Climatic and biotic extreme events moderate long‐term responses of above‐ and belowground sub‐Arctic heathland communities to climate change

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.     

Climate change and cyclic predator–prey population dynamics in the high Arctic

The high Arctic has the world's simplest terrestrial vertebrate predator–prey community, with the collared lemming being the single main prey of four predators, the snowy owl, the Arctic fox, the long-tailed skua, and the stoat. Using a 20-year-long time series of population densities for the five species and a dynamic model that has been previously parameterized for northeast Greenland, we analyzed the population and community level consequences of the ongoing and predicted climate change. Species' responses to climate change are complex, because in addition to the direct effects of climate change, which vary depending on species' life histories, species are also affected indirectly due to, e.g., predator–prey interactions. The lemming–predator community exemplifies these complications, yet a robust conclusion emerges from our modeling: in practically all likely scenarios of how climate change may influence the demography of the species, climate change increases the length of the lemming population cycle and decreases the maximum population densities. The latter change in particular is detrimental to the populations of the predators, which are adapted to make use of the years of the greatest prey abundance. Therefore, climate change will indirectly reduce the predators' reproductive success and population densities, and may ultimately lead to local extinction of some of the predator species. Based on these results, we conclude that the recent anomalous observations about lack of cyclic lemming dynamics in eastern Greenland may well be the first signs of a severe impact of climate change on the lemming–predator communities in Greenland and elsewhere in the high Arctic.     

The future of cold‐adapted plants in changing climates: Micranthes (Saxifragaceae) as a case study

Research has shown species undergoing range contractions and/or northward and higher elevational movements as a result of changing climates. Here, we evaluate how the distribution of a group of cold‐adapted plant species with similar evolutionary histories changes in response to warming climates. We selected 29 species of Micranthes (Saxifragaceae) representing the mountain and Arctic biomes of the Northern Hemisphere. For this analysis, 24,755 data points were input into ecological niche models to assess both present fundamental niches and predicted future ranges under climate change scenarios. Comparisons were made across the Northern Hemisphere between all cold‐adapted Micranthes, including Arctic species, montane species, and species defined as narrow endemics. Under future climate change models, 72% of the species would occupy smaller geographical areas than at present. This loss of habitat is most pronounced in Arctic species in general, but is also prevalent in species restricted to higher elevations in mountains. Additionally, narrowly endemic species restricted to high elevations were more susceptible to habitat loss than those species found at lower elevations. Using a large dataset and modeling habitat suitability at a global scale, our results empirically model the threats to cold‐adapted species as a result of warming climates. Although Arctic and alpine biomes share many underlying climate similarities, such as cold and short growing seasons, our results confirm that species in these climates have varied responses to climate change and that key abiotic variables differ between these two habitats.     

Nonlinear effects of climate on boreal rodent dynamics: mild winters do not negate high‐amplitude cycles

Small rodents are key species in many ecosystems. In boreal and subarctic environments, their importance is heightened by pronounced multiannual population cycles. Alarmingly, the previously regular rodent cycles appear to be collapsing simultaneously in many areas. Climate change, particularly decreasing snow quality or quantity in winter, is hypothesized as a causal factor, but the evidence is contradictory. Reliable analysis of population dynamics and the influence of climate thereon necessitate spatially and temporally extensive data. We combined data on vole abundances and climate, collected at 33 locations throughout Finland from 1970 to 2011, to test the hypothesis that warming winters are causing a disappearance of multiannual vole cycles. We predicted that vole population dynamics exhibit geographic and temporal variation associated with variation in climate; reduced cyclicity should be observed when and where winter weather has become milder. We found that the temporal patterns in cyclicity varied between climatically different regions: a transient reduction in cycle amplitude in the coldest region, low‐amplitude cycles or irregular dynamics in the climatically intermediate regions, and strengthening cyclicity in the warmest region. Our results did not support the hypothesis that mild winters are uniformly leading to irregular dynamics in boreal vole populations. Long and cold winters were neither a prerequisite for high‐amplitude multiannual cycles, nor were mild winters with reduced snow cover associated with reduced winter growth rates. Population dynamics correlated more strongly with growing season than with winter conditions. Cyclicity was weakened by increasing growing season temperatures in the cold, but strengthened in the warm regions. High‐amplitude multiannual vole cycles emerge in two climatic regimes: a winter‐driven cycle in cold, and a summer‐driven cycle in warm climates. Finally, we show that geographic climatic gradients alone may not reliably predict biological responses to climate change.     

