Impacts of long-term snow climate change on a high-elevation cold desert shrubland, California, USA

Ecological responses to 50-year old manipulations of snow depth and melt timing were assessed using snow fences arrayed across 50 km of a shrub–conifer landscape mosaic in eastern California, USA. We compared how increased, decreased, and ambient snow depth affected patterns of vegetation community composition, fire fuel accumulation, and annual tree ring growth. We also tested the effect of snow depth on soil carbon storage based on total C content under the two co-dominant shrub species (Artemisia tridentata and Purshia tridentata) in comparison with open, intershrub sites. Increased snow depth reduced the cover of the N-fixing shrub P. tridentata but not the water-redistributing shrub A. tridentata. Annual ring growth was greater on +snow plots and lower on ?snow plots for the conifer Pinus jeffreyi but not for Pinus contorta. Graminoid cover and aboveground biomass indicated higher fire fuel accumulation where snow depth was increased. Dead shrub stem biomass was greater regardless of whether snow depth was increased or decreased. Results demonstrate community shifts, altered tree growth, feedbacks on carbon storage, and altered fire fuel accumulation as a result of changes in snow depth and melt timing for this high-elevation, snow-dominated ecotone under future climate scenarios that envision increased or decreased snow depth.     

Growth responses of Sphagnum hollows to a growing season lengthening manipulation in Alaskan Arctic tundra

The global increase in surface air temperature has produced an overall lengthening of the growing season by 3–5 days/decade in the Northern Hemisphere during the last 30 years. The direct impact of a longer growing season has not been well documented for Sphagnum moss communities in the Arctic. We hypothesized that an increase in the growing season length may be detrimental to Sphagnum growth as a result of photoinhibition caused by the plants emerging from snow near the seasonal peak of solar irradiance. We conducted an experiment from 1999 to 2002, lengthening the growing season in arctic Alaska, to determine the effects that this simulation of climate change had on the growth of hollows dominated by Sphagnum angustifolium. The lengthened growing season was associated with a decrease in annual moss height increment of 78 and 69 % for 1999 and 2000, respectively. These growth reductions may be related to freeze/thaw episodes and prolonged periods of cold in those years. The growth of individuals exposed to snow removal was also reduced by high global radiation. Overall, snow removal did not significantly affect the seasonal dynamics of growth, but seasonal patterns of growth strongly differed among years. These differences in seasonal dynamics suggest that Sphagnum growth is driven by opportunistic responses to favorable conditions rather than ontogenetic drivers. In addition, we compared environmental variation and growth between control and snow removal plots. Growth of Sphagnum in both treatments was stimulated by warmer soil temperatures and drier conditions. With earlier snowmelt as a result of warmer temperatures, it is likely that S. angustifolium will be subjected to higher levels of radiation and possibly greater photoinhibition which may lead to lower growth rates and significant implications for moss production in tussock tundra.     

Impacts of extreme winter warming in the sub‐Arctic: growing season responses of dwarf shrub heathland

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.     

Snow cover and extreme winter warming events control flower abundance of some,but not all species in high arctic Svalbard

The High Arctic winter is expected to be altered through ongoing and future climate change. Winter precipitation and snow depth are projected to increase and melt out dates change accordingly. Also, snow cover and depth will play an important role in protecting plant canopy from increasingly more frequent extreme winter warming events. Flower production of many Arctic plants is dependent on melt out timing, since season length determines resource availability for flower preformation. We erected snow fences to increase snow depth and shorten growing season, and counted flowers of six species over 5 years, during which we experienced two extreme winter warming events. Most species were resistant to snow cover increase, but two species reduced flower abundance due to shortened growing seasons. Cassiope tetragona responded strongly with fewer flowers in deep snow regimes during years without extreme events, while Stellaria crassipes responded partly. Snow pack thickness determined whether winter warming events had an effect on flower abundance of some species. Warming events clearly reduced flower abundance in shallow but not in deep snow regimes of Cassiope tetragona, but only marginally for Dryas octopetala. However, the affected species were resilient and individuals did not experience any long term effects. In the case of short or cold summers, a subset of species suffered reduced reproductive success, which may affect future plant composition through possible cascading competition effects. Extreme winter warming events were shown to expose the canopy to cold winter air. The following summer most of the overwintering flower buds could not produce flowers. Thus reproductive success is reduced if this occurs in subsequent years. We conclude that snow depth influences flower abundance by altering season length and by protecting or exposing flower buds to cold winter air, but most species studied are resistant to changes.     

