yielding six climate scenarios. We studied the effects of these scenarios on flowering phenology and flower production of Andromeda polifolia (woody evergreen) and Rubus chamaemorus (perennial herb) during 2 years. The second year of our study (2002) was characterized by unusually high spring and early summer temperatures. Our winter manipulations led to consistent increases in winter snow cover. As a result, average and minimum air and soil temperatures in the high snow cover treatments were higher than in the winter ambient treatments, whereas temperature fluctuations were smaller. Spring warming resulted in higher average, minimum, and maximum soil temperatures. Summer warming led to higher air and soil temperatures in mid-summer (June-July), but not in late summer (August-September). The unusually high temperatures in 2002 advanced the median flowering date by 2 weeks for both species in all treatments. Superimposed on this effect, we found that for both Andromeda and Rubus, all our climate treatments (except summer warming for Rubus) advanced flowering by 1-4 days. The total flower production of both species showed a more or less similar response: flower production in the warm year 2002 exceeded that in 2001 by far. However, in both species flower production was only stimulated by the spring-warming treatments. Our results show that the reproductive ecology of both species is very responsive to climate change but this response is very dependent on specific climate events, especially those that occur in winter and spring. This suggests that high-latitude climate change experiments should focus more on winter and spring events than has been the case so far.

Traditionally, biogenic volatile organic compound (BVOC) emissions are often considered a unidirectional flux, from the ecosystem to the atmosphere, but recent studies clearly show the potential for bidirectional exchange. Here we aimed to investigate how warming and leaf litter addition affect the bidirectional exchange (flux) of BVOCs in a long-term field experiment in the Subarctic. We also assessed changes in net BVOC fluxes in relation to the time of day and the influence of different plant phenological stages. The study was conducted in a full factorial experiment with open top chamber warming and annual litter addition treatments in a tundra heath in Abisko, Northern Sweden. After 18 years of treatments, ecosystem-level net BVOC fluxes were measured in the experimental plots using proton-transfer-reaction time-of-flight mass spectrometry (PTR?ToF?MS). The warming treatment increased monoterpene and isoprene emissions by ≈50%. Increasing temperature, due to diurnal variations, can both increase BVOC emission and simultaneously, increase ecosystem uptake. For any given treatment, monoterpene, isoprene, and acetone emissions also increased with increasing ambient air temperatures caused by diurnal variability. Acetaldehyde, methanol, and sesquiterpenes decreased likely due to a deposition flux. For litter addition, only a significant indirect effect on isoprene and monoterpene fluxes (decrease by ~50%?75%) was observed. Litter addition may change soil moisture conditions, leading to changes in plant species composition and biomass, which could subsequently result in changes to BVOC emission compositions. Phenological stages significantly affected fluxes of methanol, isoprene and monoterpenes. We suggest that plant phenological stages differ in impacts on BVOC net emissions, but ambient air temperature and photosynthetically active radiation (PAR) also interact and influence BVOC net emissions differently. Our results may also suggest that BVOC fluxes are not only a response to changes in temperature and light intensity, as the circadian clock also affects emission rates.

The aim of this study was to predict the combined effects of enhanced nitrogen (N) deposition and warming on phytoplankton development in high latitude and mountain lakes. Consequently, we assessed, in a series of enclosureexperiments, how lake water nutrient stoichiometry and phytoplankton nutrient limitation varied over the growingseason in 11 lakes situated along an altitudinal/climate gradient with low N-deposition (<1 kg N ha1yr1) in northern subarctic Sweden. Short-term bioassay experiments with N- and P-additions revealed that phytoplankton inhigh-alpine lakes were more prone to P-limitation, and with decreasing altitude became increasingly N- andNP-colimited. Nutrient limitation was additionally most obvious in midsummer. There was also a strong positivecorrelation between phytoplankton growth and water temperature in the bioassays. Although excess nutrients wereavailable in spring and autumn, on these occasions growth was likely constrained by low water temperatures. Theseresults imply that enhanced N-deposition over the Swedish mountain areas will, with the exception of high-alpinelakes, enhance biomass and drive phytoplankton from N- to P-limitation. However, if not accompanied by warming,N-input from deposition will stimulate limited phytoplankton growth due to low water temperatures during largeparts of the growing season. Direct effects of warming, allowing increased metabolic rates and an extension of thegrowing season, seem equally crucial to synergistically enhance phytoplankton development in these lakes.

Global change affects terrestrial loadings of colored dissolved organic carbon (DOC) and nutrients to northern lakes. Still, little is known about how phytoplankton respond to changes in light and nutrient availability across gradients in lake DOC. In this study, we used results from whole‐lake studies in northern Sweden to show that annual mean phytoplankton biomass expressed unimodal curved relationships across lake DOC gradients, peaking at threshold DOC levels of around 11 mg/L. Whole‐lake single nutrient enrichment in selected lakes caused elevated biomass, with most pronounced effect at the threshold DOC level. These patterns give support to the suggested dual control by DOC on phytoplankton via nutrient (positively) and light (negatively) availability and imply that the lakes' location along the DOC axis is critical in determining to what extent phytoplankton respond to changes in DOC and/or nutrient loadings. By using data from the large Swedish Lake Monitoring Survey, we further estimated that 80% of northern Swedish lakes are below the DOC threshold, potentially experiencing increased phytoplankton biomass with browning alone, and/or combined with nutrient enrichment. The results support the previous model results on effects of browning and eutrophication on lake phytoplankton, and provide important understanding of how northern lakes may respond to future global changes.

Climate change scenarios predict an increased frequency of extreme climatic events. In Arctic regions, one of the most profound of these are extreme and sudden winter warming events in which temperatures increase rapidly to above freezing, often causing snow melt across whole landscapes and exposure of ecosystems to warm temperatures. Following warming, vegetation and soils no longer insulated below snow are then exposed to rapidly returning extreme cold. Using a new experimental facility established in sub-Arctic dwarf shrub heathland in northern Sweden, we simulated an extreme winter warming event in the field and report findings on growth, phenology and reproduction during the subsequent growing season. A 1-week long extreme winter warming event was simulated in early March using infrared heating lamps run with or without soil warming cables. Both single short events delayed bud development of Vaccinium myrtillus by up to 3 weeks in the following spring (June) and reduced flower production by more than 80%: this also led to a near-complete elimination of berry production in mid-summer. Empetrum hermaphroditum also showed delayed bud development. In contrast, Vaccinium vitis-idaea showed no delay in bud development, but instead appeared to produce a greater number of actively growing vegetative buds within plots warmed by heating lamps only. Again, there was evidence of reduced flowering and berry production in this species. While bud break was delayed, growing season measurements of growth and photosynthesis did not reveal a differential response in the warmed plants for any of the species. These results demonstrate that a single, short, extreme winter warming event can have considerable impact on bud production, phenology and reproductive effort of dominant plant species within sub-Arctic dwarf shrub heathland. Furthermore, large interspecific differences in sensitivity are seen. These findings are of considerable concern, because they suggest that repeated events may potentially impact on the biodiversity and productivity of these systems should these extreme events increase in frequency as a result of global change. Although climate change may lengthen the growing season by earlier spring snow melt, there is a profound danger for these high-latitude ecosystems if extreme, short-lived warming in winter exposes plants to initial warm temperatures, but then extreme cold for the rest of the winter. Work is ongoing to determine the longer term and wider impacts of these events.

Extreme weather events can have strong negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme, short‐lived, winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (for instance, 2–10 °C for 2–14 days) but upon return to normal winter climate exposes the ecosystem to much colder temperatures due to the loss of insulating snow. Single events have been shown to reduce plant reproduction and increase shoot mortality, but impacts of multiple events are little understood as are the broader impacts on community structure, growth, carbon balance, and nutrient cycling. To address these issues, we simulated week‐long extreme winter warming events – using infrared heating lamps and soil warming cables – for 3 consecutive years in a sub‐Arctic heathland dominated by the dwarf shrubs Empetrum hermaphroditum, Vaccinium vitis‐idaea (both evergreen) and Vaccinium myrtillus (deciduous). During the growing seasons after the second and third winter event, spring bud burst was delayed by up to a week for E. hermaphroditum and V. myrtillus, and berry production reduced by 11–75% and 52–95% for E. hermaphroditum and V. myrtillus, respectively. Greater shoot mortality occurred in E. hermaphroditum (up to 52%), V. vitis‐idaea (51%), and V. myrtillus (80%). Root growth was reduced by more than 25% but soil nutrient availability remained unaffected. Gross primary productivity was reduced by more than 50% in the summer following the third simulation. Overall, the extent of damage was considerable, and critically plant responses were opposite in direction to the increased growth seen in long‐term summer warming simulations and the ‘greening’ seen for some arctic regions. Given the Arctic is warming more in winter than summer, and extreme events are predicted to become more frequent, this generates large uncertainty in our current understanding of arctic ecosystem responses to climate change.