Functional and numerical responses of four lemming predators in high arctic Greenland

The high‐arctic tundra ecosystem has the world's simplest vertebrate predator–prey community, with only four predators preying upon one rodent species, the collared lemming (Dicrostonyx groenlandicus). We document the functional and numerical responses of all the four predators in NE Greenland. Using these data, we assess the impact of predation on the dynamics of the collared lemming with a 4 yr cycle and >100‐fold difference between maximum and minimum densities. All predator species feed mostly (>90%) on lemmings when lemming density is >1 ha^?1, but the shapes of the predators’ responses vary greatly. The snowy owl (Nyctea scandiaca) is present and breeds only when lemming densities at snowmelt are >2 ha^?1, giving rise to a step‐like numerical response. The long‐tailed skua (Stercorarius longicaudus) has a type III functional response and shifts from alternate food (mainly berries and insects) to lemmings with increasing lemming density. The skua surpasses all the other predators in summer by its total response. The type III functional response of the Arctic fox (Alopex lagopus) starts to increase at much lower lemming densities than the responses of the avian predators, but it has only a weak numerical response. Finally, the stoat (Mustela erminea) is the most specialized predator and the only one with a clearly delayed numerical response. According to their specific functional and numerical responses, each predator plays a key role at some point of the lemming cycle, but only the stoat has the potential to drive the lemming cycle. Stoat predation is greatly reduced in the winter preceding the lemming peak, and it reaches a maximum in the winter preceding the lowest lemming summer density. Stoat predation appears to maintain low lemming densities for at least two successive years. Our study provides empirical support for the specialist predator hypothesis about small mammal population cycles.     

Potential for an Arctic‐breeding migratory bird to adjust spring migration phenology to Arctic amplification

Arctic amplification, the accelerated climate warming in the polar regions, is causing a more rapid advancement of the onset of spring in the Arctic than in temperate regions. Consequently, the arrival of many migratory birds in the Arctic is thought to become increasingly mismatched with the onset of local spring, consequently reducing individual fitness and potentially even population levels. We used a dynamic state variable model to study whether Arctic long‐distance migrants can advance their migratory schedules under climate warming scenarios which include Arctic amplification, and whether such an advancement is constrained by fuel accumulation or the ability to anticipate climatic changes. Our model predicts that barnacle geese Branta leucopsis suffer from considerably reduced reproductive success with increasing Arctic amplification through mistimed arrival, when they cannot anticipate a more rapid progress of Arctic spring from their wintering grounds. When geese are able to anticipate a more rapid progress of Arctic spring, they are predicted to advance their spring arrival under Arctic amplification up to 44 days without any reproductive costs in terms of optimal condition or timing of breeding. Negative effects of mistimed arrival on reproduction are predicted to be somewhat mitigated by increasing summer length under warming in the Arctic, as late arriving geese can still breed successfully. We conclude that adaptation to Arctic amplification may rather be constrained by the (un)predictability of changes in the Arctic spring than by the time available for fuel accumulation. Social migrants like geese tend to have a high behavioural plasticity regarding stopover site choice and migration schedule, giving them the potential to adapt to future climate changes on their flyway.     

Plant functional traits mediate reproductive phenology and success in response to experimental warming and snow addition in Tibet

Global climate change is predicted to have large impacts on the phenology and reproduction of alpine plants, which will have important implications for plant demography and community interactions, trophic dynamics, ecosystem energy balance, and human livelihoods. In this article we report results of a 3‐year, fully factorial experimental study exploring how warming, snow addition, and their combination affect reproductive phenology, effort, and success of four alpine plant species belonging to three different life forms in a semiarid, alpine meadow ecosystem on the central Tibetan Plateau. Our results indicate that warming and snow addition change reproductive phenology and success, but responses are not uniform across species. Moreover, traits associated with resource acquisition, such as rooting depth and life history (early vs. late flowering), mediate plant phenology, and reproductive responses to changing climatic conditions. Specifically, we found that warming delayed the reproductive phenology and decreased number of inflorescences of Kobresia pygmaea C. B. Clarke, a shallow‐rooted, early‐flowering plant, which may be mainly constrained by upper‐soil moisture availability. Because K. pygmaea is the dominant species in the alpine meadow ecosystem, these results may have important implications for ecosystem dynamics and for pastoralists and wildlife in the region.     