Deepened winter snow cover enhances net ecosystem exchange and stabilizes plant community composition and productivity in a temperate grassland

Global warming has greatly altered winter snowfall patterns, and there is a trend towards increasing winter snow in semi‐arid regions in China. Winter snowfall is an important source of water during early spring in these water‐limited ecosystems, and it can also affect nutrient supply. However, we know little about how changes in winter snowfall will affect ecosystem productivity and plant community structure during the growing season. Here, we conducted a 5‐year winter snow manipulation experiment in a temperate grassland in Inner Mongolia. We measured ecosystem carbon flux from 2014 to 2018 and plant biomass and species composition from 2015 to 2018. We found that soil moisture increased under deepened winter snow in early growing season, particularly in deeper soil layers. Deepened snow increased the net ecosystem exchange of CO₂ (NEE) and reduced intra‐ and inter‐annual variation in NEE. Deepened snow did not affect aboveground plant biomass (AGB) but significantly increased root biomass. This suggested that the enhanced NEE was allocated to the belowground, which improved water acquisition and thus contributed to greater stability in NEE in deep‐snow plots. Interestingly, the AGB of grasses in the control plots declined over time, resulting in a shift towards a forb‐dominated system. Similar declines in grass AGB were also observed at three other locations in the region over the same time frame and are attributed to 4 years of below‐average precipitation during the growing season. By contrast, grass AGB was stabilized under deepened winter snow and plant community composition remained unchanged. Hence, our study demonstrates that increased winter snowfall may stabilize arid grassland systems by reducing resource competition, promoting coexistence between plant functional groups, which ultimately mitigates the impacts of chronic drought during the growing season.     

Initial Stages of Tundra Shrub Litter Decomposition May Be Accelerated by Deeper Winter Snow But Slowed Down by Spring Warming

The Arctic climate is projected to change during the coming century, with expected higher air temperatures and increased winter snowfall. These climatic changes might alter litter decomposition rates, which in turn could affect carbon (C) and nitrogen (N) cycling rates in tundra ecosystems. However, little is known of seasonal climate change effects on plant litter decomposition rates and N dynamics, hampering predictions of future arctic vegetation composition and the tundra C balance. We tested the effects of snow addition (snow fences), warming (open top chambers), and shrub removal (clipping), using a full-factorial experiment, on mass loss and N dynamics of two shrub tissue types with contrasting quality: deciduous shrub leaf litter (Salix glauca) and evergreen shrub shoots (Cassiope tetragona). We performed a 10.5-month decomposition experiment in a low-arctic shrub tundra heath in West-Greenland. Field incubations started in late fall, with harvests made after 249, 273, and 319 days of field incubation during early spring, summer and fall of the next year, respectively. We observed a positive effect of deeper snow on winter mass loss which is considered a result of observed higher soil winter temperatures and corresponding increased winter microbial litter decomposition in deep-snow plots. In contrast, warming reduced litter mass loss during spring, possibly because the dry spring conditions might have dried out the litter layer and thereby limited microbial litter decomposition. Shrub removal had a small positive effect on litter mass loss for C. tetragona during summer, but not for S. glauca. Nitrogen dynamics in decomposing leaves and shoots were not affected by the treatments but did show differences in temporal patterns between tissue types: there was a net immobilization of N by C. tetragona shoots after the winter incubation, while S. glauca leaf N-pools were unaltered over time. Our results support the widely hypothesized positive linkage between winter snow depth and litter decomposition rates in tundra ecosystems, but our results do not reveal changes in N dynamics during initial decomposition stages. Our study also shows contrasting impacts of spring warming and snow addition on shrub decomposition rates that might have important consequences for plant community composition and vegetation-climate feedbacks in rapidly changing tundra ecosystems.     