Extreme weather events can have negative impacts on species survival and community structure when surpassing lethal thresholds. Extreme winter warming events in the Arctic rapidly melt snow and expose ecosystems to unseasonably warm air (2–10 °C for 2–14 days), but returning to cold winter climate exposes the ecosystem to lower temperatures by the loss of insulating snow. Soil animals, which play an integral part in soil processes, may be very susceptible to such events depending on the intensity of soil warming and low temperatures following these events. We simulated week‐long extreme winter warming events – using infrared heating lamps, alone or with soil warming cables – for two consecutive years in a sub‐Arctic dwarf shrub heathland. Minimum temperatures were lower and freeze‐thaw cycles were 2–11 times more frequent in treatment plots compared with control plots. Following the second event, Acari populations decreased by 39%; primarily driven by declines of Prostigmata (69%) and the Mesostigmatic nymphs (74%). A community‐weighted vertical stratification shift occurred from smaller soil dwelling (eu‐edaphic) Collembola species dominance to larger litter dwelling (hemi‐edaphic) species dominance in the canopy‐with‐soil warming plots compared with controls. The most susceptible groups to these winter warming events were the smallest individuals (Prostigmata and eu‐edaphic Collembola). This was not apparent from abundance data at the Collembola taxon level, indicating that life forms and species traits play a major role in community assembly following extreme events. The observed shift in soil community can cascade down to the micro‐flora affecting plant productivity and mineralization rates. Short‐term extreme weather events have the potential to shift community composition through trait composition with potentially large consequences for ecosystem development.

Environmental manipulation studies are integral to determining biological consequences of climate warming. Open Top Chambers (OTCs) have been widely used to assess summer warming effects on terrestrial biota, with their effects during other seasons normally being given less attention even though chambers are often deployed year-round. In addition, their effects on temperature extremes and freeze-thaw events are poorly documented. To provide robust documentation of the microclimatic influences of OTCs throughout the year, we analysed temperature data from 20 studies distributed across polar and alpine regions. The effects of OTCs on mean temperature showed a large range (-0.9 to 2.1 degrees C) throughout the year, but did not differ significantly between studies. Increases in mean monthly and diurnal temperature were strongly related (R-2 = 0.70) with irradiance, indicating that PAR can be used to predict the mean warming effect of OTCs. Deeper snow trapped in OTCs also induced higher temperatures at soil/vegetation level. OTC-induced changes in the frequency of freeze-thaw events included an increase in autumn and decreases in spring and summer. Frequency of high-temperature events in OTCs increased in spring, summer and autumn compared with non-manipulated control plots. Frequency of low-temperature events was reduced by deeper snow accumulation and higher mean temperatures. The strong interactions identified between aspects of ambient environmental conditions and effects of OTCs suggest that a detailed knowledge of snow depth, temperature and irradiance levels enables us to predict how OTCs will modify the microclimate at a particular site and season. Such predictive power allows a better mechanistic understanding of observed biotic response to experimental warming studies and for more informed design of future experiments. However, a need remains to quantify OTC effects on water availability and wind speed (affecting, for example, drying rates and water stress) in combination with microclimate measurements at organism level.

Climate change impacts are not uniform across the Arctic region because interacting factors causes large variations in local ecosystem change. Extreme climatic events and population cycles of herbivores occur simultaneously against a background of gradual climate warming trends and can redirect ecosystem change along routes that are difficult to predict. Here, we present the results from sub-Arctic heath vegetation and its belowground micro-arthropod community in response to the two main drivers of vegetation damage in this region: extreme winter warming events and subsequent outbreaks of the defoliating autumnal moth caterpillar (Epirrita autumnata). Evergreen dwarf shrub biomass decreased (30%) following extreme winter warming events and again by moth caterpillar grazing. Deciduous shrubs that were previously exposed to an extreme winter warming event were not affected by the moth caterpillar grazing, while those that were not exposed to warming events (control plots) showed reduced (23%) biomass from grazing. Cryptogam cover increased irrespective of grazing or winter warming events. Micro-arthropods declined (46%) following winter warming but did not respond to changes in plant community. Extreme winter warming and caterpillar grazing suppressed the CO2 fluxes of the ecosystem. Evergreen dwarf shrubs are disadvantaged in a future sub-Arctic with more stochastic climatic and biotic events. Given that summer warming may further benefit deciduous over evergreen shrubs, event and trend climate change may both act against evergreen shrubs and the ecosystem functions they provide. This is of particular concern given that Arctic heath vegetation is typically dominated by evergreen shrubs. Other components of the vegetation showed variable responses to abiotic and biotic events, and their interaction indicates that sub-Arctic vegetation response to multiple pressures is not easy to predict from single-factor responses. Therefore, while biotic and climatic events may have clear impacts, more work is needed to understand their net effect on Arctic ecosystems.

Global climate and land use change are causing woody plant encroachment in arctic, alpine, and arid/semi-arid ecosystems around the world, yet our understanding of the belowground impacts of this phenomenon is limited. We conducted a globally distributed field study of 13 alpine sites across four continents undergoing woody plant encroachment and sampled soils from both woody encroached and nearby herbaceous plant community types. We found that woody plant encroachment influenced soil microbial richness and community composition across sites based on multiple factors including woody plant traits, site level climate, and abiotic soil conditions. In particular, root symbiont type was a key determinant of belowground effects, as Nitrogen-fixing woody plants had higher soil fungal richness, while Ecto/Ericoid mycorrhizal species had higher soil bacterial richness and symbiont types had distinct soil microbial community composition. Woody plant leaf traits indirectly influenced soil microbes through their impact on soil abiotic conditions, primarily soil pH and C:N ratios. Finally, site-level climate affected the overall magnitude and direction of woody plant influence, as soil fungal and bacterial richness were either higher or lower in woody encroached versus herbaceous soils depending on mean annual temperature and precipitation. All together, these results document global impacts of woody plant encroachment on soil microbial communities, but highlight that multiple biotic and abiotic pathways must be considered to scale up globally from site- and species-level patterns. Considering both the aboveground and belowground effects of woody encroachment will be critical to predict future changes in alpine ecosystem structure and function and subsequent feedbacks to the global climate system.

Northern ecosystems are experiencing some of the most dramatic impacts of global change on Earth. Rising temperatures, hydrological intensification, changes in atmospheric acid deposition and associated acidification recovery, and changes in vegetative cover are resulting in fundamental changes in terrestrial–aquatic biogeochemical linkages. The effects of global change are readily observed in alterations in the supply of dissolved organic matter (DOM)—the messenger between terrestrial and lake ecosystems—with potentially profound effects on the structure and function of lakes. Northern terrestrial ecosystems contain substantial stores of organic matter and filter or funnel DOM, affecting the timing and magnitude of DOM delivery to surface waters. This terrestrial DOM is processed in streams, rivers, and lakes, ultimately shifting its composition, stoichiometry, and bioavailability. Here, we explore the potential consequences of these global change-driven effects for lake food webs at northern latitudes. Notably, we provide evidence that increased allochthonous DOM supply to lakes is overwhelming increased autochthonous DOM supply that potentially results from earlier ice-out and a longer growing season. Furthermore, we assess the potential implications of this shift for the nutritional quality of autotrophs in terms of their stoichiometry, fatty acid composition, toxin production, and methylmercury concentration, and therefore, contaminant transfer through the food web. We conclude that global change in northern regions leads not only to reduced primary productivity but also to nutritionally poorer lake food webs, with discernible consequences for the trophic web to fish and humans.

Abstract Slow-colonizing forest understorey plants are probably not able to rapidly adjust their distribution range following large-scale climate change. Therefore, the acclimation potential to climate change within their actual occupied habitats will likely be key for their short- and long-term persistence. We combined transplant experiments along a latitudinal gradient with open-top chambers to assess the effects of temperature on phenology, growth and reproductive performance of multiple populations of slow-colonizing understorey plants, using the spring flowering geophytic forb Anemone nemorosa and the early summer flowering grass Milium effusum as study species. In both species, emergence time and start of flowering clearly advanced with increasing temperatures. Vegetative growth (plant height, aboveground biomass) and reproductive success (seed mass, seed germination and germinable seed output) of A. nemorosa benefited from higher temperatures. Climate warming may thus increase future competitive ability and colonization rates of this species. Apart from the effects on phenology, growth and reproductive performance of M. effusum generally decreased when transplanted southwards (e.g., plant size and number of individuals decreased towards the south) and was probably more limited by light availability in the south. Specific leaf area of both species increased when transplanted southwards, but decreased with open-top chamber installation in A. nemorosa. In general, individuals of both species transplanted at the home site performed best, suggesting local adaptation. We conclude that contrasting understorey plants may display divergent plasticity in response to changing temperatures which may alter future understorey community dynamics.