Rapid climate‐driven loss of breeding habitat for Arctic migratory birds

Millions of birds migrate to and from the Arctic each year, but rapid climate change in the High North could strongly affect where species are able to breed, disrupting migratory connections globally. We modelled the climatically suitable breeding conditions of 24 Arctic specialist shorebirds and projected them to 2070 and to the mid‐Holocene climatic optimum, the world's last major warming event ~6000 years ago. We show that climatically suitable breeding conditions could shift, contract and decline over the next 70 years, with 66–83% of species losing the majority of currently suitable area. This exceeds, in rate and magnitude, the impact of the mid‐Holocene climatic optimum. Suitable climatic conditions are predicted to decline acutely in the most species rich region, Beringia (western Alaska and eastern Russia), and become concentrated in the Eurasian and Canadian Arctic islands. These predicted spatial shifts of breeding grounds could affect the species composition of the world's major flyways. Encouragingly, protected area coverage of current and future climatically suitable breeding conditions generally meets target levels; however, there is a lack of protected areas within the Canadian Arctic where resource exploitation is a growing threat. Given that already there are rapid declines of many populations of Arctic migratory birds, our results emphasize the urgency of mitigating climate change and protecting Arctic biodiversity.     

Glacial cycles drive rapid divergence of cryptic field vole species

Understanding the factors that contribute to the generation of reproductively isolated forms is a fundamental goal of evolutionary biology. Cryptic species are an especially interesting challenge to study in this context since they lack obvious morphological differentiation that provides clues to adaptive divergence that may drive reproductive isolation. Geographical isolation in refugial areas during glacial cycling is known to be important for generating genetically divergent populations, but its role in the origination of new species is still not fully understood and likely to be situation dependent. We combine analysis of 35,434 single‐nucleotide polymorphisms (SNPs) with environmental niche modeling (ENM) to investigate genomic and ecological divergence in three cryptic species formerly classified as the field vole (Microtus agrestis). The SNPs demonstrate high genomic divergence (pairwise F_ST values of 0.45–0.72) and little evidence of gene flow among the three field vole cryptic species, and we argue that genetic drift may have been a particularly important mechanism for divergence in the group. The ENM reveals three areas as potential glacial refugia for the cryptic species and differing climatic niches, although with spatial overlap between species pairs. This evidence underscores the role that glacial cycling has in promoting genetic differentiation and reproductive isolation by subdivision into disjunct distributions at glacial maxima in areas relatively close to ice sheets. Future investigation of the intrinsic barriers to gene flow between the field vole cryptic species is required to fully assess the mechanisms that contribute to reproductive isolation. In addition, the Portuguese field vole (M. rozianus) shows a high inbreeding coefficient and a restricted climatic niche, and warrants investigation into its conservation status.     

Physiological and ecological effects of increasing temperature on fish production in lakes of Arctic Alaska

Lake ecosystems in the Arctic are changing rapidly due to climate warming. Lakes are sensitive integrators of climate‐induced changes and prominent features across the Arctic landscape, especially in lowland permafrost regions such as the Arctic Coastal Plain of Alaska. Despite many studies on the implications of climate warming, how fish populations will respond to lake changes is uncertain for Arctic ecosystems. Least Cisco (Coregonus sardinella) is a bellwether for Arctic lakes as an important consumer and prey resource. To explore the consequences of climate warming, we used a bioenergetics model to simulate changes in Least Cisco production under future climate scenarios for lakes on the Arctic Coastal Plain. First, we used current temperatures to fit Least Cisco consumption to observed annual growth. We then estimated growth, holding food availability, and then feeding rate constant, for future projections of temperature. Projected warmer water temperatures resulted in reduced Least Cisco production, especially for larger size classes, when food availability was held constant. While holding feeding rate constant, production of Least Cisco increased under all future scenarios with progressively more growth in warmer temperatures. Higher variability occurred with longer projections of time mirroring the expanding uncertainty in climate predictions further into the future. In addition to direct temperature effects on Least Cisco growth, we also considered changes in lake ice phenology and prey resources for Least Cisco. A shorter period of ice cover resulted in increased production, similar to warming temperatures. Altering prey quality had a larger effect on fish production in summer than winter and increased relative growth of younger rather than older age classes of Least Cisco. Overall, we predicted increased production of Least Cisco due to climate warming in lakes of Arctic Alaska. Understanding the implications of increased production of Least Cisco to the entire food web will be necessary to predict ecosystem responses in lakes of the Arctic.     

