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The Arctic is experiencing rapid climate change. This research documents changes to tundra vegetation near Atqasuk and Utqiaġvik, Alaska. At each location, 30 plots were sampled annually from 2010 to 2019 using a point frame. For every encounter, we recorded the height and classified it into eight groupings (deciduous shrubs, evergreen shrubs, forbs, graminoids, bryophytes, lichens, litter, and standing dead vegetation); for vascular plants we also identified the species. We found an increase in plant stature and cover over time, consistent with regional warming. Graminoid cover and height increased at both sites, with a 5-fold increase in cover in Atqasuk. At Atqasuk, the cover and height of shrubs and forbs increased. Species diversity decreased at both the sites. Year was generally the strongest predictor of vegetation change, suggesting a cumulative change over time; however, soil moisture and soil temperature were also predictors of vegetation change. We anticipate that plants in the region will continue to grow taller as the region warms, resulting in greater plant cover, especially of graminoids and shrubs. The increase in plant cover and accumulation of litter may negatively impact non-vascular plants. Continued changes in community structure will impact energy balance and carbon cycling and may have regional and global consequences.

The Arctic is warming twice as fast as the rest of the globe. Graminoid, deciduous shrub, and evergreen shrub cover has increased in some regions, but not others. To better understand why plant responses vary across regions, we compared change in plant cover over time with nine functional traits of 12 dominant species in three regions of northern Alaska (Utqiaġvik, Atqasuk, and Toolik Lake). Cover was measured three times from 2008 to 2018. Repeated-measures analysis of variance (ANOVA) found that one species — Carex aquatilis — showed significant change in cover over time, increasing by 12.7% at Atqasuk. Canonical correspondence analysis suggested a relationship between shifts in species cover and traits, but Pearson and Spearman rank correlations did not find a significant trend for any trait when analyzed individually. Investigation of community-weighted means (CWMs) for each trait revealed no significant changes over time for any trait in any region. By comparison, estimated ecosystem values for several traits important to ecosystem functioning showed consistent increases over time in two regions (Utqiaġvik and Atqasuk). Our results indicate that vascular plant community composition and function have remained consistent over time; however, documented increases in total plant cover have important implications for ecosystem functioning.

Increases in shrub growth and canopy cover are well documented community responses to climate warming in the Arctic. An important consequence of larger deciduous shrubs is shading of prostrate plant species, many of which are important sources of nectar and berries. Here, we present the impact of a shading experiment on two prostrate shrubs, Vaccinium vitis-idaea L. and Arctous alpina L., in northern Alaska over two growing seasons. We implemented three levels of shading (no shade, 40% shade, and 80% shade) in dry heath and moist acidic tundra. Plots were monitored for soil moisture content, surface temperature, normalized difference vegetation index (NDVI), and flowering. Shading was shown to, on average, lower surface temperature (0.7 °C to 5.3 °C) and increase soil moisture content (0.5% to 5.6%) in both communities. Both species- and plot-level NDVI values were delayed in timing of peak values (7 to 13 days) and decreased at the highest shading. Flower abundance of both species was lower in shaded plots and peak flowering was delayed (3 to 8 days) compared with controls. Changes in timing may result in phenological mismatches and can impact other trophic levels in the Arctic as both the flowers and resulting berries are important food sources for animals.

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

Open top chambers (OTCs) were adopted as the recommended warming mechanism by the International Tundra Experiment network in the early 1990s. Since then, OTCs have been deployed across the globe. Hundreds of papers have reported the impacts of OTCs on the abiotic environment and the biota. Here, we review the impacts of the OTC on the physical environment, with comments on the appropriateness of using OTCs to characterize the response of biota to warming. The purpose of this review is to guide readers to previously published work and to provide recommendations for continued use of OTCs to understand the implications of warming on low stature ecosystems. In short, the OTC is a useful tool to experimentally manipulate temperature; however, the characteristics and magnitude of warming varies greatly in different environments; therefore, it is important to document chamber performance to maximize the interpretation of biotic response. When coupled with long-term monitoring, warming experiments are a valuable means to understand the impacts of climate change on natural ecosystems. Open top chambers (OTCs) were adopted as the recommended warming mechanism by the International Tundra Experiment network in the early 1990s. Since then, OTCs have been deployed across the globe. Hundreds of papers have reported the im-pacts of OTCs on the abiotic environment and the biota. Here, we review the impacts of the OTC on the physical environment, with comments on the appropriateness of using OTCs to characterize the response of biota to warming. The purpose of this review is to guide readers to previously published work and to provide recommendations for continued use of OTCs to under -stand the implications of warming on low stature ecosystems. In short, the OTC is a useful tool to experimentally manipulate temperature; however, the characteristics and magnitude of warming varies greatly in different environments; therefore, it is important to document chamber performance to maximize the interpretation of biotic response. When coupled with long-term monitoring, warming experiments are a valuable means to understand the impacts of climate change on natural ecosystems.

