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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]

Aim: Altitudinal and latitudinal treeline ecotones have not consistently responded to climate warming in the direction and/or magnitude predicted by climate alone, suggesting that non‐climatic mechanisms (e.g. biotic interactions) also mediate treeline range dynamics. Through a collaborative research approach, we assessed environmental conditions associated with pre‐dispersal insect cone granivory and how this biotic interaction may govern the reproductive potential, and therefore range dynamics, of spruce‐dominated treelines across northern Canada. Location: In all, 10 boreal forest treelines, tundra and alpine, from Yukon to Newfoundland and Labrador, Canada. Taxa: White spruce (Picea glauca [Moench] Voss), Black spruce (Picea mariana [Mill.] B.S.P.), Strobilomyia spp., Megastigmus spp. Methods: Treeline sites were assessed for presence and magnitude of pre‐dispersal seed granivory by insects along with viable seed availability. We quantified stand density metrics, organic layer depth and understorey vegetation composition at each location and, subsequently, incorporated those variables into generalized linear mixed models to establish predictors of granivory magnitude and viability of available seed. Results: Insect granivory was widespread across sites; however, site‐specific patterns of granivory were associated with increased moss cover and decreased shrub cover and stand density. While all black spruce‐dominated sites exhibited seed viability rates > 50%, the number of seeds produced per cone varied, suggesting that within‐site abiotic conditions and biotic interaction pressures limit successful colonization of novel environments in advance of seed dispersal. Main Conclusions: The modelled relationships between granivory, seed viability and environmental conditions represent an essential step towards generalizing how and when biotic interactions across subarctic treelines influence boreal tree range dynamics before seed dispersal. Connections between granivory magnitude and site‐level treeline characteristics (e.g. stand density, understorey vegetation) will provide a more comprehensive understanding of treeline range dynamics under continued climate change. [ABSTRACT FROM AUTHOR]

Climate change is increasing the frequency and intensity of drought events in many boreal forests. Trees are sessile organisms with a long generation time, which makes them vulnerable to fast climate change and hinders fast adaptations. Therefore, it is important to know how forests cope with drought stress and to explore the genetic basis of these reactions. We investigated three natural populations of white spruce (Picea glauca) in Alaska, located at one drought-limited and two cold-limited treelines with a paired plot design of one forest and one treeline plot. We obtained individual increment cores from 458 trees and climate data to assess dendrophenotypes, in particular the growth reaction to drought stress. To explore the genetic basis of these dendrophenotypes, we genotyped the individual trees at 3000 single nucleotide polymorphisms in candidate genes and performed genotype-phenotype association analysis using linear mixed models and Bayesian sparse linear mixed models. Growth reaction to drought stress differed in contrasting treeline populations. Therefore, the populations are likely to be unevenly affected by climate change. We identified 40 genes associated with dendrophenotypic traits that differed among the treeline populations. Most genes were identified in the drought-limited site, indicating comparatively strong selection pressure of drought-tolerant phenotypes. Contrasting patterns of drought-associated genes among sampled sites and in comparison to Canadian populations in a previous study suggest that drought adaptation acts on a local scale. Our results highlight genes that are associated with wood traits which in turn are critical for the establishment and persistence of future forests under climate change. [ABSTRACT FROM AUTHOR]

Recent climate warming is expected to increase tree growth and productivity, substantially altering ecological function and boundaries in northern ecosystems. Although climate largely determines the range and growth of trees in any biome, variations in microsite conditions can influence growth on a finer scale. Red foxes (Vulpes vulpes) are found in most terrestrial ecosystems and are considered ecosystem engineers through their denning activities. Added soil nutrients from prey remains, feces, and urine could benefit tree growth on dens in subarctic regions by alleviating soil nutrient limitations. We examined growth in white spruce (Picea glauca) trees growing on eight red fox dens and paired control sites at the Arctic treeline. Radial growth was 55% higher for trees on dens than on control sites between 1897 and 2017, despite similarities in tree ages and densities. This enhanced growth was more pronounced in younger trees, consistent with the decrease in growth with age among all trees. By promoting tree growth near the treeline, red foxes may act as hotspots for tree reproduction. Although the impacts on tree growth largely depend on the spatial distribution of dens, predators can create distinct microhabitats across the landscape, leading to increased vegetation productivity, persisting over many decades. [ABSTRACT FROM AUTHOR]

