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.     

Circumpolar arctic tundra biomass and productivity dynamics in response to projected climate change and herbivory

Satellite remote sensing data have indicated a general ‘greening’ trend in the arctic tundra biome. However, the observed changes based on remote sensing are the result of multiple environmental drivers, and the effects of individual controls such as warming, herbivory, and other disturbances on changes in vegetation biomass, community structure, and ecosystem function remain unclear. We apply ArcVeg, an arctic tundra vegetation dynamics model, to estimate potential changes in vegetation biomass and net primary production (NPP) at the plant community and functional type levels. ArcVeg is driven by soil nitrogen output from the Terrestrial Ecosystem Model, existing densities of Rangifer populations, and projected summer temperature changes by the NCAR CCSM4.0 general circulation model across the Arctic. We quantified the changes in aboveground biomass and NPP resulting from (i) observed herbivory only; (ii) projected climate change only; and (iii) coupled effects of projected climate change and herbivory. We evaluated model outputs of the absolute and relative differences in biomass and NPP by country, bioclimate subzone, and floristic province. Estimated potential biomass increases resulting from temperature increase only are approximately 5% greater than the biomass modeled due to coupled warming and herbivory. Such potential increases are greater in areas currently occupied by large or dense Rangifer herds such as the Nenets‐occupied regions in Russia (27% greater vegetation increase without herbivores). In addition, herbivory modulates shifts in plant community structure caused by warming. Plant functional types such as shrubs and mosses were affected to a greater degree than other functional types by either warming or herbivory or coupled effects of the two.     

Rapid carbon turnover beneath shrub and tree vegetation is associated with low soil carbon stocks at a subarctic treeline

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

Dwarf shrub and grass vegetation resistant to long‐term experimental warming while microarthropod abundance declines on the Falkland Islands

Dwarf shrubs are a dominant plant type across many regions of the Earth and have hence a large impact on carbon and nutrient cycling rates. Climate change impacts on dwarf shrubs have been extensively studied in the Northern Hemisphere, and there appears to be large variability in response between ecosystem types and regions. In the Southern Hemisphere, less data are available despite dwarf shrub vegetation being a dominant feature of southern South America and mountainous regions of the Southern Hemisphere. Here, we present the response of an Empetrum rubrum dwarf shrub and a Poa grass community to 12 years of experimental climate manipulation achieved using open top chambers on the Falkland Islands, a cold temperate island group in the South Atlantic. The dwarf shrub and grass vegetation did not change significantly in cover, biomass or species richness over the 12 years period in response to climate warming scenarios of up to 1°C reflecting annual warming levels predicted in this region for the coming decades. The soil microarthropod community, however, responded with declines in abundance (37%) under warming conditions in the grass community, but no such changes were observed in the dwarf shrub community. Overall, our data indicate that dwarf shrub communities are resistant to the levels of climate warming predicted over the coming decades in the southern South America region and will, therefore, remain a dominant driver of local ecosystem properties.     

Variable sensitivity of plant communities in Iceland to experimental warming

Facing an increased threat of rapid climate change in cold‐climate regions, it is important to understand the sensitivity of plant communities both in terms of degree and direction of community change. We studied responses to 3–5 years of moderate experimental warming by open‐top chambers in two widespread but contrasting tundra communities in Iceland. In a species‐poor and nutrient‐deficient moss heath, dominated by Racomitrium lanuginosum, mean daily air temperatures at surface were 1–2°C higher in the warmed plots than the controls whereas soil temperatures tended to be lower in the warmed plots throughout the season. In a species‐rich dwarf shrub heath on relatively rich soils at a cooler site, dominated by Betula nana and R. lanuginosum, temperature changes were in the same direction although more moderate. In the moss heath, there were no detectable community changes while significant changes were detected in the dwarf shrub heath: the abundance of deciduous and evergreen dwarf shrubs significantly increased (>50%), bryophytes decreased (18%) and canopy height increased (100%). Contrary to some other studies of tundra communities, we detected no changes in species richness or other diversity measures in either community and the abundance of lichens did not change. It is concluded that the sensitivity of Icelandic tundra communities to climate warming varies greatly depending on initial conditions in terms of species diversity, dominant species, soil and climatic conditions as well as land‐use history.     