Increases in shrub growth and canopy cover are well documented community responses to climate warming in the Arctic. An important consequence of larger deciduous shrubs is shading of prostrate plant species, many of which are important sources of nectar and berries. Here, we present the impact of a shading experiment on two prostrate shrubs, Vaccinium vitis-idaea L. and Arctous alpina L., in northern Alaska over two growing seasons. We implemented three levels of shading (no shade, 40% shade, and 80% shade) in dry heath and moist acidic tundra. Plots were monitored for soil moisture content, surface temperature, normalized difference vegetation index (NDVI), and flowering. Shading was shown to, on average, lower surface temperature (0.7 °C to 5.3 °C) and increase soil moisture content (0.5% to 5.6%) in both communities. Both species- and plot-level NDVI values were delayed in timing of peak values (7 to 13 days) and decreased at the highest shading. Flower abundance of both species was lower in shaded plots and peak flowering was delayed (3 to 8 days) compared with controls. Changes in timing may result in phenological mismatches and can impact other trophic levels in the Arctic as both the flowers and resulting berries are important food sources for animals.

The relative contribution of bryophytes to plant diversity, primary productivity, and ecosystem functioning increases towards colder climates. Bryophytes respond to environmental changes at the species level, but because bryophyte species are relatively difficult to identify, they are often lumped into one functional group. Consequently, bryophyte function remains poorly resolved. Here, we explore how higher resolution of bryophyte functional diversity can be encouraged and implemented in tundra ecological studies. We briefly review previous bryophyte functional classifications and the roles of bryophytes in tundra ecosystems and their susceptibility to environmental change. Based on shoot morphology and colony organization, we then propose twelve easily distinguishable bryophyte functional groups. To illustrate how bryophyte functional groups can help elucidate variation in bryophyte effects and responses, we compiled existing data on water holding capacity, a key bryophyte trait. Although plant functional groups can mask potentially high interspecific and intraspecific variability, we found better separation of bryophyte functional group means compared with previous grouping systems regarding water holding capacity. This suggests that our bryophyte functional groups truly represent variation in the functional roles of bryophytes in tundra ecosystems. Lastly, we provide recommendations to improve the monitoring of bryophyte community changes in tundra study sites.

The Arctic is experiencing rapid climate change. This research documents changes to tundra vegetation near Atqasuk and Utqiaġvik, Alaska. At each location, 30 plots were sampled annually from 2010 to 2019 using a point frame. For every encounter, we recorded the height and classified it into eight groupings (deciduous shrubs, evergreen shrubs, forbs, graminoids, bryophytes, lichens, litter, and standing dead vegetation); for vascular plants we also identified the species. We found an increase in plant stature and cover over time, consistent with regional warming. Graminoid cover and height increased at both sites, with a 5-fold increase in cover in Atqasuk. At Atqasuk, the cover and height of shrubs and forbs increased. Species diversity decreased at both the sites. Year was generally the strongest predictor of vegetation change, suggesting a cumulative change over time; however, soil moisture and soil temperature were also predictors of vegetation change. We anticipate that plants in the region will continue to grow taller as the region warms, resulting in greater plant cover, especially of graminoids and shrubs. The increase in plant cover and accumulation of litter may negatively impact non-vascular plants. Continued changes in community structure will impact energy balance and carbon cycling and may have regional and global consequences.