White spruce (Picea glauca) spans a massive range, yet the variability in respiratory physiology and related implications for tree carbon balance at the extremes of this distribution remain as enigmas. Working at both the most northern and southern extents of the distribution range more than 5000 km apart, we measured the short‐term temperature response of dark respiration (R/T) at upper and lower canopy positions. R/T curves were fit to both polynomial and thermodynamic models so that model parameters could be compared among locations, canopy positions, and with previously published data. Respiration measured at 25°C (R25) was 68% lower at the southern location than at the northern location, resulting in a significantly lower intercept in R/T response in temperate trees. Only at the southern location did upper canopy leaves have a steeper temperature response than lower canopy leaves, likely reflecting canopy gradients in light. At the northern range limit respiration is nearly twice that of the average R25 reported in a global leaf respiration database. We predict that without significant thermal acclimation, respiration will increase with projected end‐of‐the‐century warming and will likely constrain the future range limits of this important boreal species. Summary Statement: White spruce (Picea glauca) needle respiration at the northern limit of the species range is three times higher than at the southern range limit (when measured at 25°C). This high carbon cost likely challenges tree survival and contributes to the location of the northern treeline. [ABSTRACT FROM AUTHOR]

Arctic Treeline is the transition from the boreal forest to the treeless tundra and may be determined by growing season temperatures. The physiological mechanisms involved in determining the relationship between the physical and biological environment and the location of treeline are not fully understood. In Northern Alaska, we studied the relationship between temperature and leaf respiration in 36 white spruce (Picea glauca) trees, sampling both the upper and lower canopy, to test two research hypotheses. The first hypothesis is that upper canopy leaves, which are more directly coupled to the atmosphere, will experience more challenging environmental conditions and thus have higher respiration rates to facilitate metabolic function. The second hypothesis is that saplings [stems that are 5–10cm DBH (diameter at breast height)] will have higher respiration rates than trees (stems ≥10cm DBH) since saplings represent the transition from seedlings growing in the more favorable aerodynamic boundary layer, to trees which are fully coupled to the atmosphere but of sufficient size to persist. Respiration did not change with canopy position, however respiration at 25°C was 42% higher in saplings compared to trees (3.43±0.19 vs. 2.41±0.14μmolm−2 s−1). Furthermore, there were significant differences in the temperature response of respiration, and seedlings reached their maximum respiration rates at 59°C, more than two degrees higher than trees. Our results demonstrate that the respiratory characteristics of white spruce saplings at treeline impose a significant carbon cost that may contribute to their lack of perseverance beyond treeline. In the absence of thermal acclimation, the rate of leaf respiration could increase by 57% by the end of the century, posing further challenges to the ecology of this massive ecotone. [ABSTRACT FROM AUTHOR]

Treeline ecosystems are of great scientific interest to study the effects of limiting environmental conditions on tree growth. However, tree growth is multidimensional, with complex interactions between height and radial growth. In this study, we aimed to disentangle effects of height and climate on xylem anatomy of white spruce [ Picea glauca (Moench) Voss] at three treeline sites in Alaska; i.e., one warm and drought-limited, and two cold, temperature-limited. To analyze general growth differences between trees from different sites, we used data on annual ring width, diameter at breast height (DBH), and tree height. A representative subset of the samples was used to investigate xylem anatomical traits. We then used linear mixed-effects models to estimate the effects of height and climatic variables on our study traits. Our study showed that xylem anatomical traits in white spruce can be directly and indirectly controlled by environmental conditions: hydraulic-related traits seem to be mainly influenced by tree height, especially in the earlywood. Thus, they are indirectly driven by environmental conditions, through the environment's effects on tree height. Traits related to mechanical support show a direct response to environmental conditions, mainly temperature, especially in the latewood. These results highlight the importance of assessing tree growth in a multidimensional way by considering both direct and indirect effects of environmental forcing to better understand the complexity of tree growth responses to the environment. [ABSTRACT FROM AUTHOR]

The ecological function of boreal forests is challenged by drastically changing climate conditions. Although an increasing number of studies are investigating how climate change is influencing growth and distribution of boreal tree species, there is a lack of studies examining the potential of these species to genetically adapt or phenotypically adjust. Here, we sampled clonally and non-clonally growing white spruce trees (Picea glauca [Moench] Voss) to investigate spatial and genetic effects on tree ring width and on six xylem anatomical traits representing growth, water transport, mechanical support, and wood density. We compared different methods for estimating broad sense heritability (H2) of each trait and we evaluated the effects of spatial grouping and genetic grouping on the xylem anatomical traits with linear models. We found that the three different methods used to estimate H2 were quite robust, showing overall consistent patterns, while our analyses were unsuccessful at fully separating genetic from spatial effects. By evaluating the effect size, we found a significant effect of genetic grouping in latewood density and earlywood hydraulic diameter. However, evaluating model performances showed that spatial grouping was a better predictor than genetic grouping for variance in earlywood density, earlywood hydraulic diameter and growth. For cell wall thickness neither spatial nor genetic grouping was significant. Our findings imply that (1) the variance in the investigated xylem anatomical traits and growth is mainly influenced by spatial clustering (most probably caused by microhabitat conditions), which (2) makes it rather difficult to estimate the heritability of these traits in naturally grown trees in situ. Yet, (3) latewood density and earlywood hydraulic diameter qualified for further analysis on the genetic background of xylem traits and (4) cell wall thickness seems a useful trait to investigate large-scale climatic effects, decoupled from microclimatic, edaphic and genetic influences. [ABSTRACT FROM AUTHOR]

