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Tree-ring data for 1942 - 2013 were used to determine if boreal and subalpine Picea albertiana S. Brown emend. Strong & Hills radial growth rates changed after 1950 in southern Yukon, northwest Canada. The latter year represented the beginning of accelerated atmospheric greenhouse gas accumulation, which has subsequently caused global climate changes. Ring-width chronologies for both boreal and subalpine trees were constructed using 80 trees. Individual ring-width sequences were crossdated, age-detrended, and converted to tree ring-width indices (RWI). In 2013, boreal and subalpine RWI were 41% and 169% greater than in 1950, respectively. In contrast, elevational treeline chronologies in the region typically decreased 20% - 70%. During 1942 - 2013, boreal and subalpine chronologies were significantly, at best modestly (r < |0.36|), and exclusively correlated with either moisture or temperature variables, respectively. Within this same time frame, RWI-climate variable correlates differed between the early and late phases of each chronology. Average pre-1969 or early-phase boreal correlations based on a 19-year moving window reached a maximum of - 0.56. Late-phase chronology correlations typically shifted towards weaker associations with climate variables (typically +0.04 to +0.30). For boreal RWI, 25 of 68 tested climate variables shifted as much as 0.40 - 0.88 correlation coefficient units between the early and late phases of the chronologies. Subalpine RWI were correlated only with July - August precipitation during the early phase of its chronology. These changes suggest responses by spruce to climate have shifted since 1970. The modest 1942 - 2013 RWI-climate variable correlations likely reflect the result of these changing ecological relationships. [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]

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]

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]

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]

Small-scale vertical aerial photographs taken in 1947 and 1948 covering 200 km2 of the Kluane Ranges, southwest Yukon, were compared with corresponding photographs taken in 1989 for the purpose of characterizing changes in the distribution and abundance of white spruce (Picea glauca (Moench) Voss) at the alpine treeline. Digital photogrammetry, including orthorectification and on-screen interpretation, was supplemented by stereoscopic inspection of the original prints. Qualitative assessment of change across nine image pairs was accompanied by quantitative analysis of changes in spruce density and elevation using 1 hectare plots and 100 m wide elevational belt transects, respectively, superimposed on the orthorectified images. Significant changes were observed over the 41 years, but the degree of change varied throughout the study area. The most common changes were an increase in canopy size of individual trees and an increase in stand density resulting from the establishment of new individuals. Several instances of treeline advance were also observed. An absence of major natural disturbances or widespread land use change indicates that treeline change is attributable to climate. Results from concurrent dendroecological studies indicate that these dynamics represent only part of the total extent of change to occur during 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]