Effects of Warming on Shrub Abundance and Chemistry Drive Ecosystem-Level Changes in a Forest–Tundra Ecotone

Tundra vegetation is responding rapidly to on-going climate warming. The changes in plant abundance and chemistry might have cascading effects on tundra food webs, but an integrated understanding of how the responses vary between habitats and across environmental gradients is lacking. We assessed responses in plant abundance and plant chemistry to warmer climate, both at species and community levels, in two different habitats. We used a long-term and multisite warming (OTC) experiment in the Scandinavian forest?Ctundra ecotone to investigate (i) changes in plant community composition and (ii) responses in foliar nitrogen, phosphorus, and carbon-based secondary compound concentrations in two dominant evergreen dwarf-shrubs (Empetrum hermaphroditum and Vaccinium vitis-idaea) and two deciduous shrubs (Vaccinium myrtillus and Betula nana). We found that initial plant community composition, and the functional traits of these plants, will determine the responsiveness of the community composition, and thus community traits, to experimental warming. Although changes in plant chemistry within species were minor, alterations in plant community composition drive changes in community-level nutrient concentrations. In view of projected climate change, our results suggest that plant abundance will increase in the future, but nutrient concentrations in the tundra field layer vegetation will decrease. These effects are large enough to have knock-on consequences for major ecosystem processes like herbivory and nutrient cycling. The reduced food quality could lead to weaker trophic cascades and weaker top down control of plant community biomass and composition in the future. However, the opposite effects in forest indicate that these changes might be obscured by advancing treeline forests.     

Herbivore impacts to the moss layer determine tundra ecosystem response to grazing and warming

Herbivory and climate are key environmental drivers, shaping ecosystems at high latitudes. Here, we focus on how these two drivers act in concert, influencing the high arctic tundra. We aim to investigate mechanisms through which herbivory by geese influences vegetation and soil processes in tundra ecosystems under ambient and warmed conditions. To achieve this, two grazing treatments, clipping plus faecal additions and moss removal, were implemented in conjunction with passive warming. Our key finding was that, in many cases, the tundra ecosystem response was determined by treatment impacts on the moss layer. Moss removal reduced the remaining moss layer depth by 30% and increased peak grass biomass by 27%. These impacts were probably due to observed higher soil temperatures and decomposition rates associated with moss removal. The positive impact of moss removal on grass biomass was even greater with warming, further supporting this conclusion. In contrast, moss removal reduced dwarf shrub biomass possibly resulting from increased exposure to desiccating winds. An intact moss layer buffered the soil to increased air temperature and as a result there was no response of vascular plant productivity to warming over the course of this study. In fact, moss removal impacts on soil temperature were nearly double those of warming, suggesting that the moss layer is a key component in controlling soil conditions. The moss layer also absorbed nutrients from faeces, promoting moss growth. We conclude that both herbivory and warming influence this high arctic ecosystem but that herbivory is the stronger driver of the two. Disturbance to the moss layer resulted in a shift towards a more grass-dominated system with less abundant mosses and shrubs, a trend that was further enhanced by warming. Thus herbivore impacts to the moss layer are key to understanding arctic ecosystem response to grazing and warming.     

Depth‐based differentiation in nitrogen uptake between graminoids and shrubs in an Arctic tundra plant community

Questions

The rapid climate warming in tundra ecosystems can increase nutrient availability in the soil, which may initiate shifts in vegetation composition. The direction in which the vegetation shifts will co‐determine whether Arctic warming is mitigated or accelerated, making the understanding of successional trajectories urgent. One of the key factors influencing the competitive relationships between plant species is their access to nutrients, depending on the depth where they take up most nutrients. However, nutrient uptake at different soil depths by tundra plant species that differ in rooting depth is unclear.

Location

Kytalyk Nature Reserve, northeast Siberia, Russia.

Methods

We injected ¹⁵N to 5 cm, 15 cm and the thaw front of the soil in a moist tussock tundra. The absorption of ¹⁵N by grasses, sedges, deciduous shrubs and evergreen shrubs from the three depths was compared.

Results

The results clearly show a vertical differentiation of N uptake by these plant functional types, corresponding to their rooting strategy. Shallow‐rooting dwarf shrubs were more capable of absorbing nutrients from the upper soil than from deeper soil. Deep‐rooting grasses and sedges were more capable of absorbing nutrients from deeper soil than the dwarf shrubs. The natural ¹⁵N abundances in control plants also indicate that graminoids can absorb more nutrients from the deeper soil than dwarf shrubs.

Conclusions

Our results show that graminoids and shrubs in the Arctic differ in their N uptake strategies, with graminoids profiting from nutrients released at the thaw front, while shrubs mainly forage in upper soil layers. Our results suggest that tundra vegetation will become graminoid‐dominated as permafrost thaw progresses and nutrient availability increases in the deep soil.     