Tree growth at northern treelines is generally temperature‐limited due to cold and short growing seasons. However, temperature‐induced drought stress was repeatedly reported for certain regions of the boreal forest in northwestern North America, provoked by a significant increase in temperature and possibly reinforced by a regime shift of the pacific decadal oscillation (PDO). The aim of this study is to better understand physiological growth reactions of white spruce, a dominant species of the North American boreal forest, to PDO regime shifts using quantitative wood anatomy and traditional tree‐ring width (TRW) analysis. We investigated white spruce growth at latitudinal treeline across a >1,000 km gradient in northwestern North America. Functionally important xylem anatomical traits (lumen area, cell‐wall thickness, cell number) and TRW were correlated with the drought‐sensitive standardized precipitation–evapotranspiration index of the growing season. Correlations were computed separately for complete phases of the PDO in the 20th century, representing alternating warm/dry (1925–1946), cool/wet (1947–1976) and again warm/dry (1977–1998) climate regimes. Xylem anatomical traits revealed water‐limiting conditions in both warm/dry PDO regimes, while no or spatially contrasting associations were found for the cool/wet regime, indicating a moisture‐driven shift in growth‐limiting factors between PDO periods. TRW reflected only the last shift of 1976/1977, suggesting different climate thresholds and a higher sensitivity to moisture availability of xylem anatomical traits compared to TRW. This high sensitivity of xylem anatomical traits permits to identify first signs of moisture‐driven growth in treeline white spruce at an early stage, suggesting quantitative wood anatomy being a powerful tool to study climate change effects in the northwestern North American treeline ecotone. Projected temperature increase might challenge growth performance of white spruce as a key component of the North American boreal forest biome in the future, when drier conditions are likely to occur with higher frequency and intensity. [ABSTRACT FROM AUTHOR]

Climate-sensitive height–age models were developed for top height trees of trembling aspen (Populus tremuloides Michx.), jack pine (Pinus banksiana Lamb.), and white spruce (Picea glauca (Moench) Voss) in natural and reclaimed oil sands stands. We used stem analysis data collected from the Athabasca oil sands region in northern Alberta, Canada, and climate data generated by the ClimateWNA model. Height–age trajectories differed between top height trees in natural and reclaimed stands for jack pine and white spruce, but not for trembling aspen. At a given age, white spruce top height trees were taller and jack pine top height trees were shorter in reclaimed stands than those in natural stands, suggesting that it is easier to achieve similar forest productivity for oil sands sites reclaimed with white spruce stands than for sites reclaimed with jack pine stands. The principal climate variables were growing season (May to September) precipitation averaged over the previous 10 years for trembling aspen and jack pine and summer (June to August) precipitation averaged over the previous 10 years for white spruce. These variables had positive effects on the height–age trajectories. [ABSTRACT FROM AUTHOR]

Herbivores can modify the rate of shrub and treeline advance. Both direct and indirect effects of herbivory may simultaneously interact to affect the growth rates of plants at this ecotone. We investigated the effect of snowshoe hare herbivory on the height of white spruce at two treeline locations in Alaska, USA. White spruce is expanding its distribution both upwards in elevation and northward in latitude because of climate warming, and snowshoe hares are already present in areas likely to be colonized by spruce. We hypothesized that herbivory would result in browsed individuals having reduced height, suggesting herbivory is a direct, negative effect on spruce treeline advance. We found an interactive effect between browsing history and spruce age. When young (under 30 years old), individuals that were browsed tended to be taller than unbrowsed individuals. However, older seedlings (over 30 years old) that had been browsed were shorter than unbrowsed individuals of the same age. Hares suppress faster growing individuals that are initially taller by preferentially browsing them as they emerge above the winter snowpack. This reduced height, in combination with increased mortality associated with browsing, is predicted to slow the advance of both latitudinal and altitudinal treeline expansions and alter the structure of treeline forests. [ABSTRACT FROM AUTHOR]

Significant treeline advance can only occur with successful establishment, growth, and survival of new seedlings. Several studies have examined microsite factors at single locations to explain the presence/absence of seedlings at treeline. We conducted a much larger observational study and included multiple factors to determine (1) which variable(s) was/were most important, (2) whether their importance differed between aspects, and (3) whether the same variables explained variation in seedling growth and damage. We analyzed five biophysical and six shrub variables along four forest-tundra ecotones in southwest Yukon at 640 points. The model that best explained seedling occurrence was similar between north- and southfacing slopes. Of all variables, seedling occurrence was best explained by the proximity, height, and upslope orientation of shrubs (relative to the seedling). The data indicate an optimal range of shrub cover, which differed with aspect. On north-facing slopes, seedlings occurred most often when shrub cover exceeded 13%, while on south-facing slopes seedlings occurred most when shrub cover was between 9% and 72%.We also found that with the exception of shrub-related factors, very few biophysical variables explained size and growth characteristics and damage of tree seedlings, suggesting that the relative strength and importance of variables change depending on the life stage and size of the tree seedling. Collectively, our results demonstrate a non-random distribution of seedlings in the forest-tundra ecotone, suggesting that as shrub distributions change with climate change, colonization sites for seedlings will also be influenced. [ABSTRACT FROM AUTHOR]

