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The evolutionary dynamics of tree species are influenced by their specific climatic environments, and their ability to persist is determined by adaptive strategies such as broad climate tolerance, phenotypic plasticity or genetic differentiation. Biogeographical predictions indicate that populations located at the edges of their ranges are more likely to experience heightened vulnerability to climatic fluctuations due to approaching tolerance limits. However, if local phenotypic acclimation or genetic adaptation has taken place, trees near the edge of their range could demonstrate comparable sensitivity to freeze and drought as the rest of the sampled population. Nevertheless, there remains uncertainty regarding the extent of developmental and evolutionary adjustments in climate sensitivity across the entire ranges of many tree species.Here, we document the biogeography of tree growth sensitivity to freeze and soil water and vapour pressure deficits during 1950–2018 using an extensive multi‐species tree‐ring dataset of 35,784 trees at 4535 sites covering boreal, temperate conifer and temperate deciduous forests of Canada. We quantify the relationships between tree radial growth increment and seasonal climate variables and explore factors driving the observed patterns of annual growth and climate sensitivity such as species, regional climate and local site conditions, and tree age and size.Freeze and drought had widespread impacts on tree growth that were contingent on the presence of focal species in interaction with tree size. An indirect growth thermal limitation towards cold/wet edges, occurring through the site moisture conditions, was observed in seven widespread species (e.g. Picea glauca, Picea mariana and Larix laricina). Moreover, six species had negative drought impacts more strongly expressed towards their warm/dry edges (e.g. Abies balsamea, Betula papyrifera and Pseudotsuga menziesii). However, widespread Picea, Pinus and Populus species showed no indication of increased sensitivity to soil water conditions at these edges.Synthesis. Our findings support the idea of evolutionary or acclimatization adjustments in the development of populations in response to long‐term climate conditions experienced in their respective locations. This underscores the importance of incorporating phenotypic and genomic data into future analyses of climate change impacts, which would enhance our ability to predict potential ecological shifts. [ABSTRACT FROM AUTHOR]

Baseline studies of small rodent populations in undisturbed ecosystems are rare. We report here 50 years of monitoring and experimentation in Yukon of a dominant rodent species in the North American boreal forest, the red‐backed vole Clethrionomys rutilus. These voles breed in summer, weigh 20–25 g, and reach a maximum density of 20 to 25 per ha. Their populations have shown consistent 3–4‐year cycles for the last 50 years with the only change being that peak densities averaged 8/ha until 2000 and 18/ha since that year. During the last 25 years, we have measured food resources, predator numbers, and winter weather, and for 1‐year social interactions, to estimate their contribution to changes in the rate of summer increase and the rate of overwinter decline. All these potential limiting factors could contribute to changes in density, and we measured their relative contributions statistically with multiple regressions. The rate of winter decline in density was related to both food supply and winter severity. The rate of summer increase was related to summer berry crops and white spruce cone production. No measure of predator numbers was related to winter or summer changes in vole abundance. There was a large signal of climate change effects in these populations. There is no density dependence in summer population growth and only a weak one in winter population declines. None of our results provide a clear understanding of what generates 3–4‐year cycles in these voles, and the major missing piece may be an understanding of social interactions at high density. [ABSTRACT FROM AUTHOR]

We investigated the genetic makeup of Lutz spruce, a natural hybrid between white and Sitka spruce on the Kenai Peninsula, Alaska. Microsatellites indicate 72% of individuals sampled had predominantly white spruce ancestry, whereas 14% had predominantly Sitka spruce ancestry; some individuals classified as white spruce had Sitka spruce plastid genotypes. As Picea mitochondria are maternally inherited and plastids are paternally inherited, it appears that white spruce was the ancestral seed parent of nearly all spruce on the western peninsula, whereas Sitka spruce alleles originated from pollen. Pollen records show that white spruce colonized the western peninsula ∼8500 YBP from glacial refugium, whereas Sitka spruce arrived on the eastern peninsula ∼4000 YBP after migrating up the Pacific coast. Our data suggest that Sitka spruce colonization west of the Kenai Mountains may have occurred not via seed dispersal but by long distance transport of wind-borne pollen and subsequent hybridization with established white spruce populations. Hybridization was an important mechanism that allowed Sitka spruce to expand the leading edge of its range in response to historical climate change. As the climate continues to warm, climate envelope modeling suggests Lutz spruce may ultimately displace white spruce on the western peninsula even as Sitka spruce is constrained to the eastern peninsula where it will continue to hybridize. [ABSTRACT FROM AUTHOR]

Under climate change circumstances, increasing studies have reported the temporal instability of tree growth responses to climate, which poses a major challenge to linearly extrapolating past climate and future growth dynamics using tree‐ring data. Space‐for‐time substitution (SFTS) is a potential solution to this problem that is widely used in the dendrochronology field to project past or future temporal growth response trajectories from contemporary spatial patterns. However, the projected accuracy of the SFTS in the climate effects on tree growth remains uncertain. Here, we empirically test the SFTS method by comparing the effect of spatial and temporal climate variations on climate responses of white spruce (Picea glauca), which has a transcontinental range in North America. We first applied a response surface regression model to capture the variations in growth responses along the spatial climate gradients. The results showed that the relationships between growth and June temperature varied along spatial climate gradients in a predictable way. And their relationships varied mainly along with local temperate condition. Then, the projected correlation coefficients between growth and climate using SFTS were compared against the observed. We found that the growth response changes caused by spatial versus temporal climate variations showed opposite trends. Moreover, the projected correlation coefficients using the SFTS were significantly lower than the observed. This finding suggests that applying the SFTS to project the growth response of white spruce might lead to an overestimation of the degree of tree maladaptation in future climate scenarios. And the overestimation is likely to get weaker from Alaska and Yukon Territory in the west to Quebec in the east. Although this is only a case study of the SFTS method for projecting tree growth response, our findings suggest that direct application of the SFTS method may not be applicable to all regions and all tree species. [ABSTRACT FROM AUTHOR]