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.     

Spatial and temporal variability in ringed seal (Pusa hispida) stable isotopes in the Beaufort Sea

Arctic ecosystem dynamics are shifting in response to warming temperatures and sea ice loss. Such ecosystems may be monitored by examining the diet of upper trophic level species, which varies with prey availability. To assess interannual variation in the Beaufort Sea ecosystem, we examined spatial and temporal trends in ringed seal (Pusa hispida) δ¹³C and δ¹⁵N in claw growth layers grown from 1964 to 2011. Stable isotopes were correlated with climate indices, environmental conditions, seal population productivity, and geographic location. Sex and age did not influence stable isotopes. Enriched ¹³C was linked to cyclonic circulation regimes, seal productivity, and westward sampling locations. Higher δ¹⁵N was linked to lower sea surface temperatures, a higher percentage of pups in the subsistence harvest, and sample locations that were eastward and further from shore. From the 1960s to 2000s, ringed seal niche width expanded, suggesting a diversification of diet due to expansion of prey and/or seal space use. Overall, trends in ringed seal stable isotopes indicate changes within the Beaufort Sea ecosystem affected by water temperatures and circulation regimes. We suggest that continued monitoring of upper trophic level species will yield insights into changing ecosystem structure with climate change.     

Testing the environmental warming responses of Brachyscome daisy species using a common garden approach

As temperatures increase in a warming world, there will be different responses among related plant species, with some species able to increase growth rate under warmer conditions and others less likely. Here, we identify survival and growth parameters in a group of 19 related Australian daisies from the genera Brachyscome and Pembertonia when exposed to higher soil temperature, focusing particularly on species from the alpine environment. We used a common garden approach to measure growth and survival under warming. We tested for the effects of evolutionary history by investigating phylogeny and testing for a phylogenetic signal, and for the effects of ecological history by considering climatic variables associated with species distributions in their native range. Evolutionary history did not have a detectable effect on warming responses. While there was a moderate signal for plant growth in the absence of warming, there was no signal for growth changes in response to warming, despite variability among species to warming that ranged from positive to negative growth responses. There was no strong effect of climate context, as species that showed a positive response to warming did not necessarily originate from hotter environments. In fact, several species from hot environments grew relatively poorly when exposed to higher soil temperature. However, species endemic to alpine areas were less likely to benefit from warming than widespread species. We found a strong phylogenetic signal for climate history, in that closely related species tend to occur in areas with similar annual variability in precipitation. Species differences in response to soil warming were variable and difficult to link to climate conditions except for the poor response of alpine endemics. There was no significant association between survival and warming responses of species. However, as some species showed weak growth responses, this may reduce their fitness into the future.     

A century of fish growth in relation to climate change,population dynamics and exploitation

Marine ecosystems, particularly in high‐latitude regions such as the Arctic, have been significantly affected by human activities and contributions to climate change. Evaluating how fish populations responded to past changes in their environment is helpful for evaluating their future patterns, but is often hindered by the lack of long‐term biological data available. Using otolith increments of Northeast Arctic cod (Gadus morhua) as a proxy for individual growth, we developed a century‐scale biochronology (1924–2014) based on the measurements of 3,894 fish, which revealed significant variations in cod growth over the last 91 years. We combined mixed‐effect modeling and path analysis to relate these growth variations to selected climate, population and fishing‐related factors. Cod growth was negatively related to cod population size and positively related to capelin population size, one of the most important prey items. This suggests that density‐dependent effects are the main source of growth variability due to competition for resources and cannibalism. Growth was also positively correlated with warming sea temperatures but negatively correlated with the Atlantic Multidecadal Oscillation, suggesting contrasting effects of climate warming at different spatial scales. Fishing pressure had a significant but weak negative direct impact on growth. Additionally, path analysis revealed that the selected growth factors were interrelated. Capelin biomass was positively related to sea temperature and negatively influenced by herring biomass, while cod biomass was mainly driven by fishing mortality. Together, these results give a better understanding of how multiple interacting factors have shaped cod growth throughout a century, both directly and indirectly.     