Phenology and species determine growing‐season albedo increase at the altitudinal limit of shrub growth in the sub‐Arctic

Arctic warming is resulting in reduced snow cover and increased shrub growth, both of which have been associated with altered land surface–atmospheric feedback processes involving sensible heat flux, ground heat flux and biogeochemical cycling. Using field measurements, we show that two common Arctic shrub species (Betula glandulosa and Salix pulchra), which are largely responsible for shrub encroachment in tundra, differed markedly in albedo and that albedo of both species increased as growing season progressed when measured at their altitudinal limit. A moveable apparatus was used to repeatedly measure albedo at six precise spots during the summer of 2012, and resampled in 2013. Contrary to the generally accepted view of shrub‐covered areas having low albedo in tundra, full‐canopy prostrate B. glandulosa had almost the highest albedo of all surfaces measured during the peak of the growing season. The higher midsummer albedo is also evident in localized MODIS albedo aggregated from 2000 to 2013, which displays a similar increase in growing‐season albedo. Using our field measurements, we show the ensemble summer increase in tundra albedo counteracts the generalized effect of earlier spring snow melt on surface energy balance by approximately 40%. This summer increase in albedo, when viewed in absolute values, is as large as the difference between the forest and tundra transition. These results indicate that near future (<50 years) changes in growing‐season albedo related to Arctic vegetation change are unlikely to be particularly large and might constitute a negative feedback to climate warming in certain circumstances. Future efforts to calculate energy budgets and a sensible heating feedback in the Arctic will require more detailed information about the relative abundance of different ground cover types, particularly shrub species and their respective growth forms and phenology.     

Range Expansion of Moose in Arctic Alaska Linked to Warming and Increased Shrub Habitat

Twentieth century warming has increased vegetation productivity and shrub cover across northern tundra and treeline regions, but effects on terrestrial wildlife have not been demonstrated on a comparable scale. During this period, Alaskan moose (Alces alces gigas) extended their range from the boreal forest into tundra riparian shrub habitat; similar extensions have been observed in Canada (A. a. andersoni) and Eurasia (A. a. alces). Northern moose distribution is thought to be limited by forage availability above the snow in late winter, so the observed increase in shrub habitat could be causing the northward moose establishment, but a previous hypothesis suggested that hunting cessation triggered moose establishment. Here, we use recent changes in shrub cover and empirical relationships between shrub height and growing season temperature to estimate available moose habitat in Arctic Alaska c. 1860. We estimate that riparian shrubs were approximately 1.1 m tall c. 1860, greatly reducing the available forage above the snowpack, compared to 2 m tall in 2009. We believe that increases in riparian shrub habitat after 1860 allowed moose to colonize tundra regions of Alaska hundreds of kilometers north and west of previous distribution limits. The northern shift in the distribution of moose, like that of snowshoe hares, has been in response to the spread of their shrub habitat in the Arctic, but at the same time, herbivores have likely had pronounced impacts on the structure and function of these shrub communities. These northward range shifts are a bellwether for other boreal species and their associated predators.     

Vegetation responses in Alaskan arctic tundra after 8 years of a summer warming and winter snow manipulation experiment

