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Biomass data was quantified through the point quadrat hits (30 x 30 cm**2).

SummaryWinter is a period of dormancy for plants of cold environments. However, winter climate is changing, leading to an increasing frequency of stochastic warm periods (winter warming events) and concomitant reductions in snow cover. These conditions can break dormancy for some plants and expose them to freeze‐and‐thaw stress. Mosses are a major component of high‐latitude ecosystems, yet the longer‐term impacts of such winter warming events on mosses remain unknown.In order to determine the longer‐term legacy effects of winter warming events on mosses, we undertook a simulation of these events over three consecutive winters in a sub‐Arctic dwarf shrub‐dominated open woodland. The mat‐forming feather moss,Hylocomium splendens(the most abundant cryptogam in this system), is one of the most widespread Arctic and boreal mosses and plays a key functional role in ecosystems. We studied the ecophysiological performance of this moss during the summers of the experimental period (2007–2009) and in the following years (2010–2013).We show that the previously reported warming‐induced reduction in segment growth and photosynthesis during the experimental years was persistent. Four years after the last event, photosynthesis and segment growth were still 30 and 36% lower than control levels, which was only a slight improvement from 44 and 43% 4 years earlier. Winter warming did not affect segment symmetry. During the years after the last simulated event, in both warmed and control plots, chlorophyll fluorescence and segment growth, but not net photosynthesis, increased slightly. The increases were probably driven by increased summer rainfall over the study years, highlighting the sensitivity of this moss to rainfall change.Overall, the legacy effects shown here demonstrate that this widespread and important moss is likely to be significantly disadvantaged in a future sub‐Arctic climate where frequent winter warming events may become the norm. Given the key importance of mosses for soil insulation, shelter and carbon sequestration in high‐latitude regions, such persistent impacts may ultimately affect important ecosystem functions.ALay Summaryis available for this article.

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.

Winter climate and snow cover are the important drivers of plant community development in polar regions. However, the impacts of changing winter climate and associated changes in snow regime have received much less attention than changes during summer. Here, we synthesize the results from studies on the impacts of extreme winter weather events on polar heathland and lichen communities. Dwarf shrubs, mosses and soil arthropods were negatively impacted by extreme warming events while lichens showed variable responses to changes in extreme winter weather events. Snow mould formation underneath the snow may contribute to spatial heterogeneity in plant growth, arthropod communities and carbon cycling. Winter snow cover and depth will drive the reported impacts of winter climate change and add to spatial patterns in vegetation heterogeneity. The challenges ahead lie in obtaining better predictions on the snow patterns across the landscape and how these will be altered due to winter climate change.

AbstractClimate 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.

Passive chambers are used to examine the impacts of summer warming in Antarctica but, so far, impacts occurring outside the growing season, or related to extreme temperatures, have not been reported, despite their potentially large biological significance. In this review, we synthesise and discuss the microclimate impacts of passive warming chambers (closed, ventilated and Open Top Chamber-OTC) commonly used in Antarctic terrestrial habitats, paying special attention to seasonal warming, during the growing season and outside, extreme temperatures and freeze-thaw events. Both temperature increases and decreases were recorded throughout the year. Closed chambers caused earlier spring soil thaw (8-28 days) while OTCs delayed soil thaw (3-13 days). Smaller closed chamber types recorded the largest temperature extremes (up to 20°C higher than ambient) and longest periods (up to 11 h) of above ambient extreme temperatures, and even OTCs had above ambient temperature extremes over up to 5 consecutive hours. The frequency of freeze-thaw events was reduced by ~25%. All chamber types experienced extreme temperature ranges that could negatively affect biological responses, while warming during winter could result in depletion of limited metabolic resources. The effects outside the growing season could be as important in driving biological responses as the mean summer warming. We make suggestions for improving season-specific warming simulations and propose that seasonal and changed temperature patterns achieved under climate manipulations should be recognised explicitly in descriptions of treatment effects.

AbstractThe regional variability in tundra and boreal carbon dioxide (CO2) fluxes can be high, complicating efforts to quantify sink‐source patterns across the entire region. Statistical models are increasingly used to predict (i.e., upscale) CO2 fluxes across large spatial domains, but the reliability of different modeling techniques, each with different specifications and assumptions, has not been assessed in detail. Here, we compile eddy covariance and chamber measurements of annual and growing season CO2 fluxes of gross primary productivity (GPP), ecosystem respiration (ER), and net ecosystem exchange (NEE) during 1990–2015 from 148 terrestrial high‐latitude (i.e., tundra and boreal) sites to analyze the spatial patterns and drivers of CO2 fluxes and test the accuracy and uncertainty of different statistical models. CO2 fluxes were upscaled at relatively high spatial resolution (1 km2) across the high‐latitude region using five commonly used statistical models and their ensemble, that is, the median of all five models, using climatic, vegetation, and soil predictors. We found the performance of machine learning and ensemble predictions to outperform traditional regression methods. We also found the predictive performance of NEE‐focused models to be low, relative to models predicting GPP and ER. Our data compilation and ensemble predictions showed that CO2 sink strength was larger in the boreal biome (observed and predicted average annual NEE −46 and −29 g C m−2 yr−1, respectively) compared to tundra (average annual NEE +10 and −2 g C m−2 yr−1). This pattern was associated with large spatial variability, reflecting local heterogeneity in soil organic carbon stocks, climate, and vegetation productivity. The terrestrial ecosystem CO2 budget, estimated using the annual NEE ensemble prediction, suggests the high‐latitude region was on average an annual CO2 sink during 1990–2015, although uncertainty remains high.