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  1. Global climate change has resulted in geographic range shifts of flora and fauna at a global scale. Extreme environments, like the Arctic, are seeing some of the most pronounced changes. This region covers 14% of the Earth’s land area, and while many arctic species are widespread, understanding ecotypic variation at the genomic level will be important for elucidating how range shifts will affect ecological processes. Tussock cottongrass ( Eriophorum vaginatum L.) is a foundation species of the moist acidic tundra, whose potential decline due to competition from shrubs may affect ecosystem stability in the Arctic. We used double-digest Restriction Site-Associated DNA sequencing to identify genomic variation in 273 individuals of E. vaginatum from 17 sites along a latitudinal gradient in north central Alaska. These sites have been part of 30 + years of ecological research and are inclusive of a region that was part of the Beringian refugium. The data analyses included genomic population structure, demographic models, and genotype by environment association. Genome-wide SNP investigation revealed environmentally associated variation and population structure across the sampled range of E. vaginatum , including a genetic break between populations north and south of treeline. This structure is likely the result of subrefugial isolation, contemporary isolation by resistance, and adaptation. Forty-five candidate loci were identified with genotype-environment association (GEA) analyses, with most identified genes related to abiotic stress. Our results support a hypothesis of limited gene flow based on spatial and environmental factors for E. vaginatum , which in combination with life history traits could limit range expansion of southern ecotypes northward as the tundra warms. This has implications for lower competitive attributes of northern plants of this foundation species likely resulting in changes in ecosystem productivity. 
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  2. Abstract

    The response of plant leaf and root phenology and biomass in the Arctic to global change remains unclear due to the lack of synchronous measurements of above- and belowground parts. Our objective was to determine the phenological dynamics of the above- and belowground parts of Eriophorum vaginatum in the Arctic and its response to warming. We established a common garden located at Toolik Lake Field Station; tussocks of E. vaginatum from three locations, Coldfoot, Toolik Lake and Sagwon, were transplanted into the common garden. Control and warming treatments for E. vaginatum were set up at the Toolik Lake during the growing seasons of 2016 and 2017. Digital cameras, a handheld sensor and minirhizotrons were used to simultaneously observe leaf greenness, normalized difference vegetation index and root length dynamics, respectively. Leaf and root growth rates of E. vaginatum were asynchronous such that the timing of maximal leaf growth (mid-July) was about 28 days earlier than that of root growth. Warming of air temperature by 1 °C delayed the timing of leaf senescence and thus prolonged the growing season, but the temperature increase had no significant effect on root phenology. The seasonal dynamics of leaf biomass were affected by air temperature, whereas root biomass was correlated with soil thaw depth. Therefore, we suggest that leaf and root components should be considered comprehensively when using carbon and nutrient cycle models, as above- and belowground productivity and functional traits may have a different response to climate warming.

     
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  3. Summary

    Some rhizomatous grass and sedge species form tussocks that impact ecosystem structure and function. Despite their importance, tussock development and size controls are poorly understood due to the decadal to centennial timescales over which tussocks form.

    We explored mechanisms regulating tussock development and size in a ubiquitous arctic tussock sedge (Eriophorum vaginatum) using field observations and a mass balance model coupled with a tiller population model. Model–data fusion was used to quantify parameter and prediction uncertainty, determine model sensitivity, and test hypotheses on the factors regulating tussock size.

    The model accurately captured the dynamics of tussock development, characteristics, and size observed in the field. Tussock growth approached maximal size within several decades, which was determined by feedbacks between the mass balance of tussock root necromass and density‐dependent tillering. The model also predicted that maximal tussock size was primarily regulated by tiller root productivity and necromass bulk density and less so by tiller demography. These predictions were corroborated by field observations of tussock biomass and root characteristics.

    The study highlights the importance of belowground processes in regulating tussock development and size and enhances our understanding of the influence of tussocks on arctic ecosystem structure and function.

     
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  4. 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. 
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  5. Leaf toughness is an important functional trait that confers resistance to herbivory and mechanical damage. We sought to determine how species composition, climate, seasonality, and nutrient availability influence leaf toughness in two types of tundra in northern Alaska. We measured leaf toughness as force to punch for 11 species of Arctic plants in tussock tundra and dry heath tundra at 17 sites distributed along a latitudinal gradient. Rubus chamaemorus L. and the graminoids occupied opposite ends of the leaf toughness spectrum, with Rubus chamaemorus requiring the least force to punch, whereas one of the graminoids, Eriophorum vaginatum L., required the most. Leaf toughness increased with mean summer temperature for Eriophorum vaginatum and Betula nana L., whereas it declined with warmer temperatures for the other species. Toughness of mature leaves of Eriophorum vaginatum did not vary through the growing season but declined significantly after senescence. Application of N and P fertilizer in an experimental site decreased leaf toughness in three species but had no effect on four others. Leaf toughness of four out of five species in dry heath was greater than for the same species in tussock tundra, but there was no difference in community-weighted mean toughness between tussock tundra and dry heath. 
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  6. The phenology of Arctic plants is an important determinant of the pattern of carbon uptake and may be highly sensitive to continued rapid climate change. Eriophorum vaginatum L. (Cyperaceae) has a disproportionate influence over ecosystem processes in moist acidic tundra, but it is unclear whether its growth and phenology will remain competitive in the future. We investigated whether northern tundra ecotypes of E. vaginatum could extend their growing season in response to direct warming and transplanting into southern ecosystems. At the same time, we examined whether southern ecotypes could adjust their growth patterns in order to thrive further north, should they disperse quickly enough. Detailed phenology measurements across three reciprocal transplant gardens over a 2-year period showed that some northern ecotypes were capable of growing for longer when conditions were favourable, but their biomass and growing season length was still shorter than those of the southern ecotype. Southern ecotypes retained large leaf length when transplanted north and mirrored the growing season length better than the others, mainly owing to immediate green-up after snowmelt. All ecotypes retained the same senescence timing, regardless of environment, indicating a strong genetic control. Eriophorum vaginatum may remain competitive in a warming world if southern ecotypes can migrate north. 
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  8. Abstract

    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 m2over 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 m2and −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 m2and +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.

     
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