Regional warming has led to increased productivity near the boreal forest margin in Alaska. To date, the effects of warming on seedling recruitment have received little attention, in spite of forecasted forest expansion. Here, we used stand structure and environmental data from 95 white spruce ( Picea glauca) plots sampled across a longitudinal gradient in southwest Alaska to explore factors influencing spruce establishment and recruitment near western treeline. We used total counts of live seedlings, saplings, and trees, representing five life stages, to evaluate whether geospatial, climate, and measured plot covariates predicted abundance, using current abundance distributions as a surrogate for climate conditions in the past. We used generalized linear models to test the null hypothesis that conditions favorable for recruitment were similar along the environmental gradient represented by longitude, by exploring relationships between per-plot counts of each life stage and the covariates hypothesized to affect abundance. We also examined the relationship between growing degree days ( GDD) and seedling establishment over a period of three decades using tree-ring chronologies obtained from cores taken at a subset of our sites ( n = 30). Our results indicated that seedling, sapling, and tree abundance were positively correlated with temperature across the study area. The response to longitude was mixed, with earlier life stages (seedlings, saplings) most abundant at the western end of the gradient, and later life stages (trees) most abundant to the east. The differential relationship between longitude and life-stage abundance suggests a moving front of white spruce establishment through time, driven by changes in environmental conditions near the species' western range limit. Likewise, we found a positive relationship between periods of seedling establishment and GDD, suggesting that longer summers and/or greater heat accumulation might enhance establishment, consistent with the positive relationship we found between life-stage abundance and temperature. [ABSTRACT FROM AUTHOR]

Treelines in Alaska are advancing in elevation and latitude because of climate warming, which is expanding the habitat available for boreal wildlife species, including snowshoe hares ( Lepus americanus). Snowshoe hares are already present in tall shrub communities beyond treeline and are the main browser of white spruce ( Picea glauca), the dominant tree species at treeline in Alaska. We investigated the processes involved in a 'snowshoe hare filter' to white spruce establishment near treeline in Denali National Park, Alaska, USA. We modeled the pattern of spruce establishment from 1970 to 2009 and found that fewer spruce established during periods of high hare abundance. Multiple factors interact to influence browsing of spruce, including the hare cycle, snow depth and the characteristics of surrounding vegetation. Hares are abundant at treeline and may exclude spruce from otherwise optimal establishment sites, particularly floodplain locations with closed shrub canopies. The expansion of white spruce treeline in response to warming climate will be strongly modified by the spatial and temporal dynamics of the snowshoe hare filter. [ABSTRACT FROM AUTHOR]

Summary: Plant‐associated mycobiomes in extreme habitats are understudied and poorly understood.We analysed Illumina‐generated ITS1 sequences from the needle mycobiome of white spruce (Picea glauca) at the northern treeline in Alaska (USA). Sequences were obtained from the same DNA that was used for tree genotyping. In the present study, fungal metabarcoding and tree microsatellite data were compared for the first time.In general, neighbouring trees shared more fungal taxa with each other than trees growing in further distance. Mycobiomes correlated strongly with phenological host traits and local habitat characteristics contrasting a dense forest stand with an open treeline site. Genetic similarity between trees did not influence fungal composition and no significant correlation existed between needle mycobiome and tree genotype.Our results suggest the pronounced influence of local habitat conditions and phenotypic tree traits on needle‐inhabiting fungi. By contrast, the tree genetic identity cannot be benchmarked as a dominant driver for needle‐inhabiting mycobiomes, at least not for white spruce in this extreme environment. [ABSTRACT FROM AUTHOR]