Growth and yield (G&Y) of forest plantations can be significantly impacted by maladaptation resulting from climate change, and assisted migration has been proposed to mitigate these impacts by restoring populations to their historic climates. However, genecology models currently used for guiding assisted migration do not account for impacts of climate change on cumulative growth and assume that responses of forest population to climate do not change with age. Using provenance trial data for interior lodgepole pine (Pinus contorta subsp. latifolia Douglas) and white spruce (Picea glauca (Moench) Voss) in western Canada, we integrated Universal Response Functions, representing the relationship of population performance with their provenance and site climates, into top height curves in a G&Y model (Growth and Yield Projection System, GYPSY) to develop population-specific climate sensitive top height curves for both species. These new models can estimate the impact of climate change on top height of local populations and populations from a range of provenances to help guide assisted migration. Our findings reveal that climate change is expected to have varying effects on forest productivity across the landscape, with some areas projected to experience a slight increase in productivity by the 2050s, while the remainder are projected to face a significant decline in productivity for both species. Adoption of assisted migration, however, with the optimal populations selected was projected to maintain and even improve productivity at the provincial scale. The findings of this study provide a novel approach to incorporating assisted migration approaches into forest management to mitigate the negative impacts of climate change. [ABSTRACT FROM AUTHOR]

As northern latitudes experience rapid winter warming, there is an urgent need to assess the effect of varying winter conditions on tree growth and forest carbon sequestration potential. We examined tree growth responses to variability in cold-season (November–- April) frequency of freeze days (FFD) over 1951 to 2018 using tree-ring data from 35,217 trees and 57 species at 4,375 sites distributed across Canada. We found that annual radial growth responses to FFD varied by species, with some commonalities across genera and clades. The growth of gymnosperms with late spring leaf-out strategies was negatively related to FFD; years with high FFD were most detrimental to the annual growth of Pinus banksiana, Pinus contorta, Larix lyalli, Abies amabilis, and Abies lasiocarpa. In contrast, the growth of angiosperms with early leaf-out strategies, namely, Populus tremuloides and Betula papyrifera, was better in the coldest years, and gymnosperms with intermediate leaf-out timing, such as widespread Picea mariana and Picea glauca, had no consistent relationship to FFD. Tree growth responses to FFD were further modulated by tree size, tree age, regional climate (i.e., mean cold-season temperature), and local site conditions. Overall, our results suggest that moderately warming winters may temporarily improve the growth of widespread pines and some high-elevation conifers in western Canada, whereas warming winters may be detrimental to the growth of widespread boreal angiosperms. Our findings also highlight the value of using species-specific climate-growth relationships to refine predictions of forest carbon dynamics. [ABSTRACT FROM AUTHOR]

Mechanistically predicting the evolutionary response of tree species to climate change requires an understanding of genetic variation in relevant traits. Here we compared the phenotypic and genetic variation in the Leaf Economics Spectrum (LES) traits and the response of gas exchange to vapour pressure deficit (VPD) in lodgepole pine (Pico) and white spruce (Pigl), an early and a late successional species dominating the boreal forests of western Canada.We measured gas exchange, foliar nitrogen and lamina mass to area ratio in 697 c. 30‐year‐old trees in two field progeny trials. We analysed the response of gas exchange rates to VPD using a novel quantitative genetic model, the function‐valued trait approach.Pico showed greater phenotypic variation in the LES traits and greater genetic variation in photosynthetic rate than Pigl, but the species showed no significant difference in their phenotypic correlations between the LES traits. Pico showed a less sensitive stomatal response to VPD than Pigl and no significant genetic variation in stomatal sensitivity. In contrast, Pigl showed a positive correlation between the genetic values of stomatal sensitivity to VPD and stomatal conductance under low VPD.Our study region is projected to see an increase in VPD with climate change; the less sensitive and genetically diverse stomatal response to VPD in Pico could make this species more vulnerable to climate‐change‐induced droughts. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

With climate change, increasingly intense and frequent drought episodes will be affecting water availability for boreal tree species, prompting tree breeders and forest managers to consider adaptation to drought stress as a priority in their reforestation efforts. We used a 19‐year‐old polycross progeny test of the model conifer white spruce (Picea glauca) replicated on two sites affected by distinct drought episodes at different ages to estimate the genetic control and the potential for improvement of drought response in addition to conventional cumulative growth and wood quality traits. Drought response components were measured from dendrochronological signatures matching drought episodes in wood ring increment cores. We found that trees with more vigorous growth during their lifespan resisted better during the current year of a drought episode when the drought had more severe effects. Phenotypic data were also analyzed using genomic prediction (GBLUP) relying on the genomic relationship matrix of multi‐locus gene SNP marker information, and conventional analysis (ABLUP) based on validated pedigree information. The accuracy of predicted breeding values for drought response components was marginally lower than that for conventional traits and comparable between GBLUP and ABLUP. Genetic correlations were generally low and nonsignificant between drought response components and conventional traits, except for resistance which was positively correlated to tree height. Heritability estimates for the components of drought response were slightly lower than for conventional traits, but similar single‐trait genetic gains could be obtained. Multi‐trait genomic selection simulations indicated that it was possible to improve simultaneously for all traits on both sites while sacrificing little on gain in tree height. In a context of rapid climate change, our results suggest that with careful phenotypic assessment, drought response may be considered in multi‐trait improvement of white spruce, with accelerated screening of large numbers of candidates and selection at young age with genomic selection. [ABSTRACT FROM AUTHOR]

Increasing air temperatures and changing precipitation patterns due to climate change can affect tree growth in boreal forests. Periodic insect outbreaks affect the growth trajectory of trees, making it difficult to quantify the climate signal in growth dynamics at scales longer than a year. We studied climate‐driven growth trends and the influence of spruce budworm (Choristoneura fumiferana Clem.) outbreaks on these trends by analyzing the basal area increment (BAI) of 2058 trees of Abies balsamea (L.) Mill., Picea glauca (Moench) Voss, Thuja occidentalis L., Populus tremuloides Michx., and Betula papyrifera Marsh, which co‐occurs in the boreal mixedwood forests of western Quebec. We used a generalized additive mixed model (GAMM) to analyze species‐specific trends in BAI dynamics from 1967 to 1991. The model relied on tree size, cambial age, degree of spruce budworm defoliation, and seasonal climatic variables. Overall, we observed a decreasing growth rate of the spruce budworm host species, A. balsamea and P. glauca between 1967 and 1991, and an increasing growth rate for the non‐host, P. tremuloides, B. papyrifera, and T. occidentalis. Our results suggest that insect outbreaks may offset growth increases resulting from a warmer climate. The observation warrants the inclusion of the spruce budworm defoliation into models predicting future forest productivity. [ABSTRACT FROM AUTHOR]