Greater Abundance of <Emphasis Type="Italic">Betula nana</Emphasis> and Early Onset of the Growing Season Increase Ecosystem CO<Subscript>2</Subscript> Uptake in West Greenland

Expansion of deciduous shrubs is a common observation throughout the Arctic, with implications for carbon (C) cycling. Shrubs may increase net ecosystem C uptake through greater leaf area and gross ecosystem photosynthesis (GEP), and/or through cooler summer soils and reduced ecosystem respiration (ER). We used a space-for-time substitution combined with experimental warming at a Low Arctic site in West Greenland to examine the biophysical effects of increased temperature and Betula nana abundance on ecosystem CO₂ exchange. Communities dominated by Betula were much stronger C sinks than graminoid communities due to greater GEP and lower ER. The warming treatment had little effect on GEP, ER, or net ecosystem CO₂ exchange (NEE). The start of the growing season has been advancing at our study site, as indicated by long-term observations of plant phenology. In a retrospective analysis, we estimate that earlier onset of the growing season has increased the strength of the ecosystem C sink at rates of 1.3 and 2.1 g C m^?2 y^?1 in Betula and graminoid tundra, respectively, since 2002. However, earlier, and presumably longer, growing seasons may be associated with greater potential for drought stress. Our data suggest that mid-summer drought-induced GEP declines may partially offset C gains associated with an earlier start to the growing season. Our results suggest greater deciduous shrub abundance and longer growing seasons will likely lead to greater net C uptake in our study area, while highlighting important complexities associated with drought and plant community composition.     

Aboveground biomass allocation,leaf growth,and photosynthesis patterns in tundra plant forms in arctic Alaska

Summary Tundra plant growth forms can generally be characterized as consisting predominantly of low-growing perennial grasses and sedges, perennial herbaceous forbs, dwarf deciduous shrubs, and dwarf evergreen shrubs. Gross aboveground carbon allocation, leaf growth, and photosynthesis pattern studies were initiated to develop a quantitative understanding of the functional importance of these particular tundra growth forms. Photosynthetic capacities of 13 species were determined under standardized exposure conditions using a¹⁴CO₂ field system and ranged between 5 and 47 mg CO₂·g dry wt^-1·h^-1. These results, in conjunction with detailed leaf growth determinations, support the generalization that species with an evergreen growth form have lower photosynthetic capacities than species with a perennial graminoid, forb, or deciduous shrub growth form. However, these low photosynthetic capacities in evergreen shrubs are associated with relatively extended leaf longevities. Conversely, deciduous shrub forms exhibited high photosynthetic capacities, but were offset by relatively short leaf longevity periods. The perennial grasses, sedges, and forbs showed patterns intermediate to these. As a result, it appears that among tundra species of different growth form, photosynthetic capacity is inversely related to leaf longevity.     

Snow-induced changes in dwarf birch chemistry increase moth larval growth rate and level of herbivory

Changes in snow cover might influence arctic ecosystems to the same extent as increased temperatures. Although the duration of snow cover is generally expected to decrease in the future as a result of global warming, the amounts of snow might increase in arctic areas where much of the elevated precipitation will fall as snow. We examined the effects of an increased snow cover, as a result of a snow fence treatment, on soil nitrogen mineralization, plant phenology, plant chemistry (nitrogen and potential defense compounds), the level of invertebrate herbivory, and performance of invertebrate herbivores in an arctic ecosystem, using dwarf birch (Betula nana) and the autumnal moth (Epirrita autumnata) as study organisms. An enhanced and prolonged snow cover increased the level of herbivory on dwarf birch leaves. Larvae feeding on plants that had experienced enhanced snow cover grew faster and pupated earlier than larvae fed with plant material from control plots, indicating that plants from enhanced snow-lie plots produce higher-quality food to herbivores. The increased larval growth rate was strongly correlated with higher leaf nitrogen concentration in plants subjected to snow manipulation, and also to certain phenolic acids. Snow manipulation did not change net nitrogen mineralization rates in the soil or total carbon concentration in leaves, but it altered the within-season fluctuating pattern of leaf phenolic compounds. This study demonstrates a positive relationship between increased snow cover and level of herbivory on deciduous shrubs, thus proposing a negative feedback on the climate-induced dwarf shrub expansion in arctic areas.     