Aims: While treeline positions are globally correlated to growing season temperatures, seedling establishment, an important process of alpine treeline dynamics, is additionally controlled by regional-scale factors such as snow cover duration, desiccating winds and biotic interactions. Knowing that alpine treelines have shown contrasting responses to climate change, we determined the relative importance of key abiotic and biotic factors involved in seedling survival and growth. Location: McGerrigle Mountains, Parc National de la Gaspésie, Appalachian Range, eastern Quebec, Canada. Methods: In two white spruce (Picea glauca) treeline sites, we used the microclimate in the vicinity of tree islands, densely packed clusters of trees isolated from each other by alpine tundra vegetation, to assess the effects of abiotic variables (sum of degree days [DD], snowpack duration and a wind exposure index) as well as the effects of biotic interactions with neighbouring vegetation on the survival and growth of transplanted white spruce seedlings. For 3 yr, we surveyed seedling survival twice a year to discriminate between winter and summer survival, and measured seedling growth at the end of each growing season. We used Bayesian hierarchical models to estimate the relative effects of covariates on survival and growth. Results: Survival probability decreased in microsites wherewinter DD was high, and increased in microsites with longer snowpack duration. In wind-exposed microsites, seedling survival increased when neighbouring vegetation was present, indicating facilitative mechanisms. Seedling growth was positively affected by the duration of snow cover and tended to increase with higher DD during the previous year. In wind-sheltered microsites, seedling growth tended to be negatively affected by neighbouring vegetation, indicating competitive mechanisms. Conclusions: Our study demonstrates that seedling establishment is more sensitive to winter conditions, notably to the length of snow cover (which protects seedlings from frost and desiccation), than to summer temperature. Biotic interactions increased seedling establishment when environmental stresses were higher. We suggest that regional-scale factors such as winter climate and biotic interactions should be included inmodelling exercises to improve future treeline location forecasts. [ABSTRACT FROM AUTHOR]

Increment cores from the boreal forest have long been used to reconstruct past climates. However, in recent years, numerous studies have revealed a deterioration of the correlation between temperature and tree growth that is commonly referred to as divergence. In the Brooks Range of northern Alaska, USA, studies of white spruce ( Picea glauca) revealed that trees in the west generally showed positive growth trends, while trees in the central and eastern Brooks Range showed mixed and negative trends during late 20th century warming. The growing season climate of the eastern Brooks Range is thought to be drier than the west. On this basis, divergent tree growth in the eastern Brooks Range has been attributed to drought stress. To investigate the hypothesis that drought-induced stomatal closure can explain divergence in the Brooks Range, we synthesized all of the Brooks Range white spruce data available in the International Tree Ring Data Bank ( ITRDB) and collected increment cores from our primary sites in each of four watersheds along a west-to-east gradient near the Arctic treeline. For cores from our sites, we measured ring widths and calculated carbon isotope discrimination (δ13C), intrinsic water-use efficiency ( iWUE), and needle intercellular CO2 concentration ( Ci) from δ13C in tree-ring alpha-cellulose. We hypothesized that trees exhibiting divergence would show a corresponding decline in δ13C, a decline in Ci, and a strong increase in iWUE. Consistent with the ITRDB data, trees at our western and central sites generally showed an increase in the strength of the temperature-growth correlation during late 20th century warming, while trees at our eastern site showed strong divergence. Divergent tree growth was not, however, associated with declining δ13C. Meanwhile, estimates of Ci showed a strong increase at all of our study sites, indicating that more substrate was available for photosynthesis in the early 21st than in the early 20th century. Our results, which are corroborated by measurements of xylem sap flux density, needle gas exchange, and measurements of growth and δ13C along moisture gradients within each watershed, suggest that drought-induced stomatal closure is probably not the cause of 20th century divergence in the Brooks Range. [ABSTRACT FROM AUTHOR]

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]

Mast-seeding conifers such as Picea glauca exhibit synchronous production of large seed crops over wide areas, suggesting climate factors as possible triggers for episodic high seed production. Rapidly changing climatic conditions may thus alter the tempo and spatial pattern of masting of dominant species with potentially far-reaching ecological consequences. Understanding the future reproductive dynamics of ecosystems including boreal forests, which may be dominated by mast-seeding species, requires identifying the specific cues that drive variation in reproductive output across landscape gradients and among years. Here we used annual data collected at three sites spanning an elevation gradient in interior Alaska, USA between 1986 and 2011 to produce the first quantitative models for climate controls over both seedfall and seed viability in P. glauca, a dominant boreal conifer. We identified positive associations between seedfall and increased summer precipitation and decreased summer warmth in all years except for the year prior to seedfall. Seed viability showed a contrasting response, with positive correlations to summer warmth in all years analyzed except for one, and an especially positive response to warm and wet conditions in the seedfall year. Finally, we found substantial reductions in reproductive potential of P. glauca at high elevation due to significantly reduced seed viability there. Our results indicate that major variation in the reproductive potential of this species may occur in different landscape positions in response to warming, with decreasing reproductive success in areas prone to drought stress contrasted with increasing success in higher elevation areas currently limited by cool summer temperatures. [ABSTRACT FROM AUTHOR]