Sphagnum mosses form a major component of northern peatlands, which are expected to experience substantially higher increases in temperature and winter precipitation than the global average. Sphagnum may play an important role in the responses of the global carbon cycle to climate change. We investigated the responses of summer length growth, carpet structure and production in Sphagnum fuscum to experimentally induced changes in climate in a sub-arctic bog. Thereto, we used open-top chambers (OTCs) to create six climate scenarios including changes in summer temperatures, and changes in winter snow cover and spring temperatures. In winter, the OTCs doubled the snow thickness, resulting in 0.5-2.8degreesC higher average air temperatures. Spring air temperatures in OTCs increased by 1.0degreesC. Summer warming had a maximum effect of 0.9degreesC, while vapor pressure deficit was not affected. The climate manipulations had strong effects on S. fuscum. Summer warming enhanced the length increment by 42-62%, whereas bulk density decreased. This resulted in a trend (P<0.10) of enhanced biomass production. Winter snow addition enhanced dry matter production by 33%, despite the fact that the length growth and bulk density did not change significantly. The addition of spring warming to snow addition alone did not significantly enhance this effect, but we may have missed part of the early spring growth. There were no interactions between the manipulations in summer and those in winter/spring, indicating that the effects were additive. Summer warming may in the long term negatively affect productivity through the adverse effects of changes in Sphagnum structure on moisture holding and transporting capacity. Moreover, the strong length growth enhancement may affect interactions with other mosses and vascular plants. Because winter snow addition enhanced the production of S. fuscum without affecting its structure, it may increase the carbon balance of northern peatlands.

Climate change can alter peatland plant community composition by promoting the growth of vascular plants. How such vegetation change affects peatland carbon dynamics remains, however, unclear. In order to assess the effect of vegetation change on carbon uptake and release, we performed a vascular plant-removal experiment in two Sphagnum-dominated peatlands that represent contrasting stages of natural vegetation succession along a climatic gradient. Periodic measurements of net ecosystem CO₂ exchange revealed that vascular plants play a crucial role in assuring the potential for net carbon uptake, particularly with a warmer climate. The presence of vascular plants, however, also increased ecosystem respiration, and by using the seasonal variation of respired CO₂ radiocarbon (bomb-¹⁴C) signature we demonstrate an enhanced heterotrophic decomposition of peat carbon due to rhizosphere priming. The observed rhizosphere priming of peat carbon decomposition was matched by more advanced humification of dissolved organic matter, which remained apparent beyond the plant growing season. Our results underline the relevance of rhizosphere priming in peatlands, especially when assessing the future carbon sink function of peatlands undergoing a shift in vegetation community composition in association with climate change.

Warming occurs in the Arctic twice as fast as the global average, which in turn leads to a large enhancement in terpenoid emissions from vegetation. Volatile terpenoids are the main class of biogenic volatile organic compounds (VOCs) that play crucial roles in atmospheric chemistry and climate. However, the biochemical mechanisms behind the temperature‐dependent increase in VOC emissions from subarctic ecosystems are largely unexplored. Using 13CO2‐labeling, we studied the origin of VOCs and the carbon (C) allocation under global warming in the soil–plant–atmosphere system of contrasting subarctic heath tundra vegetation communities characterized by dwarf shrubs of the genera Salix or Betula. The projected temperature rise of the subarctic summer by 5°C was realistically simulated in sophisticated climate chambers. VOC emissions strongly depended on the plant species composition of the heath tundra. Warming caused increased VOC emissions and significant changes in the pattern of volatiles toward more reactive hydrocarbons. The 13C was incorporated to varying degrees in different monoterpene and sesquiterpene isomers. We found that de novo monoterpene biosynthesis contributed to 40%–44% (Salix) and 60%–68% (Betula) of total monoterpene emissions under the current climate, and that warming increased the contribution to 50%–58% (Salix) and 87%–95% (Betula). Analyses of above‐ and belowground 12/13C showed shifts of C allocation in the plant–soil systems and negative effects of warming on C sequestration by lowering net ecosystem exchange of CO2 and increasing C loss as VOCs. This comprehensive analysis provides the scientific basis for mechanistically understanding the processes controlling terpenoid emissions, required for modeling VOC emissions from terrestrial ecosystems and predicting the future chemistry of the arctic atmosphere. By changing the chemical composition and loads of VOCs into the atmosphere, the current data indicate that global warming in the Arctic may have implications for regional and global climate and for the delicate tundra ecosystems.

Arctic ecosystems are important in the context of climate change because they are expected to undergo the most rapid temperature increases, and could provide a globally significant release of CO2 to the atmosphere from their extensive bulk soil organic carbon reserves. Understanding the relative contributions of bulk soil organic matter and plant-associated carbon pools to ecosystem respiration is critical to predicting the response of arctic ecosystem net carbon balance to climate change. In this study, we determined the variation in ecosystem respiration rates from birch forest understory and heath tundra vegetation types in northern Sweden through a full annual cycle. We used a plant biomass removal treatment to differentiate bulk soil organic matter respiration from total ecosystem respiration in each vegetation type. Plant-associated and bulk soil organic matter carbon pools each contributed significantly to ecosystem respiration during most phases of winter and summer in the two vegetation types. Ecosystem respiration rates through the year did not differ significantly between vegetation types despite substantial differences in biomass pools, soil depth and temperature regime. Most (76-92%) of the intra-annual variation in ecosystem respiration rates from these two common mesic subarctic ecosystems was explained using a first-order exponential equation relating respiration to substrate chemical quality and soil temperature. Removal of plants and their current year’s litter significantly reduced the sensitivity of ecosystem respiration to intra-annual variations in soil temperature for both vegetation types, indicating that respiration derived from recent plant carbon fixation was more temperature sensitive than respiration from bulk soil organic matter carbon stores. Accurate assessment of the potential for positive feedbacks from high-latitude ecosystems to CO2-induced climate change will require the development of ecosystem-level physiological models of net carbon exchange that differentiate the responses of major C pools, that account for effects of vegetation type, and that integrate over summer and winter seasons.

General circulation models consistently predict that regional warming will be most rapid in the Arctic, that this warming will be predominantly in the winter season, and that it will often be accompanied by increasing snowfall. Paradoxically, despite the strong cold season emphasis in these predictions, we know relatively little about the plot and landscape-level controls on tundra biogeochemical cycling in wintertime as compared to summertime. We investigated the relative influence of vegetation type and climate on CO2 production rates and total wintertime CO2 release in the Scandinavian subarctic. Ecosystem respiration rates and a wide range of associated environmental and substrate pool size variables were measured in the two most common vegetation types of the region (birch understorey and heath tundra) at four paired sites along a 50 km transect through a strong snow depth gradient in northern Sweden. Both climate and vegetation type were strong interactive controls on ecosystem CO2 production rates during winter. Of all variables tested, soil temperature explained by far the largest amount of variation in respiration rates (41-75%). Our results indicate that vegetation type only exerted an influence on respiration when snow depth was below a certain threshold (similar to 1 m). Thus, tall vegetation that enhanced snow accumulation within that threshold resulted in more effective thermal insulation from severe air temperatures, thereby significantly increasing respiratory activity. At the end of winter, within several days of snowmelt, gross ecosystem photosynthesis rates were of a similar magnitude to ecosystem respiration, resulting in significant net carbon gain in some instances. Finally, climate and vegetation type were also strong interactive controls on total wintertime respiration, suggesting that spatial variations in maximum snowdepth may be a primary determinant of regional patterns of wintertime CO2 release. Together, our results have important implications for predictions of how the distribution of tundra vegetation types and the carbon balances of arctic ecosystems will respond to climate change during winter because they indicate a threshold (similar to 1 m) above which there would be little effect of increased snow accumulation on wintertime biogeochemical cycling.

Soil carbon in permafrost ecosystems has the potential to become a major positive feedback to climate change if permafrost thaw increases heterotrophic decomposition. However, warming can also stimulate autotrophic production leading to increased ecosystem carbon storage—a negative climate change feedback. Few studies partitioning ecosystem respiration examine decadal warming effects or compare responses among ecosystems. Here, we first examined how 11 years of warming during different seasons affected autotrophic and heterotrophic respiration in a bryophyte-dominated peatland in Abisko, Sweden. We used natural abundance radiocarbon to partition ecosystem respiration into autotrophic respiration, associated with production, and heterotrophic decomposition. Summertime warming decreased the age of carbon respired by the ecosystem due to increased proportional contributions from autotrophic and young soil respiration and decreased proportional contributions from old soil. Summertime warming’s large effect was due to not only warmer air temperatures during the growing season, but also to warmer deep soils year-round. Second, we compared ecosystem respiration responses between two contrasting ecosystems, the Abisko peatland and a tussock-dominated tundra in Healy, Alaska. Each ecosystem had two different timescales of warming (<5 years and over a decade). Despite the Abisko peatland having greater ecosystem respiration and larger contributions from heterotrophic respiration than the Healy tundra, both systems responded consistently to short- and long-term warming with increased respiration, increased autotrophic contributions to ecosystem respiration, and increased ratios of autotrophic to heterotrophic respiration. We did not detect an increase in old soil carbon losses with warming at either site. If increased autotrophic respiration is balanced by increased primary production, as is the case in the Healy tundra, warming will not cause these ecosystems to become growing season carbon sources. Warming instead causes a persistent shift from heterotrophic to more autotrophic control of the growing season carbon cycle in these carbon-rich permafrost ecosystems.

