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Although many studies have examined the phenological mismatches between interacting organisms, few have addressed the potential for mismatches between phenology and seasonal weather conditions. In the Arctic, rapid phenological changes in many taxa are occurring in association with earlier snowmelt. The timing of snowmelt is jointly affected by the size of the late winter snowpack and the temperature during the spring thaw. Increased winter snowpack results in delayed snowmelt, whereas higher air temperatures and faster snowmelt advance the timing of snowmelt. Where interannual variation in snowpack is substantial, changes in the timing of snowmelt can be largely uncoupled from changes in air temperature. Using detailed, long-term data on the flowering phenology of four arctic plant species from Zackenberg, Greenland, we investigate whether there is a phenological component to the temperature conditions experienced prior to and during flowering. In particular, we assess the role of timing of flowering in determining pre-flowering exposure to freezing temperatures and to the temperatures experienced prior to flowering. We then examine the implications of flowering phenology for flower abundance. Earlier snowmelt resulted in greater exposure to freezing conditions, suggesting an increased potential for a mismatch between the timing of flowering and seasonal weather conditions and an increased potential for negative consequences, such as freezing damage. We also found a parabolic relationship between the timing of flowering and the temperature experienced during flowering after taking interannual temperature effects into account. If timing of flowering advances to a cooler period of the growing season, this may moderate the effects of a general warming trend across years. Flower abundance was quadratically associated with the timing of flowering, such that both early and late flowering led to lower flower abundance than did intermediate flowering. Our results indicate that shifting the timing of flowering affects the temperature experienced during flower development and flowering beyond that imposed by interannual variations in climate. We also found that phenological timing may affect flower abundance, and hence, fitness. These findings suggest that plant population responses to future climate change will be shaped not only by extrinsic climate forcing, but also by species' phenological responses. [ABSTRACT FROM AUTHOR]

The position of the Arctic treeline, which is a key regulator of surface energy exchange and carbon cycling, is widely thought to be controlled by temperature. Here, we present evidence that soil nutrient availability, rather than temperature, may be the proximate control on growth of treeline trees at our study site in northwest Alaska. We examined constraints on growth and allocation of white spruce in three contrasting habitats. The habitats had similar aboveground climates, but soil temperature declined from the riverside terrace to the forest to the treeline. We identified six lines of evidence that conflict with the hypothesis of direct temperature control and/or point to the importance of soil nutrient availability. First, the magnitude of aboveground growth declined from the terrace to the forest to the treeline, along gradients of diminishing soil nitrogen (N) availability and needle N concentration. Second, peak rates of branch extension, main stem radial and fine-root growth were generally not coincident with seasonal air and soil temperature maxima. At the treeline, in particular, rates of aboveground and fine-root growth declined well before air and soil temperatures reached their seasonal peaks. Third, in contrast with the hypothesis of temperature-limited growth, growing season average net photosynthesis was positively related to the sum of normalized branch extension, main stem radial and fine-root growth across trees and sites. Fourth, needle nonstructural carbohydrate concentration was significantly higher on the terrace, where growth was greatest. Fifth, annual branch extension growth was positively related to snow depth, consistent with the hypothesis that deeper snow promotes microbial activity and greater soil nutrient availability. Finally, the tree ring record revealed a large growth increase during late 20th-century climate warming on the terrace, where soil N availability is relatively high. Meanwhile, trees in the forest and at the treeline showed progressively smaller growth increases. Our results suggest temperature effects on tree growth at our study sites may be mediated by soil nutrient availability, making responses to climate change more complex and our ability to interpret the tree ring record more challenging than previously thought. [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. [ABSTRACT FROM AUTHOR]