Too hot to die? The effects of vegetation shading on past,present, and future activity budgets of two diurnal skinks from arid Australia

Behavioral thermoregulation is an important mechanism allowing ectotherms to respond to thermal variations. Its efficiency might become imperative for securing activity budgets under future climate change. For diurnal lizards, thermal microhabitat variability appears to be of high importance, especially in hot deserts where vegetation is highly scattered and sensitive to climatic fluctuations. We investigated the effects of a shading gradient from vegetation on body temperatures and activity timing for two diurnal, terrestrial desert lizards, Ctenotus regius, and Morethia boulengeri, and analyzed their changes under past, present, and future climatic conditions. Both species’ body temperatures and activity timing strongly depended on the shading gradient provided by vegetation heterogeneity. At high temperatures, shaded locations provided cooling temperatures and increased diurnal activity. Conversely, bushes also buffered cold temperature by saving heat. According to future climate change scenarios, cooler microhabitats might become beneficial to warm‐adapted species, such as C. regius, by increasing the duration of daily activity. Contrarily, warmer microhabitats might become unsuitable for less warm‐adapted species such as M. boulengeri for which midsummers might result in a complete restriction of activity irrespective of vegetation. However, total annual activity would still increase provided that individuals would be able to shift their seasonal timing towards spring and autumn. Overall, we highlight the critical importance of thermoregulatory behavior to buffer temperatures and its dependence on vegetation heterogeneity. Whereas studies often neglect ecological processes when anticipating species’ responses to future climate change the strongest impact of a changing climate on terrestrial ectotherms in hot deserts is likely to be the loss of shaded microhabitats rather than the rise in temperature itself. We argue that conservation strategies aiming at addressing future climate changes should focus more on the cascading effects of vegetation rather than on shifts of species distributions predicted solely by climatic envelopes.     

Climate change impact on seaweed meadow distribution in the North Atlantic rocky intertidal

The North-Atlantic has warmed faster than all other ocean basins and climate change scenarios predict sea surface temperature isotherms to shift up to 600 km northwards by the end of the 21st century. The pole-ward shift has already begun for many temperate seaweed species that are important intertidal foundation species. We asked the question: Where will climate change have the greatest impact on three foundational, macroalgal species that occur along North-Atlantic shores: Fucus serratus, Fucus vesiculosus, and Ascophyllum nodosum? To predict distributional changes of these key species under three IPCC (Intergovernmental Panel on Climate Change) climate change scenarios (A2, A1B, and B1) over the coming two centuries, we generated Ecological Niche Models with the program MAXENT. Model predictions suggest that these three species will shift northwards as an assemblage or “unit” and that phytogeographic changes will be most pronounced in the southern Arctic and the southern temperate provinces. Our models predict that Arctic shores in Canada, Greenland, and Spitsbergen will become suitable for all three species by 2100. Shores south of 45° North will become unsuitable for at least two of the three focal species on both the Northwest- and Northeast-Atlantic coasts by 2200. If these foundational species are unable to adapt to the rising temperatures, they will lose their centers of genetic diversity and their loss will trigger an unpredictable shift in the North-Atlantic intertidal ecosystem.     

Environmental drivers interactively affect individual tree growth across temperate European forests