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

Rapid climate warming is altering Arctic and alpine tundra ecosystem structure and function, including shifts in plant phenology. While the advancement of green up and flowering are well-documented, it remains unclear whether all phenophases, particularly those later in the season, will shift in unison or respond divergently to warming. Here, we present the largest synthesis to our knowledge of experimental warming effects on tundra plant phenology from the International Tundra Experiment. We examine the effect of warming on a suite of season-wide plant phenophases. Results challenge the expectation that all phenophases will advance in unison to warming. Instead, we find that experimental warming caused: (1) larger phenological shifts in reproductive versus vegetative phenophases and (2) advanced reproductive phenophases and green up but delayed leaf senescence which translated to a lengthening of the growing season by approximately 3%. Patterns were consistent across sites, plant species and over time. The advancement of reproductive seasons and lengthening of growing seasons may have significant consequences for trophic interactions and ecosystem function across the tundra., Nature Communications, 12 (1), ISSN:2041-1723

A warming Arctic has been associated with increases in aboveground plant biomass, specifically shrubs, and changes in vegetation cover. However, the magnitude and direction of changes in NDVI have not been consistent across different tundra types. Here we examine the responsiveness of fine-scale NDVI values to experimental warming at eight sites in northern Alaska, United States. Warming in our eight sites ranged in duration from 2‑23 seasons. Dry, wet and moist tundra communities were monitored for canopy surface temperatures and NDVI in ambient and experimentally-warmed plots at near-daily frequencies during the summer of 2017 to assess the impact of the warming treatment on the magnitude and timing of greening. Experimental warming increased canopy-level surface temperatures across all sites (+0.47 to +3.14˚C), with the strongest warming effect occurring during June and July and for the southernmost sites. Green-up was accelerated by warming at six sites, and autumn senescence was delayed at five sites. Warming increased the magnitude of peak NDVI values at five sites, decreased it at one site, and at two sites it did not change. Warming resulted in earlier peak NDVI at three sites and no significant change in the other sites. Shrub and graminoid cover was positively correlated with the magnitude of peak NDVI (r=0.37 to 0.60) while cryptogam influence was mixed. The magnitude and timing of peak NDVI showed considerable variability across sites. Warming extended the duration of the summer green season at most sites due to accelerated greening in the spring and delayed senescence in the autumn. We show that in a warmer Arctic (as simulated by our experiment) the timing and total period of carbon gain may change. Our results suggest these changes are dependent on community composition and abundance of specific growth forms and therefore will likely impact net primary productivity and trophic interactions.

The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world., It is unclear whether plant trait relationships found at the global scale extend to climatic extremes. Here the authors analyse six major aboveground traits to show that known plant trait relationships extend to the tundra biomes and exhibit the same two dimensions of variation detected at the global scale.

The Arctic is undergoing dramatic environmental change with rapidly rising surface temperatures, accelerating sea ice decline and changing snow regimes, all of which influence tundra plant phenology. Despite these changes, no globally consistent direction of trends in spring phenology has been reported across the Arctic. While spring has advanced at some sites, spring has delayed or not changed at other sites, highlighting substantial unexplained variation. Here, we test the relative importance of local temperatures, local snow melt date and regional spring drop in sea ice extent as controls of variation in spring phenology across different sites and species. Trends in long-term time series of spring leaf-out and flowering (average span: 18 years) were highly variable for the 14 tundra species monitored at our four study sites on the Arctic coasts of Alaska, Canada and Greenland, ranging from advances of 10.06 days per decade to delays of 1.67 days per decade. Spring temperatures and the day of spring drop in sea ice extent advanced at all sites (average 1°C per decade and 21 days per decade, respectively), but only those sites with advances in snow melt (average 5 days advance per decade) also had advancing phenology. Variation in spring plant phenology was best explained by snow melt date (mean effect: 0.45 days advance in phenology per day advance snow melt) and, to a lesser extent, by mean spring temperature (mean effect: 2.39 days advance in phenology per °C). In contrast to previous studies examining sea ice and phenology at different spatial scales, regional spring drop in sea ice extent did not predict spring phenology for any species or site in our analysis. Our findings highlight that tundra vegetation responses to global change are more complex than a direct response to warming and emphasize the importance of snow melt as a local driver of tundra spring phenology.