Permafrost thaw is a major potential feedback source to climate change as it can drive the increased release of greenhouse gases carbon dioxide (CO2) and methane (CH4). This carbon release from the decomposition of thawing soil organic material can be mitigated by increased net primary productivity (NPP) caused by warming, increasing atmospheric CO2, and plant community transition. However, the net effect on C storage also depends on how these plant community changes alter plant litter quantity, quality, and decomposition rates. Predicting decomposition rates based on litter quality remains challenging, but a promising new way forward is to incorporate measures of the energetic favorability to soil microbes of plant biomass decomposition. We asked how the variation in one such measure, the nominal oxidation state of carbon (NOSC), interacts with changing quantities of plant material inputs to influence the net C balance of a thawing permafrost peatland. We found: (1) Plant productivity (NPP) increased post-thaw, but instead of contributing to increased standing biomass, it increased plant biomass turnover via increased litter inputs to soil; (2) Plant litter thermodynamic favorability (NOSC) and decomposition rate both increased post-thaw, despite limited changes in bulk C:N ratios; (3) these increases caused the higher NPP to cycle more rapidly through both plants and soil, contributing to higher CO2 and CH4 fluxes from decomposition. Thus, the increased C-storage expected from higher productivity was limited and the high global warming potential of CH4 contributed a net positive warming effect. Although post-thaw peatlands are currently C sinks due to high NPP offsetting high CO2 release, this status is very sensitive to the plant community's litter input rate and quality. Integration of novel bioavailability metrics based on litter chemistry, including NOSC, into studies of ecosystem dynamics, is needed to improve the understanding of controls on arctic C stocks under continued ecosystem transition.

Thawing permafrost in the sub-Arctic has implications for the physical stability and biological dynamics of peatland ecosystems. This study provides an analysis of how permafrost thawing and subsequent vegetation changes in a sub-Arctic Swedish mire have changed the net exchange of greenhouse gases, carbon dioxide (CO₂) and CH₄ over the past three decades. Images of the mire (ca. 17 ha) and surroundings taken with film sensitive in the visible and the near infrared portion of the spectrum, [i.e. colour infrared (CIR) aerial photographs from 1970 and 2000] were used. The results show that during this period the area covered by hummock vegetation decreased by more than 11% and became replaced by wet-growing plant communities. The overall net uptake of C in the vegetation and the release of C by heterotrophic respiration might have increased resulting in increases in both the growing season atmospheric CO₂ sink function with about 16% and the CH₄ emissions with 22%. Calculating the flux as CO₂ equivalents show that the mire in 2000 has a 47% greater radiative forcing on the atmosphere using a 100-year time horizon. Northern peatlands in areas with thawing sporadic or discontinuous permafrost are likely to act as larger greenhouse gas sources over the growing season today than a few decades ago because of increased CH₄ emissions.

Climate change is predicted to be dramatic at high latitudes. Still, climate impact on high latitude lake ecosystems is poorly understood. We studied 15 subarctic lakes located in a climate gradient comprising an air temperature difference of about 6&DEG; C. We show that lake water productivity varied by one order of magnitude along the temperature gradient. This variation was mainly caused by variations in the length of the ice-free period and, more importantly, in the supply of organic carbon and inorganic nutrients, which followed differences in terrestrial vegetation cover along the gradient. The results imply that warming will have rapid effects on the productivity of high latitude lakes, by prolongation of ice-free periods. However, a more pronounced consequence will be a delayed stimulation of the productivity following upon changes of the lakes terrestrial surroundings and subsequent increasing input of elements that stimulate the production of lake biota.

Abstract Plant traits – the morphological, anatomical, physiological, biochemical and phenological characteristics of plants and their organs – determine how primary producers respond to environmental factors, affect other trophic levels, influence ecosystem processes and services and provide a link from species richness to ecosystem functional diversity. Trait data thus represent the raw material for a wide range of research from evolutionary biology, community and functional ecology to biogeography. Here we present the global database initiative named TRY, which has united a wide range of the plant trait research community worldwide and gained an unprecedented buy-in of trait data: so far 93 trait databases have been contributed. The data repository currently contains almost three million trait entries for 69 €ƒ000 out of the world’s 300 €ƒ000 plant species, with a focus on 52 groups of traits characterizing the vegetative and regeneration stages of the plant life cycle, including growth, dispersal, establishment and persistence. A first data analysis shows that most plant traits are approximately log-normally distributed, with widely differing ranges of variation across traits. Most trait variation is between species (interspecific), but significant intraspecific variation is also documented, up to 40% of the overall variation. Plant functional types (PFTs), as commonly used in vegetation models, capture a substantial fraction of the observed variation – but for several traits most variation occurs within PFTs, up to 75% of the overall variation. In the context of vegetation models these traits would better be represented by state variables rather than fixed parameter values. The improved availability of plant trait data in the unified global database is expected to support a paradigm shift from species to trait-based ecology, offer new opportunities for synthetic plant trait research and enable a more realistic and empirically grounded representation of terrestrial vegetation in Earth system models.

Climate warming increases nitrogen (N) mineralization in superficial soil layers (the dominant rooting zone) of subarctic peatlands. Thawing and subsequent mineralization of permafrost increases plant-available N around the thaw-front. Because plant production in these peatlands is N-limited, such changes may substantially affect net primary production and species composition. We aimed to identify the potential impact of increased N-availability due to permafrost thawing on subarctic peatland plant production and species performance, relative to the impact of increased N-availability in superficial organic layers. Therefore, we investigated whether plant roots are present at the thaw-front (45 cm depth) and whether N-uptake (15N-tracer) at the thaw-front occurs during maximum thaw-depth, coinciding with the end of the growing season. Moreover, we performed a unique 3-year belowground fertilization experiment with fully factorial combinations of deep- (thaw-front) and shallow-fertilization (10 cm depth) and controls. We found that certain species are present with roots at the thaw-front (Rubus chamaemorus) and have the capacity (R. chamaemorus, Eriophorum vaginatum) for N-uptake from the thaw-front between autumn and spring when aboveground tissue is largely senescent. In response to 3-year shallow-belowground fertilization (S) both shallow- (Empetrum hermaphroditum) and deep-rooting species increased aboveground biomass and N-content, but only deep-rooting species responded positively to enhanced nutrient supply at the thaw-front (D). Moreover, the effects of shallow-fertilization and thaw-front fertilization on aboveground biomass production of the deep-rooting species were similar in magnitude (S: 71%; D: 111% increase compared to control) and additive (S + D: 181% increase). Our results show that plant-available N released from thawing permafrost can form a thus far overlooked additional N-source for deep-rooting subarctic plant species and increase their biomass production beyond the already established impact of warming-driven enhanced shallow N-mineralization. This may result in shifts in plant community composition and may partially counteract the increased carbon losses from thawing permafrost.

Many of the world's northern peatlands are underlain by rapidly thawing permafrost. Because plant production in these peatlands is often nitrogen (N)‐limited, a release of N stored in permafrost may stimulate net primary production or change species composition if it is plant‐available. In this study, we aimed to quantify plant‐available N in thawing permafrost soils of subarctic peatlands. We compared plant‐available N‐pools and ‐fluxes in near‐surface permafrost (0–10 cm below the thawfront) to those taken from a current rooting zone layer (5–15 cm depth) across five representative peatlands in subarctic Sweden. A range of complementary methods was used: extractions of inorganic and organic N, inorganic and organic N‐release measurements at 0.5 and 11 °C (over 120 days, relevant to different thaw‐development scenarios) and a bioassay with Poa alpina test plants. All extraction methods, across all peatlands, consistently showed up to seven times more plant‐available N in near‐surface permafrost soil compared to the current rooting zone layer. These results were supported by the bioassay experiment, with an eightfold larger plant N‐uptake from permafrost soil than from other N‐sources such as current rooting zone soil or fresh litter substrates. Moreover, net mineralization rates were much higher in permafrost soils compared to soils from the current rooting zone layer (273 mg N m⁻² and 1348 mg N m⁻² per growing season for near‐surface permafrost at 0.5 °C and 11 °C respectively, compared to −30 mg N m⁻² for current rooting zone soil at 11 °C). Hence, our results demonstrate that near‐surface permafrost soil of subarctic peatlands can release a biologically relevant amount of plant available nitrogen, both directly upon thawing as well as over the course of a growing season through continued microbial mineralization of organically bound N. Given the nitrogen‐limited nature of northern peatlands, this release may have impacts on both plant productivity and species composition.