We used snow fences and small (1 m²) open‐topped fiberglass chambers (OTCs) to study the effects of changes in winter snow cover and summer air temperatures on arctic tundra. In 1994, two 60 m long, 2.8 m high snow fences, one in moist and the other in dry tundra, were erected at Toolik Lake, Alaska. OTCs paired with unwarmed plots, were placed along each experimental snow gradient and in control areas adjacent to the snowdrifts. After 8 years, the vegetation of the two sites, including that in control plots, had changed significantly. At both sites, the cover of shrubs, live vegetation, and litter, together with canopy height, had all increased, while lichen cover and diversity had decreased. At the moist site, bryophytes decreased in cover, while an increase in graminoids was almost entirely because of the response of the sedge Eriophorum vaginatum. These community changes were consistent with results found in studies of responses to warming and increased nutrient availability in the Arctic. However, during the time period of the experiment, summer temperature did not increase, but summer precipitation increased by 28%. The snow addition treatment affected species abundance, canopy height, and diversity, whereas the summer warming treatment had few measurable effects on vegetation. The interannual temperature fluctuation was considerably larger than the temperature increases within OTCs (<2°C), however. Snow addition also had a greater effect on microclimate by insulating vegetation from winter wind and temperature extremes, modifying winter soil temperatures, and increasing spring run‐off. Most increases in shrub cover and canopy height occurred in the medium snow‐depth zone (0.5–2 m) of the moist site, and the medium to deep snow‐depth zone (2–3 m) of the dry site. At the moist tundra site, deciduous shrubs, particularly Betula nana, increased in cover, while evergreen shrubs decreased. These differential responses were likely because of the larger production to biomass ratio in deciduous shrubs, combined with their more flexible growth response under changing environmental conditions. At the dry site, where deciduous shrubs were a minor part of the vegetation, evergreen shrubs increased in both cover and canopy height. These changes in abundance of functional groups are expected to affect most ecological processes, particularly the rate of litter decomposition, nutrient cycling, and both soil carbon and nitrogen pools. Also, changes in canopy structure, associated with increases in shrub abundance, are expected to alter the summer energy balance by increasing net radiation and evapotranspiration, thus altering soil moisture regimes.     

Demographic response of tundra small mammals to a snow fencing experiment

Snow cover is a key environmental component for tundra wildlife that will be affected by climate change. Change to the snow cover may affect the population dynamics of high‐latitude small mammals, which are active throughout the winter and reproduce under the snow. We experimentally tested the hypotheses that a deeper snow cover would enhance the densities and winter reproductive rates of small mammals, but that predation by mustelids could be higher in areas of increased small mammal density. We enhanced snow cover by setting out snow fences at three sites in the Canadian Arctic (Bylot Island, Nunavut, and Herschel Island and Komakuk Beach, Yukon) over periods ranging from one to four years. Densities of winter nests were higher where snow depth was increased but spring lemming densities did not increase on the experimental areas. Lemmings probably moved from areas of deep snow, their preferred winter habitat, to summer habitat during snow melt once the advantages associated with deep snow were gone. Our treatment had no effect on signs of reproduction in winter nests, proportion of lactating females in spring, or the proportion of juveniles caught in spring, which suggests that deep snow did not enhance reproduction. Results on predation were inconsistent across sites as predation by weasels was higher on the experimental area at one site but lower at two others and was not higher in areas of winter nest aggregations. Although this experiment provided us with several new insights about the impact of snow cover on the population dynamics of tundra small mammals, it also illustrates the challenges and difficulties associated with large‐scale experiments aimed at manipulating a critical climatic factor.     

Analyzing the Effects of Growing Season Length on the Net Ecosystem Production of an Alpine Grassland Using Model–Data Fusion

Alpine ecosystems are, similar to arctic ecosystems, characterized by a very long snow season. Previous studies investigating arctic or alpine ecosystems have shown that winter CO₂ effluxes can dominate the annual balance and that the timing and duration of the snow cover plays a crucial role for plant growth and phenology and might also influence the growing season ecosystem CO₂ strength and dynamics. The objective of this study was to analyze seasonal and annual CO₂ balances of a grassland site at an elevation of 2440 m a.s.l in the Swiss central Alps. We continuously measured the NEP using the eddy covariance method from June 2013 to October 2014, covering two growing seasons and one winter. We analyzed the influence of snow melt date on the CO₂ exchange dynamics at this site, because snow melt differed about 24 days between the 2 years. To this end, we employed a process-based ecosystem carbon cycling model to disentangle the co-occurring effects of growing season length, environmental conditions during the growing season, and physiological/structural properties of the canopy on the ecosystem carbon balance. During the measurement period, the site was a net sink for CO₂ although winter efflux contributed significantly to the total balance. The cumulative growing season NEP as well as mean and maximum daily CO₂ uptake rates was lower during the year with the later snow melt, and the results indicated that the differences were mainly due to differing growing season lengths.     