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

As the Arctic warms, vegetation is responding, and satellite measures indicate widespread greening at high latitudes. This ‘greening of the Arctic’ is among the world’s most important large-scale ecological responses to global climate change. However, a consensus is emerging that the underlying causes and future dynamics of so-called Arctic greening and browning trends are more complex, variable and inherently scale-dependent than previously thought. Here we summarize the complexities of observing and interpreting high-latitude greening to identify priorities for future research. Incorporating satellite and proximal remote sensing with in-situ data, while accounting for uncertainties and scale issues, will advance the study of past, present and future Arctic vegetation change. acceptedVersion

Summary To date, the majority of our knowledge regarding the impacts of herbivory on arctic ecosystem function has been restricted to short-term (

Premise of the study Understanding the relationship between plants and changing abiotic factors is necessary to document and anticipate the impacts of climate change. Methods We used data from long-term research sites at Barrow and Atqasuk, Alaska, to investigate trends in abiotic factors (snow melt and freeze-up dates, air and soil temperature, thaw depth, and soil moisture) and their relationships with plant traits (inflorescence height, leaf length, reproductive effort, and reproductive phenology) over time. Key results Several abiotic factors, including increasing air and soil temperatures, earlier snowmelt, delayed freeze-up, drier soils, and increasing thaw depths, showed nonsignificant tendencies over time that were consistent with the regional warming pattern observed in the Barrow area. Over the same period, plants showed consistent, although typically nonsignificant tendencies toward increasing inflorescence heights and reproductive efforts. Air and soil temperatures, measured as degree days, were consistently correlated with plant growth and reproductive effort. Reproductive effort was best predicted using abiotic conditions from the previous year. We also found that varying the base temperature used to calculate degree days changed the number of significant relationships between temperature and the trait: in general, reproductive phenologies in colder sites were better predicted using lower base temperatures, but the opposite held for those in warmer sites. Conclusions Plant response to changing abiotic factors is complex and varies by species, site, and trait; however, for six plant species, we have strong evidence that climate change will cause significant shifts in their growth and reproductive effort as the region continues to warm.

Motivation The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. Main types of variables included The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. Spatial location and grain BioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2). Time period and grain BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. Major taxa and level of measurement BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. Software format .csv and .SQL., Global Ecology and Biogeography, 27 (7), ISSN:1466-822X, ISSN:1466-8238

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

To address impacts of climate change on natural ecosystems, researchers need efficient and integrated ground-based sensor systems capable of detecting plant to ecosystem alterations to productivity, species composition, phenology, and structure and function over seasonal, inter-annual, and decadal time scales. Here, we introduce the Mobile Instrumented Sensor Platform (MISP), a versatile robotic sensor system that is suspended above or within the canopy and is designed to be adaptable for both short and long-term observations, and suitable for multiple ecosystems. The system is novel in that it is mobile, rather than static, the suite of sensors can be customized, and installation and operation requires minor surface distur bance relative to comparable systems already in use. MISP was developed as a contribution to the Arctic Observation Network’s International Tundra Experiment (AON-ITEX), where at five locations between the low and high Arctic we record observations of different tundra plant communities over a 50 m transect. Observations include air temperature, surface temperature, incoming and outgoing long- and short-wave radiation, albedo, Normalized Difference Vegetation Index (NDVI), three-dimensional video, two and three-dimensional photography, and hyperspectral reflectance. Data analysis has proven the system’s suitability for detecting subtle ecosystem changes across small-scale soil moisture and other gradients due to the mobile nature of the sampling. Long-term studies will benefit from this approach because sampling is repeatable with high spatial and temporal resolution and the system can be adapted to incorporate future technologies.

International audience; Environmental manipulation studies are integral to determining biological consequences of climate warming. Open Top Chambers (OTCs) have been widely used to assess summer warming effects on terrestrial biota, with their effects during other seasons normally being given less attention even though chambers are often deployed year-round. In addition, their effects on temperature extremes and freeze-thaw events are poorly documented. To provide robust documentation of the microclimatic influences of OTCs throughout the year, we analysed temperature data from 20 studies distributed across polar and alpine regions. The effects of OTCs on mean temperature showed a large range ( 0.9 to 2.1 °C) throughout the year, but did not differ significantly between studies. Increases in mean monthly and diurnal temperature were strongly related (R2 = 0.70) with irradiance, indicating that PAR can be used to predict the mean warming effect of OTCs. Deeper snow trapped in OTCs also induced higher temperatures at soil/vegetation level. OTC-induced changes in the frequency of freeze- thaw events included an increase in autumn and decreases in spring and summer. Frequency of high- temperature events in OTCs increased in spring, summer and autumn compared with non-manipulated control plots. Frequency of low-temperature events was reduced by deeper snow accumulation and higher mean temperatures. The strong interactions identified between aspects of ambient environmental conditions and effects of OTCs suggest that a detailed knowledge of snow depth, temperature and irradiance levels enables us to predict how OTCs will modify the microclimate at a particular site and season. Such predictive power allows a better mechanistic understanding of observed biotic response to experimental warming studies and for more informed design of future experiments. However, a need remains to quantify OTC effects on water availability and wind speed (affecting, for example, drying rates and water stress) in combination with microclimate measurements at organism level.

