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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.

Open top chambers (OTCs) were adopted as the recommended warming mechanism by the International Tundra Experiment (ITEX) network in the early 1990’s. 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.

Wind damage from cyclones can devastate the forest canopy, altering environmental conditions in the understory that affect seedling growth and plant community regeneration. To investigate the impact of hurricane‐induced increases in light and soil nutrients as a result of canopy defoliation, we conducted a two‐way factorial light and nutrient manipulation in a shadehouse experiment. We measured seedling growth of the dominant canopy species in the four Everglades forest communities: pine rocklands (Pinus elliottii var densa), cypress domes (Taxodium distichum), hardwood hammocks, and tree islands (Quercus virginiana and Bursera simaruba). Light levels were full sun and 50% shade, and nutrient levels coupled with an additional set of individuals that were subjected to a treatment mimicking the sudden effects of canopy opening from hurricane‐induced defoliation and the corresponding nutrient pulse. Seedlings were measured weekly for height growth and photosynthesis, with seedlings being harvested after 16 weeks for biomass, leaf area, and leaf tissue N and 13C isotope ratio. Growth rates and biomass accumulation responded more to differences in soil nutrients than differences in light availability, with largest individuals being in the high nutrient treatments. For B. simaruba and P. elliottii, the highest photosynthetic rates occurred in the high light, high nutrient treatment, while T. distichum and Q. virginiana photosynthetic rates were highest in low light, high nutrient treatment. Tissue biomass allocation patterns remained similar across treatments, except for Q. virginiana, which altered above‐ and belowground biomass allocation to increase capture of limiting soil and light resources. In response to the hurricane simulation treatment, height growth increased rapidly for Q. virginiana and B. simaruba, with nonsignificant increases for the other two species. We show here that ultimately, hurricane‐adapted, tropical species may be more likely to recolonize the forest canopy following a large‐scale hurricane disturbance. Wind damage associated with cyclones can devastate the forest canopy, strongly altering environmental conditions in the understory that affect seedling growth and canopy regeneration. To investigate the impact of hurricane‐induced increases in light and soil nutrients as a result of canopy defoliation, we conducted a two‐way factorial light and nutrient manipulation in a shadehouse. Ultimately, the ability of a species to recolonize hurricane damaged canopies may shift community dominance to more hurricane‐adapted species.

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.

Foundation species have disproportionately large impacts on ecosystem structure and function. As a result, future changes to their distribution may be important determinants of ecosystem carbon (C) cycling in a warmer world. We assessed the role of a foundation tussock sedge ( Eriophorum vaginatum ) as a climatically vulnerable C stock using field data, a machine learning ecological niche model, and an ensemble of terrestrial biosphere models (TBMs). Field data indicated that tussock density has decreased by ∼0.97 tussocks per m 2 over the past ∼38 years on Alaska’s North Slope from ∼1981 to 2019. This declining trend is concerning because tussocks are a large Arctic C stock, which enhances soil organic layer C stocks by 6.9% on average and represents 745 Tg C across our study area. By 2100, we project that changes in tussock density may decrease the tussock C stock by 41% in regions where tussocks are currently abundant (e.g. −0.8 tussocks per m 2 and −85 Tg C on the North Slope) and may increase the tussock C stock by 46% in regions where tussocks are currently scarce (e.g. +0.9 tussocks per m 2 and +81 Tg C on Victoria Island). These climate-induced changes to the tussock C stock were comparable to, but sometimes opposite in sign, to vegetation C stock changes predicted by an ensemble of TBMs. Our results illustrate the important role of tussocks as a foundation species in determining future Arctic C stocks and highlight the need for better representation of this species in TBMs.

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.

Plant phenological and growth responses to experimental warming are widely documented, but less is known about warming effects on plant–pollinator interactions. We investigated the effects of short- and long-term passive warming on flowering phenology, insect visitation, fruit production, and floral rewards in the Low Arctic in northern Alaska. To better understand the role of insect visitors in plant reproductive success, we quantified pollen loads on floral visitors and tested for pollen limitation in four species. Long-term warming advanced flowering onset in evergreen shrubs and forbs. Warming, in general, increased the duration of flowering for forbs, evergreen shrubs, and deciduous shrubs. Considering all growth forms together, long-term warming increased floral density. This pattern was primarily driven by deciduous and evergreen shrubs. Dipterans accounted for more visits than Hymenopterans, although Hymenopterans had higher pollen loads. Insect exclusion and warming decreased fruit set in the forb, Bistorta officinalis Delarbre. Nectar volume in the deciduous shrub, Vaccinium uliginosum, was higher in the warmed plots than the control, but nectar quality did not differ. Advanced flowering onset, longer flowering duration, and increased flower density and nectar volume may have important implications for the pollinator community, warranting further research on long-term warming effects on tundra ecosystems.

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.

Satellite data suggest that contemporary climate warming has already resulted in increased productivity and shrub biomass over much of the Arctic, but plot-level evidence for vegetation transformation remains sparse. Now research provides plot-scale evidence linking changes in vascular plant abundance to local summer warming in widely dispersed tundra locations across the globe. Temperature is increasing at unprecedented rates across most of the tundra biome 1 . Remote-sensing data indicate that contemporary climate warming has already resulted in increased productivity over much of the Arctic 2 , 3 , 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.

Climate change is leading to species redistributions. In the tundra biome,shrubs are generally expanding, but not all tundra shrub species will benefitfrom warming. Winner and loser species, and the characteristics that maydetermine success or failure, have not yet been fully identified. Here, weinvestigate whether past abundance changes, current range sizes and projectedrange shifts derived from species distribution models are related toplant trait values and intraspecific trait variation. We combined 17,921 traitrecords with observed past and modelled future distributions from 62 tundrashrub species across three continents. We found that species with greatervariation in seed mass and specific leaf area had larger projected range shifts,and projected winner species had greater seed mass values. However, traitvalues and variation were not consistently related to current and projectedranges, nor to past abundance change. Overall, our findings indicate thatabundance change and range shiftswill not lead to directionalmodifications inshrub trait composition, since winner and loser species share relatively similartrait spaces.