In Arctic regions, thawing permafrost soils are projected to release 50 to 250 Gt of carbon by 2100. This data is mostly derived from carbon-rich wetlands, although 71% of this carbon pool is stored in faster-thawing mineral soils, where ecosystems close to the outer boundaries of permafrost regions are especially vulnerable. Although extensive data exists from currently thawing sites and short-term thawing experiments, investigations of the long-term changes following final thaw and co-occurring drainage are scarce. Here we show ecosystem changes at two comparable tussock tundra sites with distinct permafrost thaw histories, representing 15 and 25 years of natural drainage, that resulted in a 10-fold decrease in CH4 emissions (3.2 ± 2.2 vs. 0.3 ± 0.4 mg C-CH4 m−2 day−1), while CO2 emissions were comparable. These data extend the time perspective from earlier studies based on short-term experimental drainage. The overall microbial community structures did not differ significantly between sites, although the drier top soils at the most advanced site led to a loss of methanogens and their syntrophic partners in surface layers while the abundance of methanotrophs remained unchanged. The resulting deeper aeration zones likely increased CH4 oxidation due to the longer residence time of CH4 in the oxidation zone, while the observed loss of aerenchyma plants reduced CH4 diffusion from deeper soil layers directly to the atmosphere. Our findings highlight the importance of including hydrological, vegetation and microbial specific responses when studying long-term effects of climate change on CH4 emissions and underscores the need for data from different soil types and thaw histories.

Oxygen depletion in lake bottom waters has adverse impacts on ecosystem health including decreased water quality from release of nutrients and reduced substances from sediments, and the reduction of fish growth and reproduction. Depletion occurs when oxygen is consumed during decomposition of organic matter, and oxygen replenishment is limited by water column stratification. Arctic–alpine lakes are often well mixed and oxygenated, but rapid climate change in these regions is an important driver of shifts in catchment vegetation that could affect the mixing and oxygen dynamics of lakes. Here, we analyze high-resolution time series of dissolved oxygen concentration and temperature profiles in 40 Swedish arctic–alpine lakes across the tree line ecotone. The lakes stratified for 1−125 days, and during stratification, near-bottom dissolved oxygen concentrations changed by −0.20 to +0.15 mg L−1 day−1, resulting in final concentrations of 1.1−15.5 mg L−1 at the end of the longest stratification period. Structural equation modeling revealed that lakes with taller shoreline vegetation relative to lake area had higher dissolved organic carbon concentrations and oxygen consumption rates, but also lower wind speeds and longer stratification periods, and ultimately, lower near-bottom dissolved oxygen concentrations. We use an index of shoreline canopy height and lake area to predict variations among our study lakes in near-bottom dissolved oxygen concentrations at the end of the longest stratification period (R2 = 0.41). Upscaling this relationship to 8392 Swedish arctic–alpine lakes revealed that near-bottom dissolved oxygen concentrations drop below 3, 5, and 7 mg L−1 in 15%, 32%, and 53% of the lakes and that this proportion is sensitive (5%−22%, 13%−45%, and 29%−69%) to hypothetical tree line shifts observed in the past century or reconstructed for the Holocene (±200 m elevation; ±0.5° latitude). Assuming space-for-time substitution, we predict that tree line advance will decrease near-bottom dissolved oxygen concentrations in many arctic–alpine lakes.

Climate change–driven permafrost thaw has a strong influence on pan-Arctic regions, via, for example, the formation of thermokarst ponds. These ponds are hotspots of microbial carbon cycling and greenhouse gas production, and efforts have been put on disentangling the role of bacteria and archaea in recycling the increasing amounts of carbon arriving to the ponds from degrading watersheds. However, despite the well-established role of fungi in carbon cycling in the terrestrial environments, the interactions between permafrost thaw and fungal communities in Arctic freshwaters have remained unknown. We integrated data from 60 ponds in Arctic hydro-ecosystems, representing a gradient of permafrost integrity and spanning over five regions, namely Alaska, Greenland, Canada, Sweden, and Western Siberia. The results revealed that differences in pH and organic matter quality and availability were linked to distinct fungal community compositions and that a large fraction of the community represented unknown fungal phyla. Results display a 16%–19% decrease in fungal diversity, assessed by beta diversity, across ponds in landscapes with more degraded permafrost. At the same time, sites with similar carbon quality shared more species, aligning a shift in species composition with the quality and availability of terrestrial dissolved organic matter. We demonstrate that the degradation of permafrost has a strong negative impact on aquatic fungal diversity, likely via interactions with the carbon pool released from ancient deposits. This is expected to have implications for carbon cycling and climate feedback loops in the rapidly warming Arctic.

Recent accelerated decay of discontinuous permafrost at the Stordalen Mire in northern Sweden has been attributed to increased temperature and snow depth, and has caused expansion of wet minerotrophic areas leading to significant changes in carbon cycling in the mire. In order to track these changes through time and evaluate potential forcing mechanisms, this paper analyses a peat succession and a lake sediment sequence from within the mire, providing a record for the last 100 years, and compares these with monitored climate and active layer thickness data. The peat core was analysed for testate amoebae to reconstruct changes in peatland surface moisture conditions and water table fluctuations. The lake sediment core was analysed by near infrared spectroscopy to infer changes in the total organic carbon (TOC) concentration of the lake‐water, and changes in δ¹³C and C, N and δ¹⁵N to track changes in the dissolved inorganic carbon (DIC) pool and the influence of diagenetic effects on sediment organic matter, respectively. Results showed that major shifts towards increased peat surface moisture and TOC concentration of the lake‐water occurred around 1980, one to two decades earlier than a temperature driven increase in active layer thickness. Comparison with monitored temperature and precipitation from a nearby climate station indicates that this change in peat surface moisture is related to June–September (JJAS) precipitation and that the increase in lake‐water TOC concentration reflects an increase in total annual precipitation. A significant depletion in ¹³C of sediment organic matter in the early 1980s probably reflects the effect of a single or a few consecutive years with anomalously high summer precipitation, resulting in elevated DIC content of the lake water, predominantly originating from increased export and subsequent respiration of organic carbon from the mire. Based on these results, it was not possible to link proxy data obtained on peat and lake‐sediment records directly to permafrost decay. Instead our data indicate that increased precipitation and anomalously high rainfall during summers had a significant impact on the mire and the adjacent lake ecosystem. We therefore propose that effects of increased precipitation should be considered when evaluating potential forcing mechanisms of recent changes in carbon cycling in the subarctic.

Little is known about the impact of changing temperature regimes on composition and diversity of cryptogam communities in the Arctic and Subarctic, despite the well‐known importance of lichens and bryophytes to the functioning and climate feedbacks of northern ecosystems. We investigated changes in diversity and abundance of lichens and bryophytes within long‐term (9–16 years) warming experiments and along natural climatic gradients, ranging from Swedish subarctic birch forest and subarctic/subalpine tundra to Alaskan arctic tussock tundra. In both Sweden and Alaska, lichen diversity responded negatively to experimental warming (with the exception of a birch forest) and to higher temperatures along climatic gradients. Bryophytes were less sensitive to experimental warming than lichens, but depending on the length of the gradient, bryophyte diversity decreased both with increasing temperatures and at extremely low temperatures. Among bryophytes, Sphagnum mosses were particularly resistant to experimental warming in terms of both abundance and diversity. Temperature, on both continents, was the main driver of species composition within experiments and along gradients, with the exception of the Swedish subarctic birch forest where amount of litter constituted the best explanatory variable. In a warming experiment in moist acidic tussock tundra in Alaska, temperature together with soil ammonium availability were the most important factors influencing species composition. Overall, dwarf shrub abundance (deciduous and evergreen) was positively related to warming but so were the bryophytes Sphagnum girgensohnii, Hylocomium splendens and Pleurozium schreberi; the majority of other cryptogams showed a negative relationship to warming. This unique combination of intercontinental comparison, natural gradient studies and experimental studies shows that cryptogam diversity and abundance, especially within lichens, is likely to decrease under arctic climate warming. Given the many ecosystem processes affected by cryptogams in high latitudes (e.g. carbon sequestration, N₂‐fixation, trophic interactions), these changes will have important feedback consequences for ecosystem functions and climate.

While substantial cold-season respiration has been documented in most arctic and alpine ecosystems in recent years, the significance of cold-season photosynthesis in these biomes is still believed to be small. In a mesic, subartic heath during both the cold and warm season, we measured in situ ecosystem respiration and photosynthesis with a chamber technique at ambient conditions and at artificially, increased frequency of freeze-thaw (FT) cycles during fall and spring. We fitted the measured ecosystem exchange rates to respiration and photosynthesis models with R-2-values ranging from 0.81 to 0.85. As expected, estimated cold-season (October, November, April and May) respiration was significant and accounted for at least 22% of the annual respiratory CO2 flux. More surprisingly, estimated photosynthesis during this period accounted for up to 19% of the annual gross CO2 uptake, suggesting that cold-season photosynthesis partly balanced the cold-season respiratory carbon losses and can be significant for the annual cycle of carbon. Still, during the full year the ecosystem was a significant net source of 120 +/- 12g Cm-2 to the atmosphere. Neither respiration nor photosynthetic rates were much affected by the extra FT cycles, although the mean rate of net ecosystem loss decreased slightly, but significantly, in May. The results suggest only a small response of net carbon fluxes to increased frequency of FT cycles in this ecosystem.