The Alaska Arctic Vegetation Archive (AVA-AK, GIVD-ID: NA-US-014) is a free, publically available database archive of vegetation-plot data from the Arctic tundra region of northern Alaska. The archive currently contains 24 datasets with 3,026 non-overlapping plots. Of these, 74% have geolocation data with 25-m or better precision. Species cover data and header data are stored in a Turboveg database. A standardized Pan Arctic Species List provides a consistent nomenclature for vascular plants, bryophytes, and lichens in the archive. A web-based online Alaska Arctic Geoecological Atlas (AGA-AK) allows viewing and downloading the species data in a variety of formats, and provides access to a wide variety of ancillary data. We conducted a preliminary cluster analysis of the first 16 datasets (1,613 plots) to examine how the spectrum of derived clusters is related to the suite of datasets, habitat types, and environmental gradients. Here, we present the contents of the archive, assess its strengths and weaknesses, and provide three supplementary files that include the data dictionary, a list of habitat types, an overview of the datasets, and details of the cluster analysis.

Passive warming manipulation methodologies, such as open-top chambers (OTCs), are a meaningful approach for interpretation of impacts of climate change on the Arctic tundra biome. The magnitude of OTC warming has been studied extensively, revealing an average plot-level warming of air temperature that ranges between 1 and 3 °C as measured by shielded resistive sensors or thermocouples. Studies have also shown that the amount of OTC warming depends in part on location climate, vegetation, and soil properties. While digital infrared thermometers have been employed in a few comparisons, most of the focus of the effectiveness of OTC warming has been on air or soil temperature rather than tissue or surface temperatures, which directly translate to metabolism. Here we used thermal infrared (TIR) photography to quantify tissue and surface temperatures and their spatial variability at a previously unavailable resolution (3–6 mm2). We analyzed plots at three locations that are part of the International Tundra Experiment (ITEX)-Arctic Observing Network (AON-ITEX) network along both moisture and latitudinal gradients spanning from the High Arctic (Barrow, AK, USA) to the Low Arctic (Toolik Lake, AK, USA). Our results show a range of OTC surface warming from 2.65 to 1.27 °C (31%–10%) at our three sites. The magnitude of surface warming detected by TIR imagery in this study was comparable to increases in air temperatures previously reported for these sites. However, the thermal images revealed wide ranges of surface temperatures within the OTCs, with some surfaces well above ambient unevenly distributed within the plots under sunny conditions. We note that analyzing radiometric temperature may be an alternative for future studies that examine data acquired at the same time of day from sites that are in close geographic proximity to avoid the requirement of emissivity or atmospheric correction for validation of results. We foresee future studies using TIR photography to describe species-level thermodynamics that could prove highly valuable toward a better understanding of species-specific responses to climate change in the Arctic.

The point frame method is ideal for vegetation monitoring because exact locations may be precisely resampled over time. Many researchers, including those associated with the International Tundra Experiment (ITEX), have used a modified point frame method to document vegetation change in response to climate change and experimental warming. To save, time only the uppermost and lowermost encounter (TB method) of a plant below each grid point is recorded as opposed to all contacts (AC method). Here, we investigate the ability of this shortcut to accurately assess the community change and estimate the aboveground plant biomass at four long-term sites in northern Alaska. We found that the TB method underrepresented the species cover (by omitting intermediate contacts) between 35 and 50 % of the taxa at a site; however, the magnitude was less than 2 % cover for all but 17 of the 109 taxa and site combinations, and despite the smaller number of contacts in the TB method, it was nearly equal in its ability to detect a change in cover in response to warming as the AC method (only 1 of the 109 cases showed differences in the significance of the response). Linear regressions showed the accuracy of the TB, and AC methods were also similar in estimating aboveground plant biomass (same cases were significant, and all but 2 were within r 2 of 0.1). Thus, we conclude that TB method outlined in the ITEX manual is an efficient and accurate way of assessing vegetation changes in tundra plant communities.