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]

Climate change may have spatially variable impacts on growth of trees in topographically diverse environments, making generalizing across broad spatial and temporal extents inappropriate. Therefore, topography must be considered when analyzing growth response to climate. We address these topo-climatic relationships in the Canadian Rocky Mountains, focusing on lodgepole pine (Pinus contorta Douglas ex Louden) and interior spruce (Picea glauca (Moench) Voss × Picea engelmannii hybrid Parry) growth response to climate, Palmer drought severity index (PDSI), aspect, and slope angle. Climate variables correlate with older lodgepole pine growth on south- and west-facing slopes, including previous August temperature, winter and spring precipitation, and previous late-summer and current spring PDSI, but younger lodgepole pine were generally less sensitive to climate. Climate variables correlate with interior spruce growth on all slope aspects, with winter temperature and PDSI important for young and old individuals. Numerous monthly growth–climate correlations are not temporally stable, with shifts over the past century, and response differs by slope aspect and angle. Both species are likely to be negatively affected by moisture stress in the future in some, but not all, topographic environments. Results suggest species-specific and site-specific spatiotemporally diverse climate–growth responses, indicating that climate change is likely to have spatially variable impacts on radial growth response in mountainous environments. [ABSTRACT FROM AUTHOR]

Many forest tree species have characteristics that make the study of their evolutionary ecology complex. For example, they are long‐lived and thus have long generation times, and their often large, complex genomes have hampered establishing genomic resources. One way to tackle this challenge is to access multiple complementary data sources and analytical approaches when attempting to infer patterns of adaptive evolution. In the cover article of this issue of Molecular Ecology, Depardieu et al. (2021) combine large amounts of environmental, genomic, dendrochronological, and gene expression data in a common garden to explore the polygenic basis of drought resistance in white spruce (Picea glauca), a long‐lived conifer. They identify candidate genes involved in growth and resistance to extreme drought events and show how multiple data sets may deliver complementary evidence to circumvent the manifold challenges of the research field. [ABSTRACT FROM AUTHOR]

Summary: Phenology is an important indicator of environmental variation and climate change impacts on tree responses. In conifers, monitoring phenology of photosynthesis through remote sensing has been unreliable, because needle foliage varies little throughout the year. This is challenging for modelling ecosystem carbon uptake and monitoring phenology for enhanced breeding (genomic selection) and forest health.Here, we demonstrate that drone‐based carotenoid‐sensitive spectral indices, such as the Chl/carotenoid index (CCI), can be used to track phenology in conifers by taking advantage of the close relationship between seasonally changing carotenoid levels and the variation of photosynthetic activity.Physiological ground measurements, including photosynthetic pigments and maximum quantum yield of Chl fluorescence, indicated that CCI tracked the variation of photosynthetic activity better than other vegetation indices for 30 white spruce seedlings measured over 1 yr. A machine‐learning approach, using CCI derived from drone‐based multispectral imagery, was used to model phenology of photosynthesis for the entire pedigree population (6000 seedlings).This high‐throughput drone‐based phenotyping approach is suitable for studying climate change impacts and environmental variation on the physiological status of thousands of field‐grown conifers at unprecedented speed and scale. [ABSTRACT FROM AUTHOR]

Synchronous pulses of seed masting and natural disturbance have positive feedbacks on the reproduction of masting species in disturbance‐prone ecosystems. We test the hypotheses that disturbances and proximate causes of masting are correlated, and that their large‐scale synchrony is driven by similar climate teleconnection patterns at both inter‐annual and decadal time scales.Hypotheses were tested on white spruce (Picea glauca), a masting species which surprisingly persists in fire‐prone boreal forests while lacking clear fire adaptations. We built masting, drought and fire indices at regional (Alaska, Yukon, Alberta, Quebec) and sub‐continental scales (western North America) spanning the second half of the 20th century. Superposed Epoch Analysis tested the temporal associations between masting events, drought and burnt area at the regional scale. At the sub‐continental scale, Superposed Epoch Analysis tested whether El Niño‐Southern Oscillation (ENSO) and its coupled effects with the Atlantic Multidecadal Oscillation (AMO) in the positive phase (AMO+/ENSO+) synchronize drought, burnt area and masting. We additionally tested the consistency of our synchronization hypotheses on a decadal temporal scale to verify whether long‐term oscillations in AMO+/ENSO+ are coherent to decadal variation in drought, burnt area and masting.Analyses demonstrated synchronicity between drought, fire and masting. In all regions the year before a mast event was drier and more fire‐prone than usual. During AMO+/ENSO+ events sub‐continental indices of drought and burnt area experienced significant departures from mean values. The same was observed for large‐scale masting in the subsequent year, confirming 1‐year lag between fire and masting. Sub‐continental indices of burnt area and masting showed in‐phase decadal fluctuations led by the AMO+/ENSO+. Results support the 'Environmental prediction hypothesis' for mast seeding.Synthesis. We provide evidence of large‐scale synchronicity between seed masting in Picea glauca and fire regimes in boreal forests of western North America at both inter‐annual and decadal time scales. We conclude that seed production in white spruce predicts changes in disturbance regimes by sharing the same large‐scale climate drivers with drought and fire. This gives new insides in a mechanism providing a fire‐sensitive species with higher than expected adaptability to changes in climate. [ABSTRACT FROM AUTHOR]