The prevailing hypothesis states that treeline positions are defined by the direct effects of cold temperatures on cell division and tree growth. Meanwhile, photosynthesis is thought to be relatively unrestricted in treeline trees. Support for this hypothesis comes from the global correlation between temperature and treeline position, the observation that many treelines have advanced in response to recent warming and the frequent use of increment cores from treeline trees to reconstruct past climates. However, studies of the physiology of trees at the Arctic treeline are rare, and this hypothesis remains largely untested., To improve our understanding of the relationships between temperature and performance of white spruce near the Arctic treeline, we made measurements of needle gas exchange, needle nutrition and soil nutrient availability over 2 years in three contrasting habitats: riverside terrace, hillslope forest and treeline. The sites had similar above-ground microclimates, but very different soil conditions. Soils were warm and dry on the terrace, cool and moist in the forest and cold and seasonally wet the treeline., Photosynthesis, needle nitrogen (N) concentration and soil N availability declined from the terrace to the forest to the treeline. Low N availability at the treeline was likely a consequence of limited microbial activity in the cold and seasonally wet soils. Soils at the treeline were colder than the terrace during the growing season and colder than the forest in winter, when the treeline maintains a shallow snowpack., Our results highlight the potential for an indirect effect of temperature on the growth of trees at the Arctic treeline and suggest that treeline responses to changes in climate may be more complex than previously thought. [ABSTRACT FROM AUTHOR]

Recent studies have revealed positive and negative trends in the radial growth of treeline white spruce (Picea glauca) as temperatures have warmed in recent decades. Investigators have speculated that negative growth trends reflect the increasing importance of temperature-induced drought stress, yet direct observations of drought-induced stomatal closure have not been made in white spruce near the Arctic treeline. In this study, we measured needle gas exchange, a variety of needle traits, and branch growth in contrasting growing seasons on a riverside terrace near the Arctic treeline in Noatak National Preserve, northwest Alaska. Needle gas exchange was limited by cold soils (<7 °C), nighttime frosts, large vapor pressure deficits (VPD) and/or low soil water contents during the majority of our midday measurements. Near optimal conditions for needle gas exchange were consistently found in late August, when soils were relatively warm, air temperatures were moderate, and the VPD was relatively small. Defoliation during a two-year bud moth infestation (Zeiraphera spp.) substantially reduced branch growth, obscured potential relationships between needle gas exchange and growth, and revealed the importance of whole canopy gas exchange measurements. Results of our study show there is a very narrow window of environmental conditions for near optimal needle gas exchange in white spruce near the Arctic treeline. Although we identified many abiotic constraints on needle gas exchange, a single biological factor likely had the greatest effect on annual branch growth. [ABSTRACT FROM AUTHOR]

Throughout interior Alaska (USA), a gradual warming trend in mean monthly temperatures occurred over the last few decades (∼2-4°°C). The accompanying increases in woody vegetation at many alpine treeline (hereafter treeline) locations provided an opportunity to examine how biotic and abiotic local site conditions interact to control tree establishment patterns during warming. We devised a landscape ecological approach to investigate these relationships at an undisturbed treeline in the Alaska Range. We identified treeline changes between 1953 (aerial photography) and 2005 (satellite imagery) in a geographic information system (GIS) and linked them with corresponding local site conditions derived from digital terrain data, ancillary climate data, and distance to 1953 trees. Logistic regressions enabled us to rank the importance of local site conditions in controlling tree establishment. We discovered a spatial transition in the importance of tree establishment controls. The biotic variable (proximity to 1953 trees) was the most important tree establishment predictor below the upper tree limit, providing evidence of response lags with the abiotic setting and suggesting that tree establishment is rarely in equilibrium with the physical environment or responding directly to warming. Elevation and winter sun exposure were important predictors of tree establishment at the upper tree limit, but proximity to trees persisted as an important tertiary predictor, indicating that tree establishment may achieve equilibrium with the physical environment. However, even here, influences from the biotic variable may obscure unequivocal correlations with the abiotic setting (including temperature). Future treeline expansion will likely be patchy and challenging to predict without considering the spatial variability of influences from biotic and abiotic local site conditions. [ABSTRACT FROM AUTHOR]

Aim The objectives of the study are: (1) to evaluate the dynamics of the maritime tree line and forest limit of white spruce, Picea glauca, within the dual framework of primary succession induced by the rapid post-glacial land emergence on the eastern coast of Hudson Bay and the impacts of recent and past climate changes; and (2) to determine the time lapse between land emergence and seedling, tree, and forest establishment in the context of the primary chronosequence occurring on rising, well-drained sandy beaches and terraces. Location The study area was located on the eastern coast of Hudson Bay (56°20′ N, 76°32′ W) in northern Québec, Canada. Methods We evaluated the colonization dynamics of white spruce as seedlings, tree-line trees and primary-forest trees at eight sites distributed along a 200-km latitudinal gradient based on a mean land emergence rate of 1.2 m century−1. A 30-m wide by 140–300-m long quadrat was positioned at random at the centre of each site. The elevation above sea level, position and age of all individuals of spruce present in the quadrat areas were determined, and the soils of each chronosequence were described. Results The main stages of primary succession along the emerging coast were common to all the sites, regardless of latitude, but occurred at different elevations above sea level (a.s.l.). White spruce seedlings colonized near-shore beaches 2 m a.s.l., whereas the tree line and forest limit tended to form only at about 3–4 m and 4–8 m a.s.l., corresponding approximately to 180–825 years and 310–1615 years after land emersion, respectively. White spruce establishment at the tree line occurred about 50 years ago. Climatic conditions at this time were probably more favourable to tree colonization than when the species established at the forest limit. Soil formation was influenced primarily by distance from the seashore and elevation above sea level, with podzolization being accelerated by white spruce cover. Main conclusions The current tree-line and forest-limit positions on the rising coast of eastern Hudson Bay correspond to ecological limits established during the course of primary succession within a context of changing climatic conditions. The recent establishment of trees at the tree line and forest limit at relatively old coastal sites is associated with warmer conditions over the last 100 years. Although white spruce was present nearby, coastal sites were devoid of trees before the 20th century. [ABSTRACT FROM AUTHOR]