Temperature is increasing at unprecedented rates across most of the tundra biome. Remote-sensing data indicate that contemporary climate warming has already resulted in increased productivity over much of the Arctic, but plot-based evidence for vegetation transformation is not widespread. We analysed change in tundra vegetation surveyed between 1980 and 2010 in 158 plant communities spread across 46 locations. We found biome-wide trends of increased height of the plant canopy and maximum observed plant height for most vascular growth forms; increased abundance of litter; increased abundance of evergreen, low-growing and tall shrubs; and decreased abundance of bare ground. Intersite comparisons indicated an association between the degree of summer warming and change in vascular plant abundance, with shrubs, forbs and rushes increasing with warming. However, the association was dependent on the climate zone, the moisture regime and the presence of permafrost. Our data provide plot-scale evidence linking changes in vascular plant abundance to local summer warming in widely dispersed tundra locations across the globe. © 2012 Macmillan Publishers Limited. All rights reserved.

35 Abstract Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date.

Projected increases in air temperature and precipitation due to climate change in Arctic wetlands could dramatically affect ecosystem functioning. As a consequence, it is important to define the controls on evapotranspiration, which is the major pathway of water loss from these systems. We quantified the multi-year controls on midday arctic coastal wetland evapotranspiration measured with the eddy covariance method at two vegetated drained thaw lake basins near Barrow, Alaska. Variations in near-surface soil moisture and atmospheric vapor pressure deficits were found to have nonlinear effects on midday evapotranspiration rates. Vapor pressure deficits near and above 0.3 kPa appeared to be an important hydrological threshold, allowing latent heat fluxes to persistently exceed sensible heat fluxes. Dry soils increased the bulk surface resistance (water-limited). Wet soils favored ground heat flux and therefore limited the energy available to sensible and latent heat fluxes (energy-limited). Thus, midday evapotranspiration was suppressed on both dry and wet soils through different mechanisms. We also found that wet soils (ponding excluded) combined with large atmospheric demands resulted in an increased bulk surface resistance and therefore suppressing the evapotranspiration to below its potential rate (Priestley-Taylor α < 1.26). This is likely caused by the limited ability of mosses to transfer moisture during large atmospheric demands. Ultimately, in addition to net radiation, the various controlling factors on midday evapotranspiration (near-surface soil moisture, atmospheric vapor pressure, and the limited ability of mosses that are saturated at depth to transfer water during high atmospheric vapor demands) resulted in an average evapotranspiration rate of up to 75 % of the potential evapotranspiration rate. The multiple limitations on midday evapotranspiration rates have the potential to moderate interannual variation of total evapotranspiration and dampen excessive water loss during a warmer climate. Combined with the prevailing maritime winds and the projected increase in precipitation, these dampening mechanisms will likely prevent extensive future soil drying and hence maintain the presence of coastal wetlands.

Satellite observations have shown greening trends in tundra in response to climate change, suggesting increases in productivity. To better understand the ability of remote sensing to detect climate impacts on tundra vegetation productivity, we applied a photosynthetic light use efficiency model to simulated climate change treatments of tundra vegetation. We examined changes in the Normalized Difference Vegetation Index (NDVI) and photosynthetic light use efficiency (e) in experimental warming and moisture treatments designed to simulate climate change in northern Alaska. Plots were warmed either passively, using Open Top Chambers, or actively using electric heaters in the soil. In one set of plots water table depth was actively altered, while other plots were established in locations that were naturally wet or dry. Over two growing seasons, plot-level carbon flux and spectral reflectance measurements were collected, and the results were used to derive a light use efficiency model that could explore the effects of moisture and temperature treatments using remote sensing. Warming increased values of canopy greenness (NDVI) relative to control plots, this effect being more pronounced in wet plots than in dry plots. Light use efficiency (LUE), the relationship between absorbed photosynthetically active radiation (PAR) and gross ecosystem production (GEP), was consistent across warming treatments, growing season, subsequent years, and sites. However, LUE was affected by vegetation type, which varied with moisture; plots in naturally dry locations showed reduced light use efficiency relative to moist plots. Additionally moss exhibited reduced LUE relative to vascular plants. Understory moss production, not accounted for by the usual definition of the fraction of absorbed PAR (fAPAR), was found to be a significant part of total GEP, particularly in areas with low vascular plant cover. These results support the use of light use efficiency models driven by spectral reflectance for estimating GEP in tundra vegetation, provided effects of vegetation functional type (e.g. mosses versus vascular plants) and microtopography are considered.

