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Abstract Tundra plants are widely considered to be constrained by cool growing conditions and short growing seasons. Furthermore, phenological development is generally predicted by daily heat sums calculated as growing degree days. Analyzing over a decade of seasonal flower counts of 23 plant species distributed across four plant communities, together with hourly canopy-temperature records, we show that the timing of flowering of many tundra plants are best predicted by a modified growing degree day model with a maximum temperature threshold. Threshold maximums are commonly employed in agriculture, but until recently have not been considered for natural ecosystems and to our knowledge have not been used for tundra plants. Estimated maximum temperature thresholds were found to be within the range of daily temperatures commonly experienced for many species, particularly for plants at the colder, high Arctic study site. These findings provide an explanation for why passive experimental warming—where moderate changes in mean daily temperatures are accompanied by larger changes in daily maximum temperatures—generally shifts plant phenology less than ambient warming. Our results also suggest that many plants adapted to extreme cold environments may have limits to their thermal responsiveness. 

Plot-level photography is an attractive time-saving alternative to field measurements for vegetation monitoring. However, widespread adoption of this technique relies on efficient workflows for post-processing images and the accuracy of the resulting products. Here, we estimated relative vegetation cover using both traditional field sampling methods (point frame) and semi-automated classification of photographs (plot-level photography) across thirty 1 m2 plots near Utqiaġvik, Alaska, from 2012 to 2021. Geographic object-based image analysis (GEOBIA) was applied to generate objects based on the three spectral bands (red, green, and blue) of the images. Five machine learning algorithms were then applied to classify the objects into vegetation groups, and random forest performed best (60.5% overall accuracy). Objects were reliably classified into the following classes: bryophytes, forbs, graminoids, litter, shadows, and standing dead. Deciduous shrubs and lichens were not reliably classified. Multinomial regression models were used to gauge if the cover estimates from plot-level photography could accurately predict the cover estimates from the point frame across space or time. Plot-level photography yielded useful estimates of vegetation cover for graminoids. However, the predictive performance varied both by vegetation class and whether it was being used to predict cover in new locations or change over time in previously sampled plots. These results suggest that plot-level photography may maximize the efficient use of time, funding, and available technology to monitor vegetation cover in the Arctic, but the accuracy of current semi-automated image analysis is not sufficient to detect small changes in cover. 

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. 

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. 

Vegetation change of the Arctic tundra due to global warming is a well-known process, but the implication for the belowground microbial communities, key in nutrient cycling and decomposition, is poorly understood. We characterized the fungal and bacterial abundances in litter and soil layers across 16 warming experimental sites at 12 circumpolar locations. We investigated the relationship between microbial abundances and nitrogen (N) and carbon (C) isotopic signatures, indicating shifts in microbial processes with warming. Microbial abundances were 2–3 orders of magnitude larger in litter than in soil. Local, site-dependent responses of microbial abundances were variable, and no general effect of warming was detected. The only generalizable trend across sites was a dependence between the warming response ratios and C:N ratio in controls, highlighting a legacy of the vegetation on the microbial response to warming. We detected a positive effect of warming on the litter mass and δ 15 N, which was linked to bacterial abundance under warmed conditions. This effect was stronger in experimental sites dominated by deciduous shrubs, suggesting an altered bacterial N-cycling with increased temperatures, mediated by the vegetation, and with possible consequences on ecosystem feedbacks to climate change. 

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. 

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 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. 

The International Tundra Experiment (ITEX) was founded in 1990 as a network of scientists studying responses of tundra ecosystems to ambient and experimental climate change at Arctic and alpine sites across the globe. Common measurement and experimental design protocols have facilitated synthesis of results across sites to gain biome-wide insights of climate change impacts on tundra. This special issue presents results from more than 30 years of ITEX research. The importance of snow regimes, bryophytes, and herbivory are highlighted, with new protocols and studies proposed. The increasing frequency and magnitude of extreme climate events is shown to have strong effects on plant reproduction. The most consistent plant trait response across sites is an increase in vegetation height, especially for shrubs. This will affect surface energy balance, carbon and nutrient dynamics and trophic level interactions. Common garden studies show adaptation responses in tundra species to climate change but they are species and regionally specific. Recommendations are made including establishing sites near northern communities to increase reciprocal engagement with local knowledge holders and establishing multi-factor experiments. The success of ITEX is based on collegial cooperation among researchers and the network remains focused on documenting and understanding impacts of environmental change on tundra ecosystems. 

Abstract 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.