Browsing‐mediated shrub canopy changes drive composition and species richness in forest‐tundra ecosystems

Changes in climate and in browsing pressure are expected to alter the abundance of tundra shrubs thereby influencing the composition and species richness of plant communities. We investigated the associations between browsing, tundra shrub canopies and their understory vegetation by utilizing a long‐term (10–13 seasons) experiment controlling reindeer and ptarmigan herbivory in the subarctic forest tundra ecotone in northwestern Fennoscandia. In this area, there has also been a consistent increase in the yearly thermal sum and precipitation during the study period. The cover of shrubs increased 2.8–7.8 fold in exclosures and these contrasted with browsed control areas creating a sharp gradient of canopy cover of tundra shrubs across a variety of vegetation types. Browsing exclusions caused significant shifts in more productive vegetation types, whereas little or no shift occurred in low‐productive tundra communities. The increased deciduous shrub cover was associated with significant losses of understory plant species and shifts in functional composition, the latter being clearest in the most productive plant community types. The total cover of understory vegetation decreased along with increasing shrub cover, while the cover of litter showed the opposite response. The cover of cryptogams decreased along with increasing shrub cover, while the cover of forbs was favoured by a shrub cover. Increasing shrub cover decreased species richness of understory vegetation, which was mainly due to the decrease in the cryptogam species. The effects were consistent across different types of forest tundra vegetation indicating that shrub increase may have broad impacts on arctic vegetation diversity. Deciduous shrub cover is strongly regulated by reindeer browsing pressure and altered browsing pressure may result in a profound shrub expansion over the next one or two decades. Results suggest that the impact of an increase in shrubs on tundra plant richness is strong and browsing pressure effectively counteracts the effects of climate warming‐driven shrub expansion and hence maintains species richness.     

Long-term ecosystem level experiments at Toolik Lake, Alaska, and at Abisko, Northern Sweden: generalizations and differences in ecosystem and plant type responses to global change

Long‐term ecosystem‐level experiments, in which the environment is manipulated in a controlled manner, are important tools to predict the responses of ecosystem functioning and composition to future global change. We present the results of a meta‐analysis performed on the results of long‐term ecosystem‐level experiments near Toolik Lake, Alaska, and Abisko, Sweden. We quantified aboveground biomass responses of different arctic and subarctic ecosystems to experimental fertilization, warming and shading. We not only analysed the general patterns but also the differences in responsiveness between sites and regions. Aboveground plant biomass showed a broad similarity of responses in both locations, and also showed some important differences. In both locations, aboveground plant biomass, particularly the biomass of deciduous and graminoid plants, responded most strongly to nutrient addition. The biomass of mosses and lichens decreased in both locations as the biomass of vascular plants increased. An important difference between the two regions was the smaller positive aboveground biomass response of deciduous shrubs in Abisko as compared with Toolik Lake. Whereas in Toolik Lake Betula nana increased its dominance and replaced many of the other plant types, in Abisko all vascular plant types increased in abundance without major shifts in relative abundance. The differences between the responses of the dominant vegetation types of the Toolik Lake region, i.e. tussock tundra systems, and that of the Abisko region, i.e. heath systems, may have important implications for ecosystem development under expected patterns of global change. However, there were also large site‐specific differences within each region. Several potential mechanistic explanations for the differences between sites and regions are discussed. The response patterns show the need for analyses of joint data sets from many regions and sites, in order to uncover common responses to changes in climate across large arctic regions from regional or local responses.     

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.     

Leaf mineral nutrition of Arctic plants in response to warming and deeper snow in northern Alaska

Articulating the consequences of global climate change on terrestrial ecosystem biogeochemistry is a critical component of Arctic system studies. Leaf mineral nutrition responses of tundra plants is an important measure of changes in organismic and ecosystem attributes because leaf nitrogen and carbon contents effect photosynthesis, primary production, carbon budgets, leaf litter, and soil organic matter decomposition as well as herbivore forage quality. In this study, we used a longterm experiment where snow depth and summer temperatures were increased independently and together to articulate how a series of climate change scenarios would affect leaf N, leaf C, and leaf C:N for vegetation in dry and moist tussock tundra in northern Alaska, USA. Our findings were: 1) moist tundra vegetation is much more responsive to this suite of climate change scenarios than dry tundra with up to a 25% increase in leaf N; 2) life forms exhibit divergence in leaf C, N, and C:N with deciduous shrubs and graminoids having almost identical leaf N contents; 3) for some species, leaf mineral nutrition responses to these climate change scenarios are tundra type dependent ( Betula ), but for others ( Vaccinium vitis-idaea ), strong responses are exhibited regardless of tundra type; and 4) the seasonal patterns and magnitudes of leaf C and leaf N in deciduous and evergreen shrubs were responsive to conditions of deeper snow in winter. Leaf N is was generally higher immediately after emergence from the deep snow experimental treatments and leaf N was higher during the subsequent summer and fall, and the leaf C:N were lower, especially in deciduous shrubs. These findings indicate that coupled increases in snow depth and warmer summer temperatures will alter the magnitudes and patterns of leaf mineral nutrition and that the long term consequences of these changes may feed-forward and affect ecosystem processes.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