Climate change is altering phenology; however, the magnitude of this change varies among taxa. Compared with phenological mismatch between plants and herbivores, synchronization due to climate has been less explored, despite its potential implications for trophic interactions. The earlier budburst induced by defoliation is a phenological strategy for plants against herbivores. Here, we tested whether warming can counteract defoliation‐induced mismatch by increasing herbivore‐plant phenological synchrony. We compared the larval phenology of spruce budworm and budburst in balsam fir, black spruce, and white spruce saplings subjected to defoliation in a controlled environment at temperatures of 12, 17, and 22°C. Budburst in defoliated saplings occurred 6–24 days earlier than in the controls, thus mismatching needle development from larval feeding. This mismatch decreased to only 3–7 days, however, when temperatures warmed by 5 and 10°C, leading to a resynchronization of the host with spruce budworm larvae. The increasing synchrony under warming counteracts the defoliation‐induced mismatch, disrupting trophic interactions and energy flow between forest ecosystem and insect populations. Our results suggest that the predicted warming may improve food quality and provide better growth conditions for larval development, thus promoting longer or more intense insect outbreaks in the future. [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]

A cost‐effective climate change adaptation strategy for the forestry sector is to move seed sources to more northern and higher elevation planting sites as part of ongoing reforestation programs. This is meant to match locally adapted populations with anticipated environments, but adaptive traits do not always show population differences suitable to mitigate climate change impacts. For white spruce, drought tolerance is a critical adaptive trait to prevent mortality and productivity losses. Here, we use a 40‐year‐old provenance experiment that has been exposed to severe drought periods in 1999 and 2002 to retrospectively investigate drought response and the adaptive capacity of white spruce populations across their boreal range. Relying on dendrochronological analysis under experimentally controlled environments, we evaluate population differences in resistance, resilience, and recovery to these extreme events. Results showed evidence for population differentiation in resistance and recovery parameters, but provenances conformed to approximately the same growth rates under drought conditions and had similar resilience metrics. The lack of populations with better growth rates under drought conditions is contrary to expectations for a wide‐ranging species with distinct regional climates. Populations from the wettest environments in the northeastern boreal were surprisingly drought‐tolerant, suggesting that these populations would readily resist water deficits projected for the 2080s, and supporting the view that northeastern Canada will provide a refugium for boreal species under climate change. The findings also suggest that white spruce is sensitive to growth reductions under climate change in the western boreal. The study highlights that population differentiation in adaptive capacity is species‐ and trait‐specific, and we provide a counterexample for drought tolerance traits, where assisted migration prescriptions may be ineffective to mitigate climate change impacts. For resource managers and policy makers, we provide maps where planning for widespread declines of boreal white spruce forests may be unavoidable. [ABSTRACT FROM AUTHOR]

Variable retention (VR) and partial cutting are both considered important silvicultural tools of natural disturbance or ecosystem based forest management approaches. Partial harvesting differs from VR in that post-treatment growth responses and stand regeneration are the primary objective rather than the maintenance of biodiversity. This partial cutting study is undertaken in mixed poplar (Populus spp.) – white spruce (Picea glauca (Moench) Voss) stands in the eastern Canadian boreal mixedwood forest. It compares, at the tree level, absolute growth rates (AGR) and relative growth rates (RGR) of basal area (BA) and stem survival; and at the stand level, it also compares absolute BA growth, mortality, and sapling density 10 years following treatment. The completely randomized experiment was established with four intensities of partial cutting (0, 50%, 65%, and 100% of poplar BA). All partial cutting intensities had a significant and similar positive effect on AGR of residual spruce stems. Complete poplar removal resulted not only in the highest increase in RGR of suppressed and intermediate spruce stems, but also in higher spruce mortality. Removal of 50% of the initial poplar stand BA provided the best trade-off between positive residual stem growth of spruce and poplar and limited post-treatment mortality. [ABSTRACT FROM AUTHOR]

The negative growth response of North American boreal forest trees to warm summers is well documented and the constraint of competition on tree growth widely reported, but the potential interaction between climate and competition in the boreal forest is not well studied. Because competition may amplify or mute tree climate‐growth responses, understanding the role current forest structure plays in tree growth responses to climate is critical in assessing and managing future forest productivity in a warming climate. Using white spruce tree ring and carbon isotope data from a long‐term vegetation monitoring program in Denali National Park and Preserve, we investigated the hypotheses that (a) competition and site moisture characteristics mediate white spruce radial growth response to climate and (b) moisture limitation is the mechanism for reduced growth. We further examined the impact of large reproductive events (mast years) on white spruce radial growth and stomatal regulation. We found that competition and site moisture characteristics mediated white spruce climate‐growth response. The negative radial growth response to warm and dry early‐ to mid‐summer and dry late summer conditions intensified in high competition stands and in areas receiving high potential solar radiation. Discrimination against 13C was reduced in warm, dry summers and further diminished on south‐facing hillslopes and in high competition stands, but was unaffected by climate in open floodplain stands, supporting the hypothesis that competition for moisture limits growth. Finally, during mast years, we found a shift in current year's carbon resources from radial growth to reproduction, reduced 13C discrimination, and increased intrinsic water‐use efficiency. Our findings highlight the importance of temporally variable and confounded factors, such as forest structure and climate, on the observed climate‐growth response of white spruce. Thus, white spruce growth trends and productivity in a warming climate will likely depend on landscape position and current forest structure. Using tree ring and carbon isotopes data from Denali National Park and Preserve, we investigated whether competition and site moisture characteristics mediate white spruce radial growth response to climate. We found competition and solar radiation mediate the influence of climate on the radial growth. Our carbon isotope analysis suggests the mechanism behind these modified climate‐growth responses is increased competition for moisture in high basal area stands and dry sites. Overall, our results suggest dynamic growth responses to future climate change that are dependent on landscape position and stand competition and likely to result in feedbacks on forest structure. [ABSTRACT FROM AUTHOR]

