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Much is still unknown about the growth and physiological responses of trees to global change at the northern treeline. We combined tree‐ring width data with century‐long stable carbon and oxygen isotope records to investigate growth and physiological responses of white spruce at two treeline sites in the Canadian Arctic to concurrent increases in temperature, atmospheric CO2 concentration (ca), and decline in sea ice extent over the past century. The tree‐ring records were assessed during three periods with contrasting climatic conditions: (a) the early 20th century warming, (b) the 1940–1970 cooling period, and (c) the anthropogenic late 20th century warming period. We found opposing growth trends between the two sites, but similar carbon isotope discrimination (Δ13C) and intrinsic water‐use efficiency (iWUE) trajectories. While tree growth (defined as basal area increment) increased at the site nearer to the Arctic Ocean during the 20th century following the rise in temperature and sea ice loss, growth declined after 1950 at the more interior site. At both sites, Δ13C slightly increased over these periods. However, trees showed a nonlinear response to increased ca, shifting after 1970 from a passive stomatal response (i.e., no changes in iWUE) to an active response (i.e., a moderate ∼12% increase in iWUE). Further, our isotope‐based findings do not support the idea that temperature‐induced drought stress caused the divergent growth trends at our treeline sites. This study thus highlights nonlinear and complex physiological and growth adjustments to concomitant changes in temperature, sea ice extent, and ca over the last century at the northern treeline. Plain Language Summary: In the last decades, Arctic ecosystems have experienced unprecedented warmer temperatures, higher atmospheric carbon dioxide concentrations, and a decline in sea ice extent. However, it is still unknown how these changes in environmental factors have impacted tree populations at their northern distribution limits. Here, we examine how the growth and water‐use efficiency of white spruce trees have changed over the last century in response to warmer temperatures, higher carbon dioxide concentrations, and sea ice decline at two treeline sites in the Canadian Arctic. We found that tree growth increased at the site nearer to the Arctic Ocean during the 20th century following the rise in temperature and sea ice loss, while growth declined after 1950 at the more interior site. At both sites, the water‐use efficiency of trees was stable until 1970. Following the rapid temperature and carbon dioxide increase since the 1970s, trees have become more efficient in using water. The contrasting growth trends found at our study sites highlight complex growth and physiological adjustments of white spruce trees to rapid and simultaneous changes in temperature, sea ice extent, and carbon dioxide concentrations in the Arctic. Key Points: Complex growth and physiological responses of white spruce trees to changes in temperature and sea ice extent inferred from stable isotopesWhite spruce radial growth at Arctic latitudinal treeline decreased at an interior site and increased at a site closer to the Arctic OceanWhite spruce shifted from a passive to an active water‐use efficiency response to rising CO2 concentration over the 20th century [ABSTRACT FROM AUTHOR]

The distribution and composition of Arctic vegetation are expected to shift with ongoing climate change. Global models generally predict northward shifts in high‐latitude ecotones, and analysis of remote sensing data shows widespread greening and changes in vegetation structure across the circumpolar Arctic. However, there are still uncertainties related to the timing of these shifts and variation among different plant functional types. In this paper, we investigate disequilibrium dynamics of green alder and white spruce in the Tuktoyaktuk Coastal Plain, NWT. We used high‐resolution air photographs captured in the 1970s and 2000s to quantify changes in the distribution and abundance of alder and spruce near their northern limits. We found increases in alder and spruce stem density over time, but no change in their range limits, indicating that both species are affected by leading‐edge disequilibrium. Low stand density and temperature limitation of reproduction along the northern margin likely contributed to observed disequilibrium in both species. We also observed the greatest change in species occupancy within a burned area, suggesting that the increased frequency of fire will play a significant role in the timing and magnitude of near‐term vegetation change. [ABSTRACT FROM AUTHOR]

Climate change is expected to increase woody vegetation abundance in the Arctic, yet the magnitude, spatial pattern and pathways of change remain uncertain. We compared historical orthophotos photos (1952 and 1979) with high‐resolution satellite imagery (2015) to examine six decades of change in abundance of white spruce Picea glauca and tall shrubs (Salix spp., Alnus spp.) near the Agashashok River in northwest Alaska. We established ~3000 random points within our ~5500 ha study area for classification into nine cover types. To examine physiographic controls on tree abundance, we fit multinomial log‐linear models with predictors derived from a digital elevation model and with arctic tundra, alpine tundra and 'tree' as levels of a categorical response variable. Between 1952 and 2015, points classified as arctic and alpine tundra decreased by 31% and 15%, respectively. Meanwhile, tall shrubs increased by 86%, trees mixed with tall shrubs increased by 385% and forest increased by 84%. Tundra with tall shrubs rarely transitioned to forest. The best multinomial model explained 71% of variation in cover and included elevation, slope and an interaction between slope and 'northness'. Treeline was defined as the elevation where the probability of tree presence equaled that of tundra. Mean treeline elevation in 2015 was 202 m, corresponding with a June–August mean air temperature > 11°C, which is > 4°C warmer than the 6–7°C isotherm associated with global treeline elevations. Our results show dramatic increases in the abundance of trees and tall shrubs, question the universality of air temperature as a predictor of treeline elevation and suggest two mutually exclusive pathways of vegetation change, because tundra that gained tall shrubs rarely transitioned to forest. Conversion of tundra to tall shrubs and forest has important and potentially contrasting implications for carbon cycling, surface energy exchange and wildlife habitat in the Arctic. [ABSTRACT FROM AUTHOR]

Trees growing near the Arctic treeline have long been used to reconstruct past climates. However, recent studies have shown deterioration of historically strong positive correlations between air temperature and tree growth (known as "divergence"). Divergence has important implications for confidence in paleoclimate reconstructions and ecosystem-atmosphere carbon exchange. Studies in the Brooks Range of northern Alaska showed that white spruce in the west increased growth in response to late 20th century warming, whereas those in the east failed to show a growth increase. In an earlier study across four watersheds in the Brooks Range, we tested and rejected the hypothesis that divergence in the easternmost watershed reflects moisture limitation of growth. Here, using 16 sites distributed across the same four watersheds, we tested an alternative hypothesis, that greater nutrient limitation in the east may have impeded positive growth responses to warming. Climate comparison across the four Brooks Range study watersheds revealed that, although the easternmost watershed generally had a drier growing-season climate, the most consistent difference was that winter air temperature and both winter and summer soil temperatures were much colder in the central and eastern watersheds. Soil nutrient availability, foliar nutrient concentrations, and tree growth were all generally lower in the central and eastern than in the western watersheds. Foliar phosphorus concentration was the best predictor of spatial variation in branch extension growth-a finding that is somewhat inconsistent with the theory that forest productivity on young, glacially derived soils should be strongly nitrogen limited. Experimental fertilization yielded the greatest growth increase in the eastern, an intermediate response in the central, and the smallest growth increase in the western watershed, generally mirroring trends in soil temperature, soil nutrient availability, foliar nutrient concentrations, and growth of control trees. Our results confirm that growth in the easternmost watershed is more nutrient limited and suggest that phosphorus limitation may be at least as important as nitrogen limitation of growth. We hypothesize that cold soil effects on tree access to nutrients might explain divergence in the eastern Brooks Range and elsewhere near the Arctic treeline, particularly in areas with cold winters and widespread permafrost., (© 2019 by the Ecological Society of America.)