Detecting changes in arctic tundra plant communities in response to warming over decadal time scales

Detecting the response of vegetation to climate forcing as distinct from spatial and temporal variability may be difficult, if not impossible, over the typical duration of most field studies. We analyzed the spatial and interannual variability of plant functional type biomass from field studies in low arctic tussock tundra and compared these to climate change simulations of plant community composition using a dynamic tundra vegetation model (ArcVeg). Spatial heterogeneity of peak season live aboveground biomass was estimated using field samples taken from low arctic tundra at Ivotuk, Alaska (68.5°N, 155.7°W) in 1999. Coefficients of variation for live aboveground biomass at the 1 m² scale ranged from 14.6% for deciduous shrubs, 18.5% for graminoids and 25.3% for mosses to over 57% for forbs and lichens. Spatial heterogeneity in the ArcVeg dynamic vegetation model was simulated to be greater than the field data, ranging from 37.1% for deciduous shrubs to 107.9% for forbs. Disturbances in the model, such as caribou grazing and freezing–thawing of soil, as well as demographic stochasticity, led to the greater variability in the simulated results. Temporal variances of aboveground live biomass over a 19-year period using data from Toolik Lake, AK fell within the range of field and simulation spatial variances. However, simulations using ArcVeg suggest that temporal variability can be substantially less than site-scale spatial variability. Field data coupled with ArcVeg simulations of climate change scenarios indicate that some changes in plant community composition may be detectable within two decades following the onset of warming, and shrubs and mosses might be the key indicators of community change. Model simulations also project increasing landscape scale spatial heterogeneity (particularly of shrubs) with increasing temperatures.     

Plant phenological responses to a long‐term experimental extension of growing season and soil warming in the tussock tundra of Alaska

Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis‐idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species‐specific and growth form‐specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.     

Nitrogen Uptake During Fall, Winter and Spring Differs Among Plant Functional Groups in a Subarctic Heath Ecosystem

Nitrogen (N) is a critical resource for plant growth in tundra ecosystems, and species differences in the timing of N uptake may be an important feature regulating community composition and ecosystem productivity. We added ¹⁵N-labelled glycine to a subarctic heath tundra dominated by dwarf shrubs, mosses and graminoids in fall, and investigated its partitioning among ecosystem components at several time points (October, November, April, May, June) through to the following spring/early summer. Soil microbes had acquired 65?±?7% of the ¹⁵N tracer by October, but this pool decreased through winter to 37?±?7% by April indicating significant microbial N turnover prior to spring thaw. Only the evergreen dwarf shrubs showed active ¹⁵N acquisition before early May indicating that they had the highest potential of all functional groups for acquiring nutrients that became available in early spring. The faster-growing deciduous shrubs did not resume ¹⁵N acquisition until after early May indicating that they relied more on nitrogen made available later during the spring/early summer. The graminoids and mosses had no significant increases in ¹⁵N tracer recovery or tissue ¹⁵N tracer concentrations after the first harvest in October. However, the graminoids had the highest root ¹⁵N tracer concentrations of all functional groups in October indicating that they primarily relied on N made available during summer and fall. Our results suggest a temporal differentiation among plant functional groups in the post-winter resumption of N uptake with evergreen dwarf shrubs having the highest potential for early N uptake, followed by deciduous dwarf shrubs and graminoids.     

Open tundra persist,but arctic features decline—Vegetation changes in the warming Fennoscandian tundra

In the forest‐tundra ecotone of the North Fennoscandian inland, summer and winter temperatures have increased by two to three centigrades since 1965, which is expected to result in major vegetation changes. To document the expected expansion of woodlands and scrublands and its impact on the arctic vegetation, we repeated a vegetation transect study conducted in 1976 in the Darju, spanning from woodland to a summit, 200 m above the tree line. Contrary to our expectations, tree line movement was not detected, and there was no increase in willows or shrubby mountain birches, either. Nevertheless, the stability of tundra was apparent. Small‐sized, poorly competing arctic species had declined, lichen cover had decreased, and vascular plants, especially evergreen ericoid dwarf shrubs, had gained ground. The novel climate seems to favour competitive clonal species and species thriving in closed vegetation, creating a community hostile for seedling establishment, but equally hostile for many arctic species, too. Preventing trees and shrubs from invading the tundra is thus not sufficient for conserving arctic biota in the changing climate. The only dependable cure is to stop the global warming.