1 Boundaries between forest and tundra ecosystems, tree lines, are expected to advance in altitude and latitude in response to climate warming. However, varied responses to 20th century warming suggest that in addition to temperature, tree line dynamics are mediated by species-specific traits and environmental conditions at landscape and local scales. 2 We examined recent tree line dynamics at six topographically different, but climatically similar, sites in south-west Yukon, Canada. Dendroecological techniques were used to reconstruct changes in density of the dominant tree species, white spruce ( Picea glauca), and to construct static age distributions of willow ( Salix spp.), one of two dominant shrub genera. Data were analysed to identify periods and rates of establishment and mortality and to relate these to past climate. 3 Tree line elevation and stand density increased significantly during the early to mid 20th century. However, this change was not uniform across sites. Spruce advanced rapidly on south-facing slopes and tree line rose 65–85 m in elevation. Tree line did not advance on north-facing slopes, but stand density increased 40–65%. Differences observed between aspects were due primarily to the differential presence of permafrost. Additional variability among sites was related to slope and vegetation type. Results were less conclusive for willow, but evidence for an advance was found at two sites. 4 Increases in stand density were strongly correlated with summer temperatures. The period of rapid change coincided with a 30-year period of above average temperatures, beginning in 1920. The highest correlations were obtained using a forward average of 30–50 years, supporting the hypothesis that tree line dynamics are controlled more by conditions influencing recruitment than by establishment alone. 5 The changes observed at several sites are suggestive of a threshold response and challenge the notion that tree lines respond gradually to climate warming. Overall, the results provide further evidence to support the idea that the pattern and timing of change is contingent on local, landscape, and regional-scale factors, as well as species’ biology. [ABSTRACT FROM AUTHOR]

From 2001 to 2004 we experimentally warmed 40 large, naturally established, white spruce [ Picea glauca (Moench) Voss] seedlings at alpine treeline in southwest Yukon, Canada, using passive open-top chambers (OTCs) distributed equally between opposing north and south-facing slopes. Our goal was to test the hypothesis that an increase in temperature consistent with global climate warming would elicit a positive growth response. OTCs increased growing season air temperatures by 1. 8°C and annual growing degree-days by one-third. In response, warmed seedlings grew significantly taller and had higher photosynthetic rates compared with control seedlings. On the south aspect, soil temperatures averaged 1. 0°C warmer and the snow-free period was nearly 1 month longer. These seedlings grew longer branches and wider annual rings than seedlings on the north aspect, but had reduced Photosystem-II efficiency and experienced higher winter needle mortality. The presence of OTCs tended to reduce winter dieback over the course of the experiment. These results indicate that climate warming will enhance vertical growth rates of young conifers, with implications for future changes to the structure and elevation of treeline contingent upon exposure-related differences. Our results suggest that the growth of seedlings on north-facing slopes is limited by low soil temperature in the presence of permafrost, while growth on south-facing slopes appears limited by winter desiccation and cold-induced photoinhibition. [ABSTRACT FROM AUTHOR]