Decomposition of Senesced Leaf Litter is Faster in Tall Compared to Low Birch Shrub Tundra

Many Low Arctic tundra regions are currently undergoing a vegetation shift towards increasing growth and groundcover of tall deciduous shrubs due to recent climate warming. Vegetation change directly affects ecosystem carbon balance, but it can also affect soil biogeochemical cycling through physical and biological feedback mechanisms. Recent studies indicate that enhanced snow accumulation around relatively tall shrubs has negligible physical effect on litter decomposition rates. However, these investigations were no more than 3 years, and therefore may be insufficient to detect differences in inherently slow biogeochemical processes. Here, we report a 5-year study near Daring Lake, Canada, comparing Betula neoalaskana foliar litter decay rates within unmanipulated and snowfenced low-stature birch (height: ~?0.3 m) plots to test the physical effect of experimentally deepened snow, and within tall birch (height: ~?0.8 m) plots to test the combined physical and biological effects, that is, deepened snow plus strong birch dominance. Having corrected for carbon gain by the colonizing decomposers, actual litter carbon loss increased by approximately 25% in the tall birch relative to both low birch sites. Decay of lignin-like acid unhydrolizable litter residues also accelerated in the tall birch site, and a similar but lower magnitude response in the snowfenced low birch site indicated that physical effects of deepened snow were at least partially responsible. In contrast, deepened snow alone did not affect litter carbon loss. Our findings suggest that a combination of greater litter inputs, altered soil microbial community, enhanced soil nutrient pools, and warmer winter soils together promote relatively fast decomposition of recalcitrant litter carbon in tall birch shrub environments.     

Extreme winter warming events more negatively impact small rather than large soil fauna: shift in community composition explained by traits not taxa

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.     

Modelled changes in arctic tundra snow,energy and moisture fluxes due to increased shrubs

In arctic tundra, shrubs can significantly modify the distribution and physical characteristics of snow, influencing the exchanges of energy and moisture between terrestrial ecosystems and the atmosphere from winter into the growing season. These interactions were studied using a spatially distributed, physically based modelling system that represents key components of the land–atmosphere system. Simulations were run for 4 years, over a 4‐km² tundra domain located in arctic Alaska. A shrub increase was simulated by replacing the observed moist‐tundra and wet‐tundra vegetation classes with shrub‐tundra; a procedure that modified 77% of the simulation domain. The remaining 23% of the domain, primarily ridge tops, was left as the observed dry‐tundra vegetation class. The shrub enhancement increased the averaged snow depth of the domain by 14%, decreased blowing‐snow sublimation fluxes by 68%, and increased the snowcover's thermal resistance by 15%. The shrub increase also caused significant changes in snow‐depth distribution patterns; the shrub‐enhanced areas had deeper snow, and the non‐modified areas had less snow. This snow‐distribution change influenced the timing and magnitude of all surface energy‐balance components during snowmelt. The modified snow distributions also affected meltwater fluxes, leading to greater meltwater production late in the melt season. For a region with an annual snow‐free period of approximately 90 days, the snow‐covered period decreased by 11 days on the ridges and increased by 5 days in the shrub‐enhanced areas. Arctic shrub increases impact the spatial coupling of climatically important snow, energy and moisture interactions by producing changes in both shrub‐enhanced and non‐modified areas. In addition, the temporal coupling of the climate system was modified when additional moisture held within the snowcover, because of less winter sublimation, was released as snowmelt in the spring.     