Growth of trees at their altitudinal and latitudinal range limits is expected to increase as climate warms, but trees often exhibit unexplained spatial and temporal variation in climate-growth responses, particularly in alpine regions. Until this variability is explained, predictions of future tree growth are unlikely to be accurate. We sampled Picea glauca (white spruce) growing at forest and tree line on north and south aspects in two mountain ranges of southwest Yukon to determine how and why ring-width patterns vary between topographic settings, and over time. We used multivariate statistical analysis to characterize variation in ring-width patterns between topographic factors and time periods, and calculated correlations between ring-width indices and climate variables to explain this variation. Ring-width patterns varied more between mountain ranges than elevations or aspects, particularly in recent decades when ring-widths increased in one mountain range but not the other. Growth responses to summer temperature were notably weaker during warmer time periods, but growth was not positively correlated to summer precipitation, suggesting trees may not be suffering from temperature-induced drought stress. Rather, ring-width indices began responding positively to spring snow depth after 1976. We conclude that tree growth is unlikely to increase in synchrony with rising air temperatures across subarctic tree lines in southwest Yukon. Instead, they may decline in areas that are prone to thin snowpacks or rapid spring runoff due to the negative influence warming springs will have on snow depth and, consequently, early growing season soil moisture. [ABSTRACT FROM AUTHOR]

Abstract: The boreal biome represents approximately one third of the world's forested area and plays an important role in global biogeochemical and energy cycles. Numerous studies in boreal Alaska have concluded that growth of black and white spruce is declining as a result of temperature‐induced drought stress. The combined evidence of declining spruce growth and changes in the fire regime that favor establishment of deciduous tree species has led some investigators to suggest the region may be transitioning from dominance by spruce to dominance by deciduous forests and/or grasslands. Although spruce growth trends have been extensively investigated, few studies have evaluated long‐term radial growth trends of the dominant deciduous species (Alaska paper birch and trembling aspen) and their sensitivity to moisture availability. We used a large and spatially extensive sample of tree cores from interior Alaska to compare long‐term growth trends among contrasting tree species (white and black spruce vs. birch and aspen). All species showed a growth peak in the mid‐1940s, although growth following the peak varied strongly across species. Following an initial decline from the peak, growth of white spruce showed little evidence of a trend, while black spruce and birch growth showed slight growth declines from ~1970 to present. Aspen growth was much more variable than the other species and showed a steep decline from ~1970 to present. Growth of birch, black and white spruce was sensitive to moisture availability throughout most of the tree‐ring chronologies, as evidenced by negative correlations with air temperature and positive correlations with precipitation. However, a positive correlation between previous July precipitation and aspen growth disappeared in recent decades, corresponding with a rise in the population of the aspen leaf miner (Phyllocnistis populiella), an herbivorous moth, which may have driven growth to a level not seen since the early 20th century. Our results provide important historical context for recent growth and raise questions regarding competitive interactions among the dominant tree species and exchanges of carbon and energy in the warming climate of interior Alaska. [ABSTRACT FROM AUTHOR]

Since 2001, climatic conditions have been notably drier than normal across large areas of the western Canadian interior, leading to widespread impacts on the forests of this region. This poses a major concern for the future, given climate change projections for continued warming and drying. We conducted tree-ring analysis in 75 pure stands of white spruce ( Picea glauca) across Alberta and west-central Saskatchewan to examine the effects of recent climatic drying on the growth of this important boreal tree species. Allometric equations were used to calculate annual growth in aboveground tree biomass ( GBM) from ring width measurements. Results showed an increasing trend in GBM from the 1960s to the 1990s, followed by a sharp decline during the severe drought of 2001-2002. Of the 75 stands, only 18 recovered sufficiently to cause an increase in mean GBM from the predrought decade of 1991-2000 to the subsequent decade of 2001-2010. The remaining 57 stands exhibited a decline in mean GBM between these decades. Climatic drying was a major cause of the growth decline, as shown by the significant stand-level relationship between percentage change in decadal mean GBM and the change in decadal mean values of a climate moisture index from 1991-2000 to 2001-2010. Subsequent analyses of boreal stands sampled across Alberta during 2015 revealed that white spruce growth had declined even further as drought conditions intensified during 2014-2015. Overall, there was a 38% decrease in mean GBM between 1997 and 2015, but surprisingly, the percentage decrease was not significantly different for young, productive stands compared with older, less productive stands. Thus, stand ageing cannot explain the observed decline in white spruce growth during the past quarter century, suggesting that these forests are at risk if the trend towards more frequent, severe drought continues in the region. [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]

Climate change is expected to increase winter temperatures in boreal climates. White spruce ( Picea glauca (Moench) Voss) is vulnerable to spring frost damage due to its habit of early budbreak, which may be exacerbated or lessened with increasingly warm winters at its southern range edge. We tested the effects of episodic warming during the quiescent stage on budbreak time and growth of seven seed sources grown in a common garden setting in Minnesota, USA. Treatment plots were warmed with infrared lamps for 4 days each in February, March, or February and March to simulate a midwinter thaw. Control plots for each treatment and an overall control were included for comparison. Trees warmed in February experienced a slight delay in spring budbreak, but differences in budbreak time were generally not significant. Terminal growth was significantly and negatively correlated with time of budbreak but not with time to growth cessation. Our results suggest that white spruce is relatively resilient to the effects of intermittent warming but that warming early in the season may delay budbreak time, which is expected to reduce terminal growth. [ABSTRACT FROM AUTHOR]

Climate warming is expected to increase the seasonal duration of photosynthetic carbon fixation and tree growth in high-latitude forests. However, photoperiod, a crucial cue for seasonality, will remain constant, which may constrain tree responses to warming. We investigated the effects of temperature and photoperiod on weekly changes in photosynthetic capacity, leaf biochemistry and growth in seedlings of a boreal evergreen conifer, white spruce [ Picea glauca (Moench) Voss]. Warming delayed autumn declines in photosynthetic capacity, extending the period when seedlings had high carbon uptake. While photoperiod was correlated with photosynthetic capacity, short photoperiods did not constrain the maintenance of high photosynthetic capacity under warming. Rubisco concentration dynamics were affected by temperature but not photoperiod, while leaf pigment concentrations were unaffected by treatments. Respiration rates at 25 °C were stimulated by photoperiod, although respiration at the growth temperatures was increased in warming treatments. Seedling growth was stimulated by increased photoperiod and suppressed by warming. We demonstrate that temperature is a stronger control on the seasonal timing of photosynthetic down-regulation than is photoperiod. Thus, while warming can stimulate carbon uptake in boreal conifers, the extra carbon may be directed towards respiration rather than biomass, potentially limiting carbon sequestration under climate change. [ABSTRACT FROM AUTHOR]