Climate change is warming the temperatures and lengthening the Arctic growing season with potentially important effects on plant phenology. The ability of plant species to acclimate to changing climatic conditions will dictate the level to which their spatial coverage and habitat-type dominance is different in the future. While the effect of changes in temperature on phenology and species composition have been observed at the plot and at the regional scale, a systematic assessment at medium spatial scales using new noninvasive sensor techniques has not been performed yet. At four sites across the North Slope of Alaska, changes in the Normalized Difference Vegetation Index (NDVI) signal were observed by Mobile Instrumented Sensor Platforms (MISP) that are suspended over 50 m transects spanning local moisture gradients. The rates of greening (measured in June) and senescence (measured in August) in response to the air temperature was estimated by changes in NDVI measured as the difference between the NDVI on a specific date and three days later. In June, graminoid- and shrub-dominated habitats showed the greatest rates of NDVI increase in response to the high air temperatures, while forb- and lichen-dominated habitats were less responsive. In August, the NDVI was more responsive to variations in the daily average temperature than spring greening at all sites. For graminoid- and shrub-dominated habitats, we observed a delayed decrease of the NDVI, reflecting a prolonged growing season, in response to high August temperatures. Consequently, the annual C assimilation capacity of these habitats is increased, which in turn may be partially responsible for shrub expansion and further increases in net summer CO2 fixation. Strong interannual differences highlight that long-term and noninvasive measurements of such complex feedback mechanisms in arctic ecosystems are critical to fully articulate the net effects of climate variability and climate change on plant community and ecosystem processes.

Climate warming is expected to differentially affect CO2 exchange of the diverse ecosystems in the Arctic. Quantifying responses of CO2 exchange to warming in these ecosystems will require coordinated experimentation using standard temperature manipula- tions and measurements. Here, we used the International Tundra Experiment (ITEX) standard warming treatment to determine CO2 flux responses to growing-season warming for ecosystems spanning natural temperature and moisture ranges across the Arctic biome. We used the four North American Arctic ITEX sites (Toolik Lake, Atqasuk, and Barrow (USA) and Alexandra Fiord (Canada)) that span 108 of latitude. At each site, we investigated the CO2 responses to warming in both dry and wet or moist ecosystems. Net ecosystem CO2 exchange (NEE), ecosystem respiration (ER), and gross ecosystem photosynthesis (GEP) were assessed using chamber techniques conducted over 24-h periods sampled regularly throughout the summers of two years at all sites. At Toolik Lake, warming increased net CO2 losses in both moist and dry ecosystems. In contrast, at Atqasuk and Barrow, warming increased net CO2 uptake in wet ecosystems but increased losses from dry ecosystems. At Alexandra Fiord, warming improved net carbon uptake in the moist ecosystem in both years, but in the wet and dry ecosystems uptake increased in one year and decreased the other. Warming generally increased ER, with the largest increases in dry ecosystems. In wet ecosystems, high soil moisture limited increases in respiration relative to increases in photosynthesis. Warming generally increased GEP, with the notable exception of the Toolik Lake moist ecosystem, where warming unexpectedly decreased GEP .25%. Overall, the respiration response determined the effect of warming on ecosystem CO2 balance. Our results provide the first multiple-site comparison of arctic tundra CO2 flux responses to standard warming treatments across a large climate gradient. These results indicate that (1) dry tundra may be initially the most responsive ecosystems to climate warming by virtue of strong increases in ER, (2) moist and wet tundra responses are dampened by higher water tables and soil water contents, and (3) both GEP and ER are responsive to climate warming, but the magnitudes and directions are ecosystem-dependent.