高山林线交错带高山杜鹃的凋落物分解

凋落物分解是维持生态系统生产力、养分循环、土壤有机质形成的关键生态过程。高山林线交错带是陆地生态系统中对气候变化响应的敏感区域。季节变化和海拔梯度上的植被类型差异可能会影响该区域凋落物的分解,进而对高山生态系统的碳氮循环产生重要影响。采用凋落物分解袋的方法,研究了川西高山林线交错带优势种高山杜鹃(Rhododendron lapponicum)凋落叶在雪被期和生长季的分解特征。结果显示:(1)季节变化和植被类型对高山杜鹃凋落物的分解均具有显著影响(P0.05),凋落叶的质量损失主要发生在生长季且在高山林线最大,暗针叶林中雪被期的质量损失略高于生长季,但差异不显著;(2)林线交错带上高山杜鹃凋落叶分解缓慢,一年干物质失重率为9.62%,拟合分解系数k为0.145;(3)高山杜鹃凋落叶的质量变化主要体现在纤维素降解显著且集中在雪被期,木质素无明显降解,在高山林线上C/N、C/P、木质素/N变化幅度较小且C、N、P的释放表现得稳定而持续。结果表明,季节性雪被对林线交错带内高山杜鹃分解的影响不仅局限在雪被期内,雪被融化期间频繁的冻融作用和雪融水淋洗作用可能会促进高山杜鹃凋落物在生长季初期的分解。总的来看,在气候变暖的情景下,雪被的缩减、生长季的延长和高山杜鹃群落的扩张可能加速高山林线交错带高山杜鹃凋落物的分解。     

Experimental icing affects growth,mortality, and flowering in a high Arctic dwarf shrub

Effects of climate change are predicted to be greatest at high latitudes, with more pronounced warming in winter than summer. Extreme mid‐winter warm spells and heavy rain‐on‐snow events are already increasing in frequency in the Arctic, with implications for snow‐pack and ground‐ice formation. These may in turn affect key components of Arctic ecosystems. However, the fitness consequences of extreme winter weather events for tundra plants are not well understood, especially in the high Arctic. We simulated an extreme mid‐winter rain‐on‐snow event at a field site in high Arctic Svalbard (78°N) by experimentally encasing tundra vegetation in ice. After the subsequent growing season, we measured the effects of icing on growth and fitness indices in the common tundra plant, Arctic bell‐heather (Cassiope tetragona). The suitability of this species for retrospective growth analysis enabled us to compare shoot growth in pre and postmanipulation years in icing treatment and control plants, as well as shoot survival and flowering. Plants from icing treatment plots had higher shoot mortality and lower flowering success than controls. At the individual sample level, heavily flowering plants invested less in shoot growth than nonflowering plants, while shoot growth was positively related to the degree of shoot mortality. Therefore, contrary to expectation, undamaged shoots showed enhanced growth in ice treatment plants. This suggests that following damage, aboveground resources were allocated to the few remaining undamaged meristems. The enhanced shoot growth measured in our icing treatment plants has implications for climate studies based on retrospective analyses of Cassiope. As shoot growth in this species responds positively to summer warming, it also highlights a potentially complex interaction between summer and winter conditions. By documenting strong effects of icing on growth and reproduction of a widespread tundra plant, our study contributes to an understanding of Arctic plant responses to projected changes in winter climatic conditions.     

Shrub shading moderates the effects of weather on arthropod activity in arctic tundra

1. Rapid warming has facilitated an increase in deciduous shrub cover in arctic tundra. Because shrubs create a cooler microclimate during the growing season, shrub cover could modulate the effects of global warming on the phenology and activity of ectotherms, including arthropods. This possibility was explored here using two dominant arthropod groups (flies and wolf spiders) in Alaskan tundra. 2. We monitored arthropods with pitfall traps over five summers at four sites that differed in shrub abundance, and used generalised additive mixed models (GAMMs) to separate the two underlying components of pitfall trap catch: the seasonal trend in arthropod density and the effects of short‐term weather variation (air temperature, wind speed, rainfall, solar radiation) on arthropod activity. 3. We found that shrub cover significantly altered the seasonal trend in the abundance of flies by reducing early‐season pitfall catch, in line with observed later snowmelt in shrub‐dominated plots at these sites. 4. Additionally, shrub cover modulated the effects of many weather variables on arthropod activity: shrub cover shifted wolf spiders' temperature–activity relationship, dampened the positive effect of solar radiation on the activity of arthropods in total, and ameliorated the negative effect of wind on the activity of flies. 5. Thus, these results indicate that shrub encroachment will probably be accompanied by altered arthropod responses to warming and other key weather variables. Because the rate of key ecological processes – herbivory, decomposition, predation – are controlled by activity at the organismal level, these effects on arthropods will have long‐term ecosystem‐level consequences.     