• Drought response of three disparate shelterbelt species is explored. • Severe and extreme 3-month droughts most greatly affect tree. • Scots pine and white spruce recover more quickly than green ash. Across western Canada, more frequent and severe droughts are expected to occur as a result of climate change. Forests are particularly vulnerable to drought as it often results in mortality events even years following a moisture deficit. Similar events have been observed in Saskatchewan shelterbelts, where many species are planted far outside their native range and are likely more vulnerable to environmental changes. Understanding how common shelterbelt species respond to drought will help landowners make better management decisions under a changing climate. This study looks at how three common shelterbelt species (green ash, Scots pine and white spruce) respond to different drought characteristics (severity, timing, and length) for up to five years following a moisture deficit using superposed epoch analysis. We identified that severe and extreme 3-month droughts greatly impact all three species. However, Scots pine and white spruce tend to recover more rapidly than green ash. Green ash is typically considered one of the most drought tolerant shelterbelt species, but its long recovery times suggest that this may be false. Scots pine was impacted by late summer droughts and white spruce was impacted by June droughts. Both Scots pine and white spruce also saw compensatory growth after their recovery period. This research augments our understanding of how drought characteristics impact three common shelterbelt species in Saskatchewan and how they are likely to recover under more frequent and severe drought events given projected climate change. [ABSTRACT FROM AUTHOR]

Premise of research: Understanding the relative contribution of environmental cues (photoperiod and temperature) as determinants of the transition from active growth to dormancy is important for better predictions of tree responses to anticipated climate change. The aim of this study was to identify the anatomical and structural changes that occur in the cambium of white spruce during the transition from active growth to dormancy triggered by short-day signals. Methodology: Anatomical features at the structural and ultrastructural levels were compared between actively and nonactively growing cambium to better understand how white spruce responds to short photoperiod. Pivotal results: The characteristics observed in the dormant cambium revealed that the organelles are densely and uniformly distributed throughout the cytoplasm. The abundance of small vacuoles, smooth endoplasmic reticulum, and starch granules were the main histological features characterizing the dormant state. In contrast to the dormant cambium cells, the active cells were characterized by abundant ribosomes, golgi bodies, and rough endoplasmic reticulum. Active cambium had cells with smaller radial dimensions and thinner cell walls than those in dormant cambium. Amyloplast-type plastids were also frequently seen and appeared to be empty. Cytoplasm in the active cambium was localized at the periphery of cambial cells and occupied by few large vacuoles in contrast to the numerous small vacuoles that appeared in the dormant state. Large and abundant lipid bodies were present in the active cambium cells. No differences were observed in the nucleus and mitochondria between active and dormant states. Conclusions: The number of cell layers and the morphology of cells in the cambium enabled us to define whether it is dormant or active. In active and dormant states, some dissimilarities in the shape and intensity of organelles were observed during the growth-to-dormancy transition, emphasizing the importance of stress protection in the tree's adaptive response to overwintering. Endoplasmic reticulum, starch granules, vacuoles, and lipid droplets showed great distinguishing characteristics between active and dormant cambium cells in white spruce. [ABSTRACT FROM AUTHOR]

Aim A critical concern for boreal ecosystems centres on broad-scale responses to warming, i.e. declining growth and mortality, or enhanced growth and greater productivity. However, few studies have synthesized tree growth along biogeographic gradients to address this issue. This study develops a broader understanding of how growth of a dominant conifer has responded to recent warming near the western forest margin of Alaska. Location Alaska, United States. Methods Thirty Picea glauca sites in south-west Alaska (1216 trees ≥ 4 cm dbh) were evaluated for growth differences from the southern boreal forest to the western forest-tundra margin across low-elevation forests and woodlands, and altitudinal tree line. Regional climate records were used to evaluate (1) whether tree growth near western tree line showed greater sensitivity to temperature and/or precipitation than southern boreal sites, (2) if the climate-growth response varied through time, across ecoregions and ecosystems, with spruce beetle disturbance, and by tree age, and (3) if there was a temperature threshold that limited growth. Results Positive growth trends since the 1980s in many open stands were consistent with the predicted expansion of western and altitudinal tree line. However, growth levelled off with temperatures ≥ 13°C at all but altitudinal tree line. An increasingly positive effect of precipitation on growth occurred after 1985, particularly at boreal tree line and closed forest sites. Closed-canopy forests showed lower rates of growth, greater spruce beetle activity and less potential for resiliency to warming than other ecosystem types. Main Conclusions Warming has led to markedly different growth responses according to ecosystem type and ecoregion near the forest-tundra margin. Altitudinal tree line sites showed consistently positive growth with warmer temperatures in recent decades, whereas low-elevation forests had reduced growth and greater beetle mortality. Strong positive correlations between growth and summer-fall precipitation since the mid-1980s suggest that precipitation will become an increasingly important factor with further warming. [ABSTRACT FROM AUTHOR]

Many studies have already addressed the existence of unstable and nonlinear relationships between radial growth of white spruce ( Picea glauca ) and climate variables in boreal forests along the high latitudes (>60° N). However, along the mid-latitudes, the climate-growth relationship is still poorly understood. In this study, we used a network of ring-width chronologies from 40 white spruce sites along a wide latitudinal gradients from 52° N to 58° N in Alberta, Canada and attempted to understand the complicated response of tree growth to climatic variables and to identify the main limiting factor for the radial growth of white spruce. We combined the empirical linear statistics with the process-based Vaganov-Shashkin Lite (VS-Lite) model requiring only latitude, monthly mean temperature, and monthly total precipitation information together to better clarify growth-climate relationship. The linear statistical methods indicated that the previous summer temperature imposed a strong negative impact on the radial growth of white spruce while the precipitation and climate moisture index in prior and current summer both had significant positive effects on the radial growth. Similarly, the VS-Lite model showed that the radial growth of white spruce was limited by soil moisture. This suggests that temperature-induced drought is the main limiting factor for the radial growth of white spruce. Furthermore, climate-growth relationship varied along different elevations, latitudes, and growing degree days (GDD >5 °C). The radial growth of white spruce in northern stands was often more strongly limited by temperature-induced drought due to the higher temperature and lower precipitation. As the global climate change is in progress, we suggest that more large-scale and continuous investigations are needed to address the spatial variation in growth-climate relationship due to the temperature-induced drought. [ABSTRACT FROM AUTHOR]