The goal of this study is to determine if the response of arctic plants to warming is consistent across species, locations and time. This study examined the impact of experimental warming and natural temperature variation on plants at Barrow and Atqasuk, Alaska beginning in 1994. We considered observations of plant performance collected from 1994-2000 "short-term" and those from 2007-2012 "long-term". The plant traits reported are the number of inflorescences, inflorescence height, leaf length, and day of flower emergence. These traits can inform us about larger scale processes such as plant reproductive effort, plant growth, and plant phenology, and therefore provide valuable insight into community dynamics, carbon uptake, and trophic interactions. We categorized traits of all species monitored at each site into temperature response types. We then compared response types across traits, plant growth forms, sites, and over time to analyze the consistency of plant response to warming. Graminoids were the most responsive to warming and showed a positive response to temperature, while shrubs were generally the least responsive. Almost half (49%) of response types (across all traits, species, and sites combined) changed from short-term to long-term. The percent of plants responsive to warming decreased from 57% (short-term) to 46% (long-term). These results indicate that the response of plants to warming varies over time and has diminished overall in recent years.

This study is presented against the background that climate warming is predicted to continue in much of the Arctic through the next century and that small greenhouse chambers have been used widely to warm tundra communities in order to forecast climate-related changes. It reports results from up to 8 years of experimental warming with ∼1 m2 open-top chambers (OTCs) at four tundra communities near Barrow (71°18′N, 156°40′W) and Atqasuk (70°29′N, 157°25′W) in northern Alaska. Between 1994 and 2002 the OTCs increased the mean growing season air temperature by between 0.6 and 2.2°C, depending on the site and year. The change in average July soil temperature recorded over 3 years at 10 cm depth due to the OTCs varied between −0.8 and 0.7°C, depending on the site. Changes in soil temperature did not result in detectable differences in thaw depth at any site. This is interpreted to be the result of the small size of the OTCs and possibly changes in vegetation. The differences in warming profiles between...

Inference about future climate change impacts typically relies on one of three approaches: manipulative experiments, historical comparisons (broadly defined to include monitoring the response to ambient climate fluctuations using repeat sampling of plots, dendroecology, and paleoecology techniques), and space-for-time substitutions derived from sampling along environmental gradients. Potential limitations of all three approaches are recognized. Here we address the congruence among these three main approaches by comparing the degree to which tundra plant community composition changes (i) in response to in situ experimental warming, (ii) with interannual variability in summer temperature within sites, and (iii) over spatial gradients in summer temperature. We analyzed changes in plant community composition from repeat sampling (85 plant communities in 28 regions) and experimental warming studies (28 experiments in 14 regions) throughout arctic and alpine North America and Europe. Increases in the relative abundance of species with a warmer thermal niche were observed in response to warmer summer temperatures using all three methods; however, effect sizes were greater over broad-scale spatial gradients relative to either temporal variability in summer temperature within a site or summer temperature increases induced by experimental warming. The effect sizes for change over time within a site and with experimental warming were nearly identical. These results support the view that inferences based on space-for-time substitution overestimate the magnitude of responses to contemporary climate warming, because spatial gradients reflect long-term processes. In contrast, in situ experimental warming and monitoring approaches yield consistent estimates of the magnitude of response of plant communities to climate warming.

The Arctic climate is changing. Permafrost is warming, hydrological processes are chang- ing and biological and social systems are also evolving in response to these changing conditions. Knowing how the structure and function of arctic terrestrial ecosystems are responding to recent and persistent climate change is paramount to understanding the future state of the Earth system and how humans will need to adapt. Our holistic review presents a broad array of evidence that illustrates con- vincingly; the Arctic is undergoing a system-wide response to an altered climatic state. New extreme and seasonal surface climatic conditions are being experienced, a range of biophysical states and pro- cesses influenced by the threshold and phase change of freezing point are being altered, hydrological and biogeochemical cycles are shifting, and more regularly human sub-systems are being affected. Importantly, the patterns, magnitude and mechanisms of change have sometimes been unpredictable or difficult to isolate due to compounding factors. In almost every discipline represented, we show

The response of plants to temperature has gained renewed interest as researchers speculate on the biotic response to climate change. It is of particular interest in the Arctic, due to recent warming trends and anticipated continued warming for the region. This long-term, multispecies study confirms that changes in temperature affect the functioning of plants in their natural environment. It also demonstrates that the influence of temperature should be considered in the context of natural variability within a given location. The study examined natural temperature gradients, interannual climate variation, and experimental warming at sites near Barrow (71°18′ N, 156°40′ W) and Atqasuk (70°29′ N, 157°25′ W) in northern Alaska, USA. At each of the four sites, 24 plots were experimentally warmed for 5–7 years with small, open-top chambers, and plant growth and phenology were monitored; an equal number of unmanipulated control plots were monitored. The response of seven traits from 32 plant species occurring in ...