Global environmental changes are causing widespread nutrient depletion, declines in the ratio of dissolved inorganic nitrogen (N) to total phosphorus (DIN:TP), and increases in both water temperature and terrestrial colored dissolved organic carbon (DOC) concentration (browning) in high-latitude northern lakes. Declining lake DIN:TP, warming, and browning alter the nutrient limitation regime and biomass of phytoplankton, but how these stressors together affect the nutritional quality in terms of polyunsaturated fatty acid (PUFA) contents of the pelagic food web components remains unknown. We assessed the fatty acid compositions of seston and zooplankton in 33 lakes across south-to-north and boreal-to-subarctic gradients in Sweden. Data showed higher lake DIN:TP in the south than in the north, and that boreal lakes were warmer and browner than subarctic lakes. Lake DIN:TP strongly affected the PUFA contents—especially eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA)—in seston, calanoids, and copepods (as a group), but not in cladocerans. The EPA+DHA contents increased by 123% in seston, 197% in calanoids, and 230% in copepods across a lake molar DIN:TP gradient from 0.17 to 14.53, indicating lower seston and copepod nutritional quality in the more N-limited lakes (those with lower DIN:TP). Water temperature affected EPA+DHA contents of zooplankton, especially cladocerans, but not seston. Cladoceran EPA+DHA contents were reduced by ca. 6% for every 1°C increase in surface water. Also, the EPA, DHA, or EPA+DHA contents of Bosmina, cyclopoids, and copepods increased in lakes with higher DOC concentrations or aromaticity. Our findings indicate that zooplankton food quality for higher consumers will decrease with warming alone (for cladocerans) or in combination with declining lake DIN:TP (for copepods), but impacts of these stressors are moderated by lake browning. Global environmental changes that drive northern lakes toward more N-limited, warmer, and browner conditions will reduce PUFA availability and nutritional quality of the pelagic food web components.

Current analyses and predictions of spatially explicit patterns and processes in ecology most often rely on climate data interpolated from standardized weather stations. This interpolated climate data represents long‐term average thermal conditions at coarse spatial resolutions only. Hence, many climate‐forcing factors that operate at fine spatiotemporal resolutions are overlooked. This is particularly important in relation to effects of observation height (e.g. vegetation, snow and soil characteristics) and in habitats varying in their exposure to radiation, moisture and wind (e.g. topography, radiative forcing or cold‐air pooling). Since organisms living close to the ground relate more strongly to these microclimatic conditions than to free‐air temperatures, microclimatic ground and near‐surface data are needed to provide realistic forecasts of the fate of such organisms under anthropogenic climate change, as well as of the functioning of the ecosystems they live in. To fill this critical gap, we highlight a call for temperature time series submissions to SoilTemp, a geospatial database initiative compiling soil and near‐surface temperature data from all over the world. Currently, this database contains time series from 7,538 temperature sensors from 51 countries across all key biomes. The database will pave the way toward an improved global understanding of microclimate and bridge the gap between the available climate data and the climate at fine spatiotemporal resolutions relevant to most organisms and ecosystem processes.

Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1-km² resolution for 0?5 and 5?15 cm soil depth. These maps were created by calculating the difference (i.e., offset) between in-situ soil temperature measurements, based on time series from over 1200 1-km² pixels (summarized from 8500 unique temperature sensors) across all the world?s major terrestrial biomes, and coarse-grained air temperature estimates from ERA5-Land (an atmospheric reanalysis by the European Centre for Medium-Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (-0.7 ± 2.3°C). The observed substantial and biome-specific offsets emphasize that the projected impacts of climate and climate change on near-surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil-related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in-situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.

Recent studies from mountainous areas of small spatial extent (<2500km2) suggest that fine-grained thermal variability over tens or hundreds of metres exceeds much of the climate warming expected for the coming decades. Such variability in temperature provides buffering to mitigate climate-change impacts. Is this local spatial buffering restricted to topographically complex terrains? To answer this, we here study fine-grained thermal variability across a 2500-km wide latitudinal gradient in Northern Europe encompassing a large array of topographic complexities. We first combined plant community data, Ellenberg temperature indicator values, locally measured temperatures (LmT) and globally interpolated temperatures (GiT) in a modelling framework to infer biologically relevant temperature conditions from plant assemblages within <1000-m2 units (community-inferred temperatures: CiT). We then assessed: (1) CiT range (thermal variability) within 1-km2 units; (2) the relationship between CiT range and topographically and geographically derived predictors at 1-km resolution; and (3) whether spatial turnover in CiT is greater than spatial turnover in GiT within 100-km2 units. Ellenberg temperature indicator values in combination with plant assemblages explained 4672% of variation in LmT and 9296% of variation in GiT during the growing season (June, July, August). Growing-season CiT range within 1-km2 units peaked at 6065 degrees N and increased with terrain roughness, averaging 1.97 degrees C (SD=0.84 degrees C) and 2.68 degrees C (SD=1.26 degrees C) within the flattest and roughest units respectively. Complex interactions between topography-related variables and latitude explained 35% of variation in growing-season CiT range when accounting for sampling effort and residual spatial autocorrelation. Spatial turnover in growing-season CiT within 100-km2 units was, on average, 1.8 times greater (0.32 degrees Ckm1) than spatial turnover in growing-season GiT (0.18 degrees Ckm1). We conclude that thermal variability within 1-km2 units strongly increases local spatial buffering of future climate warming across Northern Europe, even in the flattest terrains.

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum , strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.

Climate change is predicted to cause milder winters and thus exacerbate soil freeze?thaw perturbations in the subarctic, recasting the environmental challenges that soil microorganisms need to endure. Historical exposure to environmental stressors can facilitate the microbial resilience to new cycles of that same stress. However, whether and how such microbial memory or stress legacy can modulate microbial responses to cycles of frost remains untested. Here, we conducted an in situ field experiment in a subarctic birch forest, where winter warming resulted in a substantial increase in the number and intensity of freeze?thaw events. After one season of winter warming, which raised mean surface and soil (?8?cm) temperatures by 2.9 and 1.4°C, respectively, we investigated whether the in situ warming-induced increase in frost cycles improved soil microbial resilience to an experimental freeze?thaw perturbation. We found that the resilience of microbial growth was enhanced in the winter warmed soil, which was associated with community differences across treatments. We also found that winter warming enhanced the resilience of bacteria more than fungi. In contrast, the respiration response to freeze?thaw was not affected by a legacy of winter warming. This translated into an enhanced microbial carbon-use efficiency in the winter warming treatments, which could promote the stabilization of soil carbon during such perturbations. Together, these findings highlight the importance of climate history in shaping current and future dynamics of soil microbial functioning to perturbations associated with climate change, with important implications for understanding the potential consequences on microbial-mediated biogeochemical cycles.

Arctic plants are adapted to climatic variability, but their long-term responses to warming remain unclear. Responses may occur by range shifts, phenological adjustments in growth and reproduction, or both. Here, we compare distribution and phenology of 83 arctic and boreal mountain species, sampled identically in the early 20th (1917–1919) and 21st centuries (2017–2018) from a region of northern Sweden that has warmed significantly. We test two compensatory hypotheses to high-latitude warming—upward shifts in distribution, and earlier or extended growth and reproduction. For distribution, we show dramatic upward migration by 69% of species, averaging 6.1 m per decade, especially boreal woodland taxa whose upward expansion has reduced arctic montane habitat by 30%. Twenty percent of summit species showed distributional shifts but downward, especially moisture-associated snowbed flora. For phenology, we detected wide inter-annual variability in the onset of leafing and flowering in both eras. However, there was no detectable change in growing-season length, relating to two mechanisms. First, plot-level snow melt data starting in 1917 demonstrated that melt date, rather than vernal temperatures, better predicts plant emergence, with snow melt influenced by warmer years having greater snowfall—warmer springs did not always result in earlier emergence because snowbeds can persist longer. Second, the onset of reproductive senescence between eras was similar, even when plant emergence was earlier by a month, possibly due to intensified summer heat stress or hard-wired ‘canalization’ where senescence occurs regardless of summer temperature. Migrations in this system have possibly buffered arctic species against displacement by boreal expansion and warming, but ongoing temperature increases, woody plant invasion, and a potential lack of flexibility in timing of senescence may foreshadow challenges.

This study deals with changes in the plant cover and its net carbon sequestration over 30 years on a subarctic Sphagnum-mire with permafrost near Abisko, northernmost Sweden, in relation to climatic variations during the same period. Aerial colour infrared images from 1970 and 2000 were compared to reveal changes in surface structure and vegetation over the whole mire, while the plant populations were studied within a smaller, mainly ombrotrophic part. The results demonstrated two processes, namely (1) that wet sites dominated by graminoids expanded while hummock sites dominated by dwarf shrubs receded, and (2) that on the hummocks lichens expanded while evergreen dwarf shrubs and mosses decreased, both processes creating an instability in the surface structure. A successive degradation of the permafrost is the likely reason for the increase in wet areas, while the changes in the hummock vegetation might have resulted from higher spring temperatures giving rise to an intensified snow melt, exposing the vegetation to frost drought. Because of the vegetation changes, the annual litter input of carbon to the mire has increased slightly, by 4 g m(-2) a(-1) (7.3%), over these years while an increased erosion has resulted in a loss of 40-80 Mg carbon or 7-17 g m(-2) a(-1) for the entire mire over the same period. As the recalcitrant proportion of the litter has decreased, the decay rate in the acrotelm might be expected to increase in the future.

Climate change is rapidly reshaping arctic landscapes through shifts in vegetation cover and productivity, soil resource mobilization, and hydrological regimes. The implications of these changes for stream ecosystems and food webs is unclear and will depend largely on microbial biofilm responses to concurrent shifts in temperature, light, and resource supply from land. To study those responses, we used nutrient diffusing substrates to manipulate resource supply to biofilm communities along regional gradients in stream temperature, riparian shading, and dissolved organic carbon (DOC) loading in arctic Sweden. We found strong nitrogen (N) limitation across this gradient for gross primary production, community respiration and chlorophyll-a accumulation. For unamended biofilms, activity and biomass accrual were not driven by any single physical or chemical driver across this region. However, the magnitude of biofilm response to N addition did: in tundra streams, biofilm response was constrained by thermal regimes, whereas variation in light availability regulated this response in birch and coniferous forest streams. Furthermore, heterotrophic responses to experimental N addition increased across the region with greater stream water concentrations of DOC relative to inorganic N. Thus, future shifts in resource supply to these ecosystems are likely to interact with other concurrent environmental changes to regulate stream productivity. Indeed, our results suggest that in the absence of increased nutrient inputs, arctic streams will be less sensitive to future changes in other habitat variables such as temperature and DOC loading.This article is protected by copyright. All rights reserved.

Permafrost thaw in peatlands has the potential to alter catchment export of dissolved organic carbon (DOC) and thus influence downstream aquatic C cycling. Subarctic peatlands are often mosaics of different peatland types, where permafrost conditions regulate the hydrological setting of each type. We show that hydrological setting is key to observed differences in magnitude, timing, and chemical composition of DOC export between permafrost and nonpermafrost peatland types, and that these differences influence the export of DOC of larger catchments even when peatlands are minor catchment components. In many aspects, DOC export from a studied peatland permafrost plateau was similar to that of a forested upland catchment. Similarities included low annual export (2–3 g C m−2) dominated by the snow melt period ( 70%), and how substantial DOC export following storms required wet antecedent conditions. Conversely, nonpermafrost fens had higher DOC export (7 g C m−2), resulting from sustained hydrological connectivity during summer. Chemical composition of catchment DOC export arose from the mixing of highly aromatic DOC from organic soils from permafrost plateau soil water and upland forest surface horizons with nonaromatic DOC from mineral soil groundwater, but was further modulated by fens. Increasing aromaticity from fen inflow to outlet was substantial and depended on both water residence time and water temperature. The role of fens as catchment biogeochemical hotspots was further emphasized by their capacity for sulfate retention. As a result of fen characteristics, a 4% fen cover in a mixed catchment was responsible for 34% higher DOC export, 50% higher DOC concentrations and 10% higher DOC aromaticity at the catchment outlet during summer compared to a nonpeatland upland catchment. Expansion of fens due to thaw thus has potential to influence landscape C cycling by increasing fen capacity to act as biogeochemical hotspots, amplifying aquatic C cycling, and increasing catchment DOC export.

Methane (CH4) emissions from the northern high-latitude region represent potentially significant biogeochemical feedbacks to the climate system. We compiled a database of growing-season CH4 emissions from terrestrial ecosystems located across permafrost zones, including 303 sites described in 65 studies. Data on environmental and physical variables, including permafrost conditions, were used to assess controls on CH4 emissions. Water table position, soil temperature, and vegetation composition strongly influenced emissions and had interacting effects. Sites with a dense sedge cover had higher emissions than other sites at comparable water table positions, and this was an effect that was more pronounced at low soil temperatures. Sensitivity analysis suggested that CH4 emissions from ecosystems where the water table on average is at or above the soil surface (wet tundra, fen underlain by permafrost, and littoral ecosystems) are more sensitive to variability in soil temperature than drier ecosystems (palsa dry tundra, bog, and fen), whereas the latter ecosystems conversely are relatively more sensitive to changes of the water table position. Sites with near-surface permafrost had lower CH4 fluxes than sites without permafrost at comparable water table positions, a difference that was explained by lower soil temperatures. Neither the active layer depth nor the organic soil layer depth was related to CH4 emissions. Permafrost thaw in lowland regions is often associated with increased soil moisture, higher soil temperatures, and increased sedge cover. In our database, lowland thermokarst sites generally had higher emissions than adjacent sites with intact permafrost, but emissions from thermokarst sites were not statistically higher than emissions from permafrost-free sites with comparable environmental conditions. Overall, these results suggest that future changes to terrestrial high-latitude CH4 emissions will be more proximately related to changes in moisture, soil temperature, and vegetation composition than to increased availability of organic matter following permafrost thaw.

Thicker snowpacks and their insulation effects cause winter-warming and invoke thaw of permafrost ecosystems. Temperature-dependent decomposition of previously frozen carbon (C) is currently considered one of the strongest feedbacks between the Arctic and the climate system, but the direction and magnitude of the net C balance remains uncertain. This is because winter effects are rarely integrated with C fluxes during the snow-free season and because predicting the net C balance from both surface processes and thawing deep layers remains challenging. In this study, we quantified changes in the long-term net C balance (net ecosystem production) in a subarctic peat plateau subjected to 10 years of experimental winter-warming. By combining(210)Pb and(14)Cdating of peat cores with peat growth models, we investigated thawing effects on year-round primary production and C losses through respiration and leaching from both shallow and deep peat layers. Winter-warming and permafrost thaw had no effect on the net C balance, but strongly affected gross C fluxes. Carbon losses through decomposition from the upper peat were reduced as thawing of permafrost induced surface subsidence and subsequent waterlogging. However, primary production was also reduced likely due to a strong decline in bryophytes cover while losses from the old C pool almost tripled, caused by the deepened active layer. Our findings highlight the need to estimate long-term responses of whole-year production and decomposition processes to thawing, both in shallow and deep soil layers, as they may contrast and lead to unexpected net effects on permafrost C storage.

Responses of the mycorrhizal fungal community in terrestrial ecosystems to global change factors are not well understood. However, virtually all land plants form symbiotic associations with mycorrhizal fungi, with approximately 20% of the plants' net primary production transported down to the fungal symbionts. In this study, we investigated how ericoid mycorrhiza (ErM), fine endophytes (FE) and dark septate endophytes (DSE) in roots responded to elevated atmospheric CO2 concentrations and warming in the dwarf shrub understory of a birch forest in the subarctic region of northern Sweden. To place the belowground results into an ecosystem context we also investigated how plant cover and nutrient concentrations in leaves responded to elevated atmospheric CO2 concentrations and warming. The ErM colonization in ericaceous dwarf shrubs increased under elevated atmospheric CO2 concentrations, but did not respond to warming following 6 years of treatment. This suggests that the higher ErM colonization under elevated CO2 might be due to increased transport of carbon belowground to acquire limiting resources such as N, which was diluted in leaves of ericaceous plants under enhanced CO2. The elevated CO2 did not affect total plant cover but the plant cover was increased under warming, which might be due to increased N availability in soil. FE colonization in grass roots decreased under enhanced CO2 and under warming, which might be due to increased root growth, to which the FE fungi could not keep up, resulting in proportionally lower colonization. However, no responses in aboveground cover of Deschampsia flexuosa were seen. DSE hyphal colonization in grass roots significantly increased under warmer conditions, but did not respond to elevated CO2. This complex set of responses by mycorrhizal and other root-associated fungi to global change factors of all the fungal types studied could have broad implications for plant community structure and biogeochemistry of subarctic ecosystems.

Climate warming at high northern latitudes has caused substantial increases in plant productivity of tundra vegetation and an expansion of the range of deciduous shrub species. However significant the increase in carbon (C) contained within above-ground shrub biomass, it is modest in comparison with the amount of C stored in the soil in tundra ecosystems. Here, we use a ‘space-for-time’ approach to test the hypothesis that a shift from lower-productivity tundra heath to higher-productivity deciduous shrub vegetation in the sub-Arctic may lead to a loss of soil C that out-weighs the increase in above-ground shrub biomass. We further hypothesize that a shift from ericoid to ectomycorrhizal systems coincident with this vegetation change provides a mechanism for the loss of soil C. We sampled soil C stocks, soil surface CO2 flux rates and fungal growth rates along replicated natural transitions from birch forest (Betula pubescens), through deciduous shrub tundra (Betula nana) to tundra heaths (Empetrum nigrum) near Abisko, Swedish Lapland. We demonstrate that organic horizon soil organic C (SOCorg) is significantly lower at shrub (2.98 ± 0.48 kg m−2) and forest (2.04 ± 0.25 kg m−2) plots than at heath plots (7.03 ± 0.79 kg m−2). Shrub vegetation had the highest respiration rates, suggesting that despite higher rates of C assimilation, C turnover was also very high and less C is sequestered in the ecosystem. Growth rates of fungal hyphae increased across the transition from heath to shrub, suggesting that the action of ectomycorrhizal symbionts in the scavenging of organically bound nutrients is an important pathway by which soil C is made available to microbial degradation. The expansion of deciduous shrubs onto potentially vulnerable arctic soils with large stores of C could therefore represent a significant positive feedback to the climate system.

The Continuous Plankton Recorder survey has monitored plankton in the Northwest Atlantic at monthly intervals since 1962, with an interegnum between 1978 and 1990. In May 1999, large numbers of the Pacific diatom Neodenticula seminae were found in Continuous Plankton Recorder (CPR) samples in the Labrador Sea as the first record in the North Atlantic for more than 800 000 years. The event coincided with modifications in Arctic hydrography and circulation, increased flows of Pacific water into the Northwest Atlantic and in the previous year the exceptional occurrence of extensive ice-free water to the North of Canada. These observations indicate that N. seminae was carried in a pulse of Pacific water in 1998/early 1999 via the Canadian Arctic Archipelago and/or Fram Strait. The species occurred previously in the North Atlantic during the Pleistocene from similar to 1.2 to similar to 0.8 Ma as recorded in deep sea sediment cores. The reappearance of N. seminae in the North Atlantic is an indicator of the scale and speed of changes that are taking place in the Arctic and North Atlantic oceans as a consequence of regional climate warming. Because of the unusual nature of the event it appears that a threshold has been passed, marking a change in the circulation between the North Pacific and North Atlantic Oceans via the Arctic. Trans-Arctic migrations from the Pacific into the Atlantic are likely to occur increasingly over the next 100 years as Arctic ice continues to melt affecting Atlantic biodiversity and the biological pump with consequent feedbacks to the carbon cycle.

Climate change increases the insect abundance, especially in Arctic ecosystems. Insect herbivory also significantly increases plant emissions of volatile organic compounds (VOCs), which are highly reactive in the atmosphere and play a crucial role in atmospheric chemistry and physics. However, it is unclear how the effects of insect herbivory on VOC emissions interact with climatic changes, such as warming and increased cloudiness. We assessed how experimental manipulations of temperature and light availability in subarctic tundra, that had been maintained for 30 years at the time of the measurements, affect the VOC emissions from a widespread dwarf birch (Betula nana) when subjected to herbivory by local geometrid moth larvae, the autumnal moth (Epirrita autumnata) and the winter moth (Operophtera brumata). Warming and insect herbivory on B. nana stimulated VOC emission rates and altered the VOC blend. The herbivory-induced increase in sesquiterpene and homoterpene emissions were climate-treatment-dependent. Many herbivory-associated VOCs were more strongly induced in the shading treatment than in other treatments. We showed generally enhanced tundra VOC emissions upon insect herbivory and synergistic effects on the emissions of some VOC groups in a changing climate, which can have positive feedbacks on cloud formation. Furthermore, the acclimation of plants to long-term climate treatments affects VOC emissions and strongly interacts with plant responses to herbivory. Such acclimation complicates predictions of how climate change, together with interacting biotic stresses, affects VOC emissions in the high latitudes.

Soil microbial biomass in arctic heaths has been shown to be largely unaffected by treatments simulating climate change with temperature, nutrient and light manipulations. Here, we demonstrate that more than 10 years is needed for development of significant responses, and that changes in microbial biomass are accompanied with strong alterations in microbial community composition. In contrast to slight or nonsignificant responses after 5, 6 and 10 treatment years, 15 years of inorganic NPK fertilizer addition to a subarctic heath had strong effects on the microbial community and, as observed for the first time, warming and shading also led to significant responses, often in opposite direction to the fertilization responses. The effects were clearer in the top 5 cm soil than at the 5-10 cm depth. Fertilization increased microbial biomass C and more than doubled microbial biomass P compared to the non-fertilized plots. However, it only increased microbial biomass N at the 5-10 cm depth. Fertilization increased fungal biomass and the relative abundance of phospholipid fatty acid (PLFA) markers of gram-positive bacteria. Warming and shading decreased the relative abundance of fungal PLFAs, and shading also altered the composition of the bacterial community. The long time lag in responses may be associated with indirect effects of the gradual changes in the plant biomass and community composition. The contrasting responses to warming and fertilization treatments show that results from fertilizer addition may not be similar to the effects of increased nutrient mineralization and availability following climatic warming.

Streams play an important role in the global carbon (C) cycle, accounting for a large portion of CO2 evaded from inland waters despite their small areal coverage. However, the relative importance of different terrestrial and aquatic processes driving CO2 production and evasion from streams remains poorly understood. In this study, we measured O-2 and CO2 continuously in streams draining tundra-dominated catchments in northern Sweden, during the summers of 2015 and 2016. From this, we estimated daily metabolic rates and CO2 evasion simultaneously and thus provide insight into the role of stream metabolism as a driver of C dynamics in Arctic streams. Our results show that aquatic biological processes regulate CO2 concentrations and evasion at multiple timescales. Photosynthesis caused CO2 concentrations to decrease by as much as 900 ppm during the day, with the magnitude of this diel variation being strongest at the low-turbulence streams. Diel patterns in CO2 concentrations in turn influenced evasion, with up to 45% higher rates at night. Throughout the summer, CO2 evasion was sustained by aquatic ecosystem respiration, which was one order of magnitude higher than gross primary production. Furthermore, in most cases, the contribution of stream respiration exceeded CO2 evasion, suggesting that some stream reaches serve as net sources of CO2, thus creating longitudinal heterogeneity in C production and loss within this stream network. Overall, our results provide the first link between stream metabolism and CO2 evasion in the Arctic and demonstrate that stream metabolic processes are key drivers of the transformation and fate of terrestrial organic matter exported from these landscapes.

Nitrogen (N) fixation in moss-associated cyanobacteria is one of the main sources of available N for N-limited ecosystems such as subarctic tundra. Yet, N2 fixation in mosses is strongly influenced by soil moisture and temperature. Thus, temporal scaling up of low-frequency in situ measurements to several weeks, months or even the entire growing season without taking into account changes in abiotic conditions cannot capture the variation in moss-associated N2 fixation. We therefore aimed to estimate moss-associated N2 fixation throughout the snow-free period in subarctic tundra in field experiments simulating climate change: willow (Salix myrsinifolia) and birch (Betula pubescens spp. tortuosa) litter addition, and warming. To achieve this, we established relationships between measured in situ N2 fixation rates and soil moisture and soil temperature and used high-resolution measurements of soil moisture and soil temperature (hourly from May to October) to model N2 fixation. The modelled N2 fixation rates were highest in the warmed (2.8 ± 0.3 kg N ha−1) and birch litter addition plots (2.8 ± 0.2 kg N ha−1), and lowest in the plots receiving willow litter (1.6 ± 0.2 kg N ha−1). The control plots had intermediate rates (2.2 ± 0.2 kg N ha−1). Further, N2 fixation was highest during the summer in the warmed plots, but was lowest in the litter addition plots during the same period. The temperature and moisture dependence of N2 fixation was different between the climate change treatments, indicating a shift in the N2 fixer community. Our findings, using a combined empirical and modelling approach, suggest that a longer snow-free period and increased temperatures in a future climate will likely lead to higher N2 fixation rates in mosses. Yet, the consequences of increased litter fall on moss-associated N2 fixation due to shrub expansion in the Arctic will depend on the shrub species’ litter traits.

Half the global soil carbon (C) is held in high-latitude systems. Climate change will expose these to warming and a shift towards plant communities with more labile C input. Labile C can also increase the rate of loss of native soil organic matter (SOM); a phenomenon termed ‘priming’. We investigated how warming (+1.1 °C over ambient using open top chambers) and litter addition (90 g m−2 yr−1) treatments in the subarctic influenced the susceptibility of SOM mineralization to priming, and its microbial underpinnings. Labile C appeared to inhibit the mineralization of C from SOM by up to 60% within hours. In contrast, the mineralization of N from SOM was stimulated by up to 300%. These responses occurred rapidly and were unrelated to microbial successional dynamics, suggesting catabolic responses. Considered separately, the labile C inhibited C mineralization is compatible with previously reported findings termed ‘preferential substrate utilization’ or ‘negative apparent priming’, while the stimulated N mineralization responses echo recent reports of ‘real priming’ of SOM mineralization. However, C and N mineralization responses derived from the same SOM source must be interpreted together: This suggested that the microbial SOM-use decreased in magnitude and shifted to components richer in N. This finding highlights that only considering SOM in terms of C may be simplistic, and will not capture all changes in SOM decomposition. The selective mining for N increased in climate change treatments with higher fungal dominance. In conclusion, labile C appeared to trigger catabolic responses of the resident microbial community that shifted the SOM mining to N-rich components; an effect that increased with higher fungal dominance. Extrapolating from these findings, the predicted shrub expansion in the subarctic could result in an altered microbial use of SOM, selectively mining it for N-rich components, and leading to a reduced total SOM-use.