We examined modulus of elasticity (MoE) derived from SilviScan in white spruce ( Picea glauca (Moench) Voss) at the site, tree, and ring levels across the Boreal Plains ecozone in Canada. Area-weighted averages of MoE were calculated for juvenile and mature wood and were examined in relation to radial growth, climate, and location. Correlations indicated that there was a negative relationship between radial growth and MoE in the juvenile wood that was detectable at the site and tree levels; however, the relationship weakened in the mature wood, particularly at the site level. Few climate variables were correlated with MoE in juvenile wood, whereas multiple summer and fall climate variables showed a significant correlation with MOE in mature wood. A model describing the radial profile of MoE explained 58% of the variability in MoE, with 10% of the random variability attributed to between-tree differences. Elevation and summer water balance accounted for nearly all of the between-site variability. A decrease in MoE with increasing elevation was noted and has been previously linked to a decrease in cell wall thickness due to a shorter growing season at higher elevations. Integration of the MoE profile model into an individual-tree growth and yield simulator is the next logical step. [ABSTRACT FROM AUTHOR]

1. Local adaptation is a fundamental assumption in delineating seed zones and developing seed transfer guidelines to safeguard climatic adaptation of tree and plant species during forest regeneration and ecological restoration. It is also broadly assumed for forest tree species that show genetic differentiation among geographic populations, especially for those with widespread natural distributions that occur in distinct environments. 2. However, due to a scarcity of suitable data, the inference of local adaptation has rarely been validated for forest tree species through reciprocal transplant experiments (RTEs). In this study, we illustrate a novel approach to restructure tree provenance test data to conform to RTEs and use recently proposed statistical models to detect local adaptation, using white spruce Picea glauca (Moench) Voss as an example. Our research demonstrates how similar studies can be conducted to validate local adaptation in other forest species and populations, for which RTEs are lacking, but where large, high-quality provenance test data sets are available. 3. Contrary to common belief, our results show that local adaptation is absent in survival and height for white spruce populations from Ontario, Canada, although they have evolved in considerably different climatic habitats, and exhibit substantial and clinal genetic differentiation and significant genotype-by-environment interactions in these two adaptive traits. 4. Synthesis and applications. Our results show that the common assumption of local adaptation in forest tree species may not necessarily be correct within significant portions of their natural range. In forest genetic studies, population differentiation in adaptive traits has often been attributed to local adaptation without rigorous validation. Our results caution against such interpretation of experimental results. In the absence of local adaptation as shown by reciprocal transplant experiments, assisted migration may be considered as an option for enhancing forest adaptation to climate change. [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]

The late-Holocene shift from Picea glauca (white spruce) to Picea mariana (black spruce) forests marked the establishment of modern boreal forests in Alaska. To understand the patterns and drivers of this vegetational change and the associated late-Holocene environmental dynamics, we analyzed radiocarbon-dated sediments from Grizzly Lake for chironomids, diatoms, pollen, macrofossils, charcoal, element composition, particle size, and magnetic properties for the period 4100–1800 cal BP. Chironomid assemblages reveal two episodes of decreased July temperature, at ca. 3300–3150 (ca −1 °C) and 2900–2550 cal BP (ca −2 °C). These episodes coincided with climate change elsewhere in the Northern Hemisphere, atmospheric reorganization, and low solar activity. Diatom-inferred lake levels dropped by ca. 5 m at 3200 cal BP, suggesting dry conditions during the period 3200–1800 cal BP. P. glauca declined and P. mariana expanded at ca. 3200 cal BP; this vegetational change was linked to diatom-inferred low lake levels and thus decreased moisture availability. Forest cover declined at 3300–3100, 2800–2500 and 2300–2100 cal BP and soil erosion as inferred from increased values of Al, K, Si, Ti, and Ca intensified, when solar irradiance was low. Plant taxa adapted to disturbance and cold climate (e.g. Alnus viridis , shrub Betula , Epilobium ) expanded during these periods of reduced forest cover. This open vegetation type was associated with high fire activity that peaked at 2800 cal BP, when climatic conditions were particularly cold and dry. Forest recovery lagged behind subsequent climate warming (≤+3 °C) by ca. 75–225 years. Our multiproxy data set suggests that P. glauca was dominant under warm-moist climatic conditions, whereas P. mariana prevailed under cold-dry and warm-dry conditions. This pattern implies that climatic warming, as anticipated for this century, may promote P. glauca expansions, if moisture availability will be sufficiently high, while P. mariana may expand under dry conditions, possibly exacerbating climate impacts on the fire regime. [ABSTRACT FROM AUTHOR]

Climate-related variations in functional traits of boreal tree species can result both from physiological acclimation and genetic adaptation of local populations to their biophysical environment. To improve our understanding and prediction of the physiological and growth responses of populations to climate change, we studied the role of climate of seed origin in determining variations in functional traits and its implications for tree improvement programs for a commonly reforested boreal conifer, white spruce (Picea glauca (Moench) Voss). We evaluated growth, root-to-shoot ratio (R/S), specific leaf area (SLA), needle nitrogen (Nmass), total non-structural carbohydrates (NSC) and photosynthetic traits of 3-year-old seedlings in a greenhouse experiment using seed from six seed orchards (SO) representing the different regions where white spruce is reforested in Québec. Height and total dry mass (TDM) were positively correlated with photosynthetic capacity (Amax), stomatal conductance (gs) and mesophyll conductance (gm). Total dry mass, but not height growth, was strongly correlated with latitude of seed origin (SO) and associated climate variables. Amax, gs, gm and more marginally, photosynthetic nitrogen-use efficiency (PNUE) were positively associated with the mean July temperature of the SO, while water use efficiency (WUE) was negatively associated. Maximum rates of carboxylation (Vcmax), maximum rates of electron transport (Jmax), SLA, Nmass, NSC and R/S showed no pattern. Our results did not demonstrate a higher Amax for northern seed orchards, although this has been previously hypothesized as an adaptation mechanism for maintaining carbon uptake in northern regions. We suggest that gs, gm, WUE and PNUE are the functional traits most associated with fine-scale geographic clines and with the degree of local adaptation of white spruce populations to their biophysical environments. These geographic patterns may reflect in situ adaptive genetic differences in photosynthetic efficiency along the cline. [ABSTRACT FROM AUTHOR]