Forecasting the growth of tree species to future environmental changes requires a better understanding of its determinants. Tree growth is known to respond to global‐change drivers such as climate change or atmospheric deposition, as well as to local land‐use drivers such as forest management. Yet, large geographical scale studies examining interactive growth responses to multiple global‐change drivers are relatively scarce and rarely consider management effects. Here, we assessed the interactive effects of three global‐change drivers (temperature, precipitation and nitrogen deposition) on individual tree growth of three study species (Quercus robur/petraea, Fagus sylvatica and Fraxinus excelsior). We sampled trees along spatial environmental gradients across Europe and accounted for the effects of management for Quercus. We collected increment cores from 267 trees distributed over 151 plots in 19 forest regions and characterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. We demonstrate that growth responds interactively to global‐change drivers, with species‐specific sensitivities to the combined factors. Simultaneously high levels of precipitation and deposition benefited Fraxinus, but negatively affected Quercus’ growth, highlighting species‐specific interactive tree growth responses to combined drivers. For Fagus, a stronger growth response to higher temperatures was found when precipitation was also higher, illustrating the potential negative effects of drought stress under warming for this species. Furthermore, we show that past forest management can modulate the effects of changing temperatures on Quercus’ growth; individuals in plots with a coppicing history showed stronger growth responses to higher temperatures. Overall, our findings highlight how tree growth can be interactively determined by global‐change drivers, and how these growth responses might be modulated by past forest management. By showing future growth changes for scenarios of environmental change, we stress the importance of considering multiple drivers, including past management and their interactions, when predicting tree growth.     

The devil is in the detail: Nonadditive and context‐dependent plant population responses to increasing temperature and precipitation

In climate change ecology, simplistic research approaches may yield unrealistically simplistic answers to often more complicated problems. In particular, the complexity of vegetation responses to global climate change begs a better understanding of the impacts of concomitant changes in several climatic drivers, how these impacts vary across different climatic contexts, and of the demographic processes underlying population changes. Using a replicated, factorial, whole‐community transplant experiment, we investigated regional variation in demographic responses of plant populations to increased temperature and/or precipitation. Across four perennial forb species and 12 sites, we found strong responses to both temperature and precipitation change. Changes in population growth rates were mainly due to changes in survival and clonality. In three of the four study species, the combined increase in temperature and precipitation reflected nonadditive, antagonistic interactions of the single climatic changes for population growth rate and survival, while the interactions were additive and synergistic for clonality. This disparity affects the persistence of genotypes, but also suggests that the mechanisms behind the responses of the vital rates differ. In addition, survival effects varied systematically with climatic context, with wetter and warmer + wetter transplants showing less positive or more negative responses at warmer sites. The detailed demographic approach yields important mechanistic insights into how concomitant changes in temperature and precipitation affect plants, which makes our results generalizable beyond the four study species. Our comprehensive study design illustrates the power of replicated field experiments in disentangling the complex relationships and patterns that govern climate change impacts across real‐world species and landscapes.     

Arctic browning: Impacts of extreme climatic events on heathland ecosystem CO2 fluxes

Extreme climatic events are among the drivers of recent declines in plant biomass and productivity observed across Arctic ecosystems, known as “Arctic browning.” These events can cause landscape‐scale vegetation damage and so are likely to have major impacts on ecosystem CO₂ balance. However, there is little understanding of the impacts on CO₂ fluxes, especially across the growing season. Furthermore, while widespread shoot mortality is commonly observed with browning events, recent observations show that shoot stress responses are also common, and manifest as high levels of persistent anthocyanin pigmentation. Whether or how this response impacts ecosystem CO₂ fluxes is not known. To address these research needs, a growing season assessment of browning impacts following frost drought and extreme winter warming (both extreme climatic events) on the key ecosystem CO₂ fluxes Net Ecosystem Exchange (NEE), Gross Primary Productivity (GPP), ecosystem respiration (R_eco) and soil respiration (R_soil) was carried out in widespread sub‐Arctic dwarf shrub heathland, incorporating both mortality and stress responses. Browning (mortality and stress responses combined) caused considerable site‐level reductions in GPP and NEE (of up to 44%), with greatest impacts occurring at early and late season. Furthermore, impacts on CO₂ fluxes associated with stress often equalled or exceeded those resulting from vegetation mortality. This demonstrates that extreme events can have major impacts on ecosystem CO₂ balance, considerably reducing the carbon sink capacity of the ecosystem, even where vegetation is not killed. Structural Equation Modelling and additional measurements, including decomposition rates and leaf respiration, provided further insight into mechanisms underlying impacts of mortality and stress on CO₂ fluxes. The scale of reductions in ecosystem CO₂ uptake highlights the need for a process‐based understanding of Arctic browning in order to predict how vegetation and CO₂ balance will respond to continuing climate change.     

Assessing forest vulnerability to climate warming using a process‐based model of tree growth: bad prospects for rear‐edges

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO₂ emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of ?10.7% and ?16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.