Aim The species-specific response of tree-line species to climatic forcing is a crucial topic in modelling climate-driven ecosystem dynamics. In northern Québec, Canada, black spruce ( Picea mariana) is the dominant species at the tree line, but white spruce ( Picea glauca) also occurs along the maritime coast of Hudson Bay, and is expanding along the coast and on lands that have recently emerged because of isostatic uplift. Here we outline the present distribution, structure, dynamics and recent spread of white spruce from the tree line up to its northernmost position in the shrub tundra along the Hudson Bay coast. We aimed to obtain a minimum date of the arrival of the species in the area and to evaluate its dynamics relative to recent climate changes. Location White spruce populations and individuals were sampled along a latitudinal transect from the tree line to the northernmost individual in the shrub tundra along the Hudson Bay coast and in the Nastapoka archipelago in northern Québec and Nunavut, Canada (56°06′–56°32′ N). Methods White spruce populations were mapped, and the position, dimension, growth form and origin (seed or layering) of every individual recorded. Tree-ring analyses of living and dead trees allowed an estimation of the population structure, past recruitment, growth trends and growth rate of the species. A macrofossil analysis was performed of the organic horizon of the northernmost white spruce stands and individuals. Radiocarbon dates of white spruce remains and organic matter were obtained. The rate of isostatic uplift was assessed by radiocarbon dating of drifted wood fragments. Results The first recorded establishment of white spruce was almost synchronous at all sites and occurred aroundad 1660. Spruce recruitment was rather continuous at the tree line, while it showed a gap in the northern shrub tundra during the first decades of the 19th century. A vigorous, recent establishment of seedlings was observed in the shrub tundra; only wind-exposed, low krummholz (stunted individuals) did not show any sexual regeneration. A period of suppressed growth occurred from the 1810s to the 1850s in most sites. A growth increase was evident from the second half of the 19th century and peaked in the 1880s and the 20th century. A shift from stunted to tree growth form has occurred since the mid-19th century. No sample associated with white spruce remains gave a date older than 300 14C yearsbp [calibrated age (cal.)ad 1430–1690]. Main conclusions White spruce probably arrived recently in the coastal tundra of Hudson Bay due to a delayed post-glacial spread. The arrival of the species probably occurred during the Little Ice Age. The established individuals survived by layering during unfavourable periods, but acted as nuclei for sexual recruitment almost continuously, except in the northernmost and most exposed sites. Warmer periods were marked by strong seedling recruitment and a shift to tree growth form. Unlike white spruce, black spruce showed no evidence of an ongoing change in growth form and sexual recruitment. Ecological requirements and recent history of tree-line species should be taken into account in order to understand the present dynamics of high-latitude ecosystems. [ABSTRACT FROM AUTHOR]

The treeline in Alaska is usually mapped as the limit of white spruce Picea glauca (Moench) Voss along the south slope of the Brooks Range and in western Alaska and as the limit of Sitka spruce Picea sitchensis (Bong.) Can. on Kodiak Island and in the Alaska Peninsula. In some localities black spruce Picea mariana (Mill.) B.S.P. and paper birch Bemia papyrifera Marsh, fonn components of the treeline community with white spruce. Of special interest are the groves of balsam poplar Populus balsamifera L. occurring on the north slope of the Brooks Range and to the west and southwest of the spruce treeline. There is some question as to whether trees survived in unglaciated portions of Alaska during the last glacial period. Spruce trees became prevalent about 10000 yr ago and may still be expanding to their climatic limits. In some areas of Alaska the treeline was more expanded during the Hypsithermal period than it is at present, but in other regions evidence for such an expansion is lacking. Treetine in some areas seems to have been stable for the last several centuries, but is has been expanding rapidly for the last 40 yr in entral and western Alaska and for a longer period on Kodiak Island. [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]

The article offers information on the Yukon–Charley Rivers National Preserve (YCNP), a highly continental region with warmer growing seasons, a topographically diverse lowland landscape, and a considerably greater footprint of recent fire disturbance.

High northern latitude regions are experiencing some of the most rapid environmentalchanges. Carbon sequestration might be enhanced by increased forest productivity due towarming temperatures and potentially higher atmospheric CO2, but declines in tree growthand mortality due to drought stress could counteract this enhancement by releasing CO2 tothe atmosphere. Observed changes in boreal forest dynamics are already altering theglobal carbon and hydrological cycles, albedo and land surface feedbacks. Here, wecombined ring-width data with annually resolved stable carbon (δ13C) and oxygenisotope (δ18O) records to investigate the physiological responses of white spruce(Picea glauca [Moench] Voss) trees to warming and increased atmospheric CO2concentrations at northern treeline sites. We developed an extensive network of δ13Cand δ18O tree-ring chronologies covering the 20th century derived from 10 sitesacross North America (NA) spanning from 60.5º to 68.7ºN and 104º to 162ºW. Weused a range of observational data and a climate model to interpret signals in theseannually-resolved tree-ring series. At most sites, the inter-tree correlations werehigher for δ18O than δ13C series. Both δ13C and δ18O were highly influenced bytemperature variations, although the dominant season was different: summer and spring,respectively. This suggests that different processes associated with temperature variationsmay be operating. Physiological processes occurring at leaf level during summermainly influenced δ13C. Spring climate had the strongest influence on tree-ringδ18O via temperature effects on the δ18O of source water, and to a lesser degree,in summer, physiological process at leaf level may also occur. Accordingly, wefound weak relationships between the tree-ring δ18O and atmospheric circulation(such as integrated moisture transport) during summer using the GISS ModelE2isotopically-equipped general circulation model. Calibration and simulations of tree-ring datain the process-based mechanistic model MAIDENiso will help us to elucidate thecomplex linkages between increases in temperature and CO2 driven tree responses.This tree-ring isotope network provides a unique opportunity to disentangle theclimatic and physiological controls driving growth patterns at the northern treeline. [ABSTRACT FROM AUTHOR]