Tree improvement programs aim to develop families that are well-adapted to future growing conditions. To gain insight into the stability of the family genetic response to climate change, white spruce (Picea glauca) seedlings from 60 full-sib families were subjected to a combination of two temperature regimes and two levels of CO2over two growing seasons. There was positive effect of warmer temperatures and higher CO2on some growth variables but no significant family × treatment interactions. Instantaneous water use efficiency was the only physiological trait that was affected positively by the CO2treatment, showing a 51% increase that was consistent across families. [ABSTRACT FROM PUBLISHER]

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]

Intraspecific assisted migration ( ISAM) through seed transfer during artificial forest regeneration has been suggested as an adaptation strategy to enhance forest resilience and productivity under future climate. In this study, we assessed the risks and benefits of ISAM in white spruce based on long-term and multilocation, rangewide provenance test data. Our results indicate that the adaptive capacity and growth potential of white spruce varied considerably among 245 range-wide provenances sampled across North America; however, the results revealed that local populations could be outperformed by nonlocal ones. Provenances originating from south-central Ontario and southwestern Québec, Canada, close to the southern edge of the species' natural distribution, demonstrated superior growth in more northerly environments compared with local populations and performed much better than populations from western Canada and Alaska, United States. During the 19-28 years between planting and measurement, the southern provenances have not been more susceptible to freezing damage compared with local populations, indicating they have the potential to be used now for the reforestation of more northerly planting sites; based on changing temperature, these seed sources potentially could maintain or increase white spruce productivity at or above historical levels at northern sites. A universal response function ( URF), which uses climatic variables to predict provenance performance across field trials, indicated a relatively weak relationship between provenance performance and the climate at provenance origin. Consequently, the URF from this study did not provide information useful to ISAM. The ecological and economic importance of conserving white spruce genetic resources in south-central Ontario and southwestern Québec for use in ISAM is discussed. [ABSTRACT FROM AUTHOR]

Background EST (expressed sequence tag) sequences and their annotation provide a highly valuable resource for gene discovery, genome sequence annotation, and other genomics studies that can be applied in genetics, breeding and conservation programs for non-model organisms. Conifers are long-lived plants that are ecologically and economically important globally, and have a large genome size. Black spruce (Picea mariana), is a transcontinental species of the North American boreal and temperate forests. However, there are limited transcriptomic and genomic resources for this species. The primary objective of our study was to develop a black spruce transcriptomic resource to facilitate on-going functional genomics projects related to growth and adaptation to climate change. Results We conducted bidirectional sequencing of cDNA clones from a standard cDNA library constructed from black spruce needle tissues. We obtained 4,594 high quality (2,455 5' end and 2,139 3' end) sequence reads, with an average read-length of 532 bp. Clustering and assembly of ESTs resulted in 2,731 unique sequences, consisting of 2,234 singletons and 497 contigs. Approximately two-thirds (63%) of unique sequences were functionally annotated. Genes involved in 36 molecular functions and 90 biological processes were discovered, including 24 putative transcription factors and 232 genes involved in photosynthesis. Most abundantly expressed transcripts were associated with photosynthesis, growth factors, stress and disease response, and transcription factors. A total of 216 full-length genes were identified. About 18% (493) of the transcripts were novel, representing an important addition to the Genbank EST database (dbEST). Fifty-seven di-, tri-, tetra- and penta-nucleotide simple sequence repeats were identified. Conclusions We have developed the first high quality EST resource for black spruce and identified 493 novel transcripts, which may be species-specific related to life history and ecological traits. We have also identified full-length genes and microsatellite-containing ESTs. Based on EST sequence similarities, black spruce showed close evolutionary relationships with conspecific Picea glauca and Picea sitchensis compared to other Pinaceae members and angiosperms. The EST sequences reported here provide an important resource for genome annotation, functional and comparative genomics, molecular breeding, conservation and management studies and applications in black spruce and related conifer species. [ABSTRACT FROM AUTHOR]

Abstract: We present a reconstruction of June–July minimum temperatures since AD1245 for the Mackenzie Delta region based on a 29-site network of white spruce (Picea glauca) ring-width series. Most but not all trees experienced a divergent temperature–growth response, similar to the divergence that has affected other white spruce trees across Yukon and Alaska. However, divergence in the study region began as early as AD1900 and we have documented our methods to avoid including divergent signals in the reconstruction. Calibration/verification testing based on local temperature data, and multi-century coherence with nearby and large-scale temperature proxy records, confirm that our reconstruction is robust. The reconstruction shows cool conditions in the late 13th, early 18th and early 19th centuries, corresponding with solar minima and increased volcanism. These cool periods are interrupted by warm periods consistent with early to mid-20th century warmth. The late 20th century is the warmest interval, and the last decade is estimated to be 1.4°C warmer than any decade before the mid-20th century. The reconstructed climate history corroborates other proxy-based inferences and supports the notion that high-latitude regions such as the Mackenzie Delta have experienced rapid warming in recent decades that is exceptional in the last eight centuries. [Copyright &y& Elsevier]

Abstract: Sustainable forest management (SFM) requires the balancing of diverse values and conflicting management objectives. Climate change adds a further degree of uncertainty to this complex issue. In this study we analysed a Strategic Forest Management Plan (SFMP) from the southwest Yukon, Canada. The SFMP was developed to enable the salvage harvesting of beetle-killed white spruce stands and encourage fuel-abatement treatments to reduce fire risk to local communities. It did not, however, provide a long-term strategy to achieve SFM in the region. In this study, the SFMP served as the basis to develop and evaluate alternative forest management strategