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Observations of changes in phenology have provided some of the strongest signals of the effects of climate change on terrestrial ecosystems. The International Tundra Experiment (ITEX), initiated in the early 1990s, established a common protocol to measure plant phenology in tundra study areas across the globe. Today, this valuable collection of phenology measurements depicts the responses of plants at the colder extremes of our planet to experimental and ambient changes in temperature over the past decades. The database contains 150,434 phenology observations of 278 plant species taken at 28 study areas for periods of 1 to 26 years. Here we describe the full dataset to increase the visibility and use of these data in global analyses, and to invite phenology data contributions from underrepresented tundra locations. Portions of this tundra phenology database have been used in three recent syntheses, some datasets are expanded, others are from entirely new study areas, and the entirety of these data are now available at the Polar Data Catalogue., Arctic Science, 8 (3), ISSN:2368-7460

The largest alpine–nival vegetation permanent plot site in the Alps, the GLORIA mastersite Schrankogel (Tirol, Austria), provided evidence of warming-driven vegetation changes already 10 years after its establishment in 1994. Another decade later, in 2014, substantial compositional changes with increasing ratios of warmth-demanding to cold-adapted species have been found. The current study deals with species-specific responses involved in an ongoing vegetation transformation across the alpine–nival ecotone on Schrankogel by using presence/absence as well as cover data from permanent plots, situated between 2900 and 3400 masl. The number of occupied plots per species remained constant or even increased during the first decade, whereas disappearance events became more frequent during the second one, especially for cold-adapted specialists (subnival–nival species). Remarkably, the latter was accompanied by continued strong losses in cover of all subnival–nival species. These losses were more frequent in plots with a more thermophilous species composition, suggesting an increasing maladaptation of subnival–nival species to warmer habitat conditions and a successive trailing-edge decline. Several species with a distribution centre at lower elevations (alpine–subnival) markedly increased in cover, comparatively more so in colder plots, indicating a leading-edge expansion. Moreover, our findings show an increase in occupied plots and cover of almost all snowbed species, suggesting that areas previously with a too long snowpack period are now becoming suitable snowbed habitats. Vegetation gaps arising from population dieback of cold-adapted species, however, could only be partly filled by advancing species, indicating that species declines have occurred already before the onset of strong competition pressure.

Accepted manuscript version, licensed CC BY-NC-ND 4.0. Arctic tundra active-layer soils are at risk of soil organic carbon (SOC) depletion and degradation upon global climate warming because they are in a stage of relatively early decomposition. Non-growing season (NGS) warming is particularly pronounced, and observed increases of CO2 emissions during experimentally warmed NGSs give concern for great SOC losses to the atmosphere. Here, we used snow fences in Arctic Spitsbergen dwarf shrub tundra to simulate 1.86 °C NGS warming for 9 consecutive years, while growing season temperatures remained unchanged. In the snow fence treatment, the 4-11 cm thick A-horizon had a 2% lower SOC concentration and a 0.48 kg C m−2 smaller pool size than the controls, indicating SOC pool depletion. The snow fence treatment's A-horizon's alkyl/O-alkyl ratio was also significantly increased, indicating an advance of SOC degradation. The underlying 5 cm of B/C-horizon did not show these effects. Our results support the hypothesis that SOC depletion and degradation are connected to the long-term transience of observed ecosystem respiration (ER) increases upon soil warming. We suggest that the bulk of warming induced ER increases may originate from surface and not deep active layer or permafrost horizons. The observed losses of SOC might be significant for the ecosystem in question, but are in magnitude comparatively small relative to anthropogenic greenhouse gas enrichment of the atmosphere. We conclude that a positive feedback of carbon losses from surface soils of Arctic dwarf shrub tundra to anthropogenic forcing will be minor, but not negligible.

Climate driven species’ range shifts may interfere with existing protected area (PA) networks, resulting in a mismatch between places where species are currently protected and places where they can thrive in the future. Here, we assess the climate-smartness of the Austrian PA network by focusing on endemic species’ climatic niches and their future representation within PAs. We calculated endemic species’ climatic niches and climate space available in PAs within their dispersal reach under current and future climates, with the latter represented by three climate change scenarios and three time-steps (2030, 2050, and 2080). Niches were derived from the area of occupancy of species and the extent of PAs, respectively, and calculated as bivariate density kernels on gradients of mean annual temperature and annual precipitation. We then computed climatic representation of species’ niches in PAs as the proportion of the species’ kernel covered by the PA kernel. We found that under both a medium (RCP 4.5) and severe (RCP 8.5) climate change scenario, representation of endemic species’ climatic niches by PAs will decrease to a sixth for animals and to a third for plants, on average, toward the end of the century. Twenty to thirty percent of Austrian endemic species will then have no representation of their climatic niches in PAs anymore. Species with larger geographical and wider elevational ranges will lose less climatic niche representation. The declining representation of climatic niches in PAs implies that, even if PAs may secure the persistence of a part of these endemics, only a small portion of intraspecific diversity of many species may be represented in PAs in the future. We discuss our findings in the context of the varied elevational gradients found in Austria and suggest that the most promising strategies for safeguarding endemic species’ evolutionary potential are to limit the magnitude of climate change and to reduce other pressures that additionally threaten their survival.

In the Arctic, fungal mycelial growth takes place mainly during the cold season and beginning of growing season. Climate change induced increases of cold season temperatures may, hence, benefit fungal growth and increase their abundance. This is of particular importance for parasitic fungi, which may significantly shape Arctic vegetation composition. Here, we studied two contrasting plant parasitic fungi’s occurrences (biotrophic Exobasidium hypogenum Nannf. on the vascular plant Cassiope tetragona (L.) D. Don., and necrotrophic Pythium polare Tojo, van West & Hoshino on the moss Sanionia uncinata (Hedw.) Loeske) in response to increased snow depth, a method primarily used to increase cold season temperatures, after 7–13 years of snow manipulation in Adventdalen, Svalbard. We show that enhanced snow depth increased occurrences of both fungi tested here and indicate that increased fungal infections of host plants were at least partly responsible for decreases of host occurrences. Although bryophyte growth, in general, may be influenced by increased soil moisture and reduced competition from vascular plants, Pythium polare is likely enhanced by the combination of milder winter temperatures and moister environment provided by the snow. The relationships between host plants and fungal infection indicate ongoing processes involved in the dynamics of compositional adjustment to changing climate.

Advancing phenology is one of the most visible effects of climate change on plant communities, and has been especially pronounced in temperature-limited tundra ecosystems. However, phenological responses have been shown to differ greatly between species, with some species shifting phenology more than others. We analysed a database of 42,689 tundra plant phenological observations to show that warmer temperatures are leading to a contraction of community-level flowering seasons in tundra ecosystems due to a greater advancement in the flowering times of late-flowering species than early-flowering species. Shorter flowering seasons with a changing climate have the potential to alter trophic interactions in tundra ecosystems. Interestingly, these findings differ from those of warmer ecosystems, where early-flowering species have been found to be more sensitive to temperature change, suggesting that community-level phenological responses to warming can vary greatly between biomes.

Many Arctic regions currently experience an increase in winter snowfall as a result of climate change. Deepened snow can enhance thermal insulation of the underlying soil during winter, resulting in warmer soil temperatures that promote soil microbial nitrogen (N)-cycle processes and the availability of N and other nutrients. We conducted an ex situ study comparing the effects of deepened snow (using snow fences that have been installed for 3–13 years) on microbial N-cycle processes in late summer (late growing season) and winter (late snow-covered season) among five tundra sites in three different geographic locations across the Arctic (Greenland (dry and wet tundra), Canada (mesic tundra), and Svalbard, Norway (heath and meadow tundra)). Soil gross N cycling rates (mineralization, nitrification, immobilization of ammonium (NH4+) and nitrate (NO3−), and denitrification) were determined using a15N pool dilution. Potential denitrification activity (PDA) and nitrous oxide reductase activity (N2OR) were measured to assess denitrifying enzyme activities. The deepened snow treatment across all sites had a significant effect of the potential soil capacity of accelerating N cycling rates in late winter, including quadrupled gross nitrification, tripled NO3−-N immobilization, and doubled denitrification as well as significantly enhanced late summer gross N mineralization, denitrification (two-fold) and NH4+-N availability. The increase in gross N mineralization and nitrification rates were primarily driven by the availability of dissolved organic carbon (DOC) and nitrogen (DON) across sites. The largest increases in winter DOC and DON concentrations due to deepened snow were observed at the two wetter sites (wet and mesic tundra), and N cycling rates were also more strongly affected by deepened snow at these two sites than at the three other drier sites. Together, these results suggest that the potential effects of deepened winter snow in stimulating microbial N-cycling activities will be most pronounced in relatively moist tundra ecosystems. Hence, this study provides support to prior observations that growing season biogeochemical cycles in the Arctic are sensitive to snow depth with altered nutrient availability for microorganisms and vegetation. It can be speculated that on the one hand growing season N availability will increase and promote plant growth, but on the other hand foster increased water- and gaseous (e.g. N2 and N2O) N-losses with implications for overall nutrient status.

Recent warming in the Arctic, which has been amplified during the winter1–3, greatly enhances microbial decomposition of soil organic matter and subsequent release of carbon dioxide (CO2)4. However, the amount of CO2 released in winter is not known and has not been well represented by ecosystem models or empirically based estimates5,6. Here we synthesize regional in situ observations of CO2 flux from Arctic and boreal soils to assess current and future winter carbon losses from the northern permafrost domain. We estimate a contemporary loss of 1,662 TgC per year from the permafrost region during the winter season (October–April). This loss is greater than the average growing season carbon uptake for this region estimated from process models (−1,032 TgC per year). Extending model predictions to warmer conditions up to 2100 indicates that winter CO2 emissions will increase 17% under a moderate mitigation scenario—Representative Concentration Pathway 4.5—and 41% under business-as-usual emissions scenario—Representative Concentration Pathway 8.5. Our results provide a baseline for winter CO2 emissions from northern terrestrial regions and indicate that enhanced soil CO2 loss due to winter warming may offset growing season carbon uptake under future climatic conditions. Winter warming in the Arctic will increase the CO2 flux from soils. A pan-Arctic analysis shows a current loss of 1,662 TgC per year over the winter, exceeding estimated carbon uptake in the growing season; projections suggest a 17% increase under RCP 4.5 and a 41% increase under RCP 8.5 by 2100.

In the version of this Article originally published, the following sentence was missing from the Acknowledgements: “This work was supported by the Norwegian Research Council SnoEco project, grant number 230970”. This text has now been added.

Tundra soils store large amounts of carbon (C) that could be released through enhanced ecosystem respiration (ER) as the arctic warms. Over time, this may change the quantity and quality of available soil C pools, which in-turn may feedback and regulate ER responses to climate warming. Therefore, short-term increases in ER rates due to experimental warming may not be sustained over longer periods, as observed in other studies. One important aspect, which is often overlooked, is how climatic changes affecting ER in one season may carry-over and determine ER in following seasons. Using snow fences, we increased snow depth and thereby winter soil temperatures in a high-arctic site in Svalbard (78°N) and a low-arctic site in the Northwest Territories, Canada (64°N), for 5 and 9 years, respectively. Deepened snow enhanced winter ER while having negligible effect on growing-season soil temperatures and soil moisture. Growing-season ER at the high-arctic site was not affected by the snow treatment after 2 years. However, surprisingly, the deepened snow treatments significantly reduced growing-season ER rates after 5 years at the high-arctic site and after 8–9 years at the low-arctic site. We speculate that the reduction in ER rates, that became apparent only after several years of experimental manipulation, may, at least in part, be due to prolonged depletion of labile C substrate as a result of warmer soils over multiple cold seasons. Long-term changes in winter climate may therefore significantly influence annual net C balance not just because of increased wintertime C loss but also because of “legacy” effects on ER rates during the following growing seasons.

Altres ajuts europeus: P.A.W. was additionally supported by the European Union Fourth Environment and Climate Framework Programme (Project Number ENV4-CT970586)P.A.W. was additionally supported by the European Union Fourth Environment and Climate Framework Programme (Project Number ENV4-CT970586). The tundra is warming more rapidly than any other biome on Earth, and the potential ramifications are far-reaching because of global feedback effects between vegetation and climate. A better understanding of how environmental factors shape plant structure and function is crucial for predicting the consequences of environmental change for ecosystem functioning. Here we explore the biome-wide relationships between temperature, moisture and seven key plant functional traits both across space and over three decades of warming at 117 tundra locations. Spatial temperature-trait relationships were generally strong but soil moisture had a marked influence on the strength and direction of these relationships, highlighting the potentially important influence of changes in water availability on future trait shifts in tundra plant communities. Community height increased with warming across all sites over the past three decades, but other traits lagged far behind predicted rates of change. Our findings highlight the challenge of using space-for-time substitution to predict the functional consequences of future warming and suggest that functions that are tied closely to plant height will experience the most rapid change. They also reveal the strength with which environmental factors shape biotic communities at the coldest extremes of the planet and will help to improve projections of functional changes in tundra ecosystems with climate warming.

High mountain ecosystems and their biota are governed by low-temperature conditions and thus can be used as indicators for climate warming impacts on natural ecosystems, provided that long-term data exist. We used data from the largest alpine to nival permanent plot site in the Alps, established in the frame of the Global Observation Research Initiative in Alpine Environments (GLORIA) on Schrankogel in the Tyrolean Alps, Austria, in 1994, and resurveyed in 2004 and 2014. Vascular plant species richness per plot increased over the entire period, albeit to a lesser extent in the second decade, because disappearance events increased markedly in the latter period. Although presence/absence data could only marginally explain range shift dynamics, changes in species cover and plant community composition indicate an accelerating transformation towards a more warmth-demanding and more drought-adapted vegetation, which is strongest at the lowest, least rugged subsite. Divergent responses of vertical distribution groups of species suggest that direct warming effects, rather than competitive displacement, are the primary causes of the observed patterns. The continued decrease in cryophilic species could imply that trailing edge dynamics proceed more rapidly than successful colonisation, which would favour a period of accelerated species declines.

Published version. Source at http://dx.doi.org/10.1088/1748-9326/11/12/125006 The duration of specific periods within a plant’s life cycle are critical for plant growth and performance. In the High Arctic, the start of many of these phenological periods is determined by snowmelt date, which may change in a changing climate. It has been suggested that the end of these periods during late-season are triggered by external cues, such as day length, light quality or temperature, leading to the hypothesis that earlier or later snowmelt dates will lengthen or shorten the duration of these periods, respectively, and thereby affect plant performance.Wetested whether snowmelt date controls phenology and phenological period duration in High Arctic Svalbard using a melt timing gradient from natural and experimentally altered snow depths.Weinvestigated the response of early- and late-season phenophases from both vegetative and reproductive phenological periods of eight common species.Wefound that all phenophases follow snowmelt patterns, irrespective of timing of occurrence, vegetative or reproductive nature. Three of four phenological period durations based on these phenophases were fixed for most species, defining the studied species as periodic. Periodicity can thus be considered an evolutionary trait leading to disadvantages compared with aperiodic species and we conclude that the mesic and heath vegetation types in Svalbard are at risk of being outcompeted by invading, aperiodic species from milder biomes.

Cry-toxin genes originating from Bacillus thuringiensis are inserted into genetically modified (GM) plants, often called Bt-plants, to provide insect resistance to pests. Significant amounts of Bt-plant residues, and thus Cry-toxins, will be shed to soil and aquatic environments. We exposed Daphnia magna to purified Cry1Ab and Cry2Aa toxins for the full life-span of the animals. We used single toxins in different doses and combinations of toxins and Roundup(®), another potential stressor on the rise in agricultural ecosystems. Animals exposed to 4.5 mg/L (ppm) of Cry1Ab, Cry2Aa and the combination of both showed markedly higher mortality, smaller body size and very low juvenile production compared to controls. Animals exposed to 0.75 mg/L also showed a tendency towards increased mortality but with increased early fecundity compared to the controls. Roundup(®) stimulated animals to strong early reproductive output at the cost of later rapid mortality. We conclude that i) purified Cry-toxins in high concentrations are toxic to D. magna, indicating alternative modes-of-action for these Cry-toxins; ii) Cry-toxins act in combination, indicating that 'stacked events' may have stronger effects on non-target organisms; iii) further studies need to be done on combinatorial effects of multiple Cry-toxins and herbicides that co-occur in the environment.

Published version. Also available at http://dx.doi.org/10.3389/fmicb.2015.00399 The active layer of soil overlaying permafrost in the Arctic is subjected to dramatic annual changes in temperature and soil chemistry, which likely affect bacterial activity and community structure. We studied seasonal variations in the bacterial community of active layer soil from Svalbard (78°N) by co-extracting DNA and RNA from 12 soil cores collected monthly over a year. PCR amplicons of 16S rRNA genes (DNA) and reverse transcribed transcripts (cDNA) were quantified and sequenced to test for the effect of low winter temperature and seasonal variation in concentration of easily degradable organic matter on the bacterial communities. The copy number of 16S rRNA genes and transcripts revealed no distinct seasonal changes indicating potential bacterial activity during winter despite soil temperatures well below −10°C. Multivariate statistical analysis of the bacterial diversity data (DNA and cDNA libraries) revealed a season-based clustering of the samples, and, e.g., the relative abundance of potentially active Cyanobacteria peaked in June and Alphaproteobacteria increased over the summer and then declined from October to November. The structure of the bulk (DNA-based) community was significantly correlated with pH and dissolved organic carbon, while the potentially active (RNA-based) community structure was not significantly correlated with any of the measured soil parameters. A large fraction of the 16S rRNA transcripts was assigned to nitrogen-fixing bacteria (up to 24% in June) and phototrophic organisms (up to 48% in June) illustrating the potential importance of nitrogen fixation in otherwise nitrogen poor Arctic ecosystems and of phototrophic bacterial activity on the soil surface.

In tundra areas where the growing season is short, any delay in the start of summer may have a considerable effect on plant development, growth and reproductive success. Climate models suggest long-term changes in winter precipitation in the Arctic, which may lead to deeper snow cover and a resultant delay in date of snow melt. In this paper, we investigated the role of snow depth and melt out date on the phenological development and reproductive success of vascular plants in Adventdalen, Svalbard (78° 10'N, 16° 06'E). Effects of natural variations in snow accumulation were demonstrated using two vegetation types (snow depth: meadow 21 cm, heath 32 cm), and fences were used to experimentally increase snow depth by over 1m. Phenological delay was greatest directly after snowmelt in the earlier phenological phases, and had the largest effect on the early development of those species which normally green-up early (i.e. Dryas, Papaver, Salix, Saxifraga). Compressed growing seasons and length of the reproductive period led to a reduced reproductive success in some of the study species. There were fewer flowers, fewer plots with dispersing seeds, and lower germination rates. This can have consequences for plant establishment and community composition in the long-term.

The Arctic is one of the ecosystems most affected by climate change; in particular, winter temperatures and precipitation are predicted to increase with consequent changes to snow cover depth and duration. Whether the snow-free period will be shortened or prolonged depends on the extent and temporal patterns of the temperature and precipitation rise; resulting changes will likely affect plant growth with cascading effects throughout the ecosystem. We experimentally manipulated snow regimes using snow fences and shoveling and assessed aboveground size of eight common high arctic plant species weekly throughout the summer. We demonstrated that plant growth responded to snow regime, and that air temperature sum during the snow free period was the best predictor for plant size. The majority of our studied species showed periodic growth; increases in plant size stopped after certain cumulative temperatures were obtained. Plants in early snow-free treatments without additional spring warming were smaller than controls. Response to deeper snow with later melt-out varied between species and categorizing responses by growth forms or habitat associations did not reveal generic trends. We therefore stress the importance of examining responses at the species level, since generalized predictions of aboveground growth responses to changing snow regimes cannot be made.