Spatio-temporal patterns of ptarmigan occupancy relative to shrub cover in the Arctic

Rock and willow ptarmigan are abundant herbivores that require shrub habitats in arctic and alpine areas. Shrub expansion is likely to increase winter habitat availability for ptarmigan, which in turn influence shrub architecture and growth through browsing. Despite their ecological role in the Arctic, the distribution and movement patterns of ptarmigan are not well known, particularly in northern Alaska where shrub expansion is occurring. We used multi-season occupancy models to test whether ptarmigan occupancy varied within and among years, and the degree to which colonization and extinction probabilities were related to shrub cover and latitude. Aerial surveys were conducted from March to May in 2011 and April to May 2012 in a 21,230 km² area in northeastern Alaska. In areas with at least 30 % shrub cover, the probability of colonization by ptarmigan was >0.90, indicating that moderate to extensive patches of shrubs (typically associated with riparian areas) had a high probability of becoming occupied by ptarmigan. Occupancy increased throughout the spring in both years, providing evidence that ptarmigan migrated from southern wintering areas to breeding areas north of the Brooks Range. Occupancy was higher in the moderate snow year than the high snow year, and this was likely due to higher shrub cover in the moderate snow year. Ptarmigan distribution and migration in the Arctic are linked to expanding shrub communities on a wide geographic scale, and these relationships may be shaping ptarmigan population dynamics, as well as rates and patterns of shrub expansion.     

Shrub expansion may reduce summer permafrost thaw in Siberian tundra

Climate change is expected to cause extensive vegetation changes in the Arctic: deciduous shrubs are already expanding, in response to climate warming. The results from transect studies suggest that increasing shrub cover will impact significantly on the surface energy balance. However, little is known about the direct effects of shrub cover on permafrost thaw during summer. We experimentally quantified the influence of Betula nana cover on permafrost thaw in a moist tundra site in northeast Siberia with continuous permafrost. We measured the thaw depth of the soil, also called the active layer thickness (ALT), ground heat flux and net radiation in 10 m diameter plots with natural B. nana cover (control plots) and in plots in which B. nana was removed (removal plots). Removal of B. nana increased ALT by 9% on average late in the growing season, compared with control plots. Differences in ALT correlated well with differences in ground heat flux between the control plots and B. nana removal plots. In the undisturbed control plots, we found an inverse correlation between B. nana cover and late growing season ALT. These results suggest that the expected expansion of deciduous shrubs in the Arctic region, triggered by climate warming, may reduce summer permafrost thaw. Increased shrub growth may thus partially offset further permafrost degradation by future temperature increases. Permafrost models need to include a dynamic vegetation component to accurately predict future permafrost thaw.     

Winter warming pulses differently affect plant performance in temperate heathland and grassland communities

Winter air temperature variability is projected to increase in the temperate zone whereas snow cover is projected to decrease, leading to more variable soil temperatures. In a field experiment winter warming pulses were applied and aboveground biomass and root length of four plant species were quantified over two subsequent growing seasons in monocultures and mixtures of two species. The experiment was replicated at two sites, a colder upland site with more snow and a warmer, dryer lowland site. Aboveground biomass of Holcus lanatus declined (?29 %) in the growing season after the warming pulse treatment. Its competitor in the grassland mixture, Plantago lanceolata, profited from this decline by increased biomass production (+18 %). These effects disappeared in the second year. There was a strong decline in biomass for P. lanceolata at the lowland site in the second year. These two species also showed a decline in leaf carbohydrate content during the manipulation. Aboveground productivity and carbohydrate content of the heathland species was not affected by the treatment. The aboveground effects of the treatment did not differ significantly between the two sites, thereby implying some generality for different temperate ecosystems with little and significant amount of snowfall. Root length increased directly after the treatment for H. lanatus and for Calluna vulgaris with a peak at the end of the first growing season. The observed species-specific effects emphasize the ecological importance of winter temperature variability in the temperate zone and appear important for potential shifts in community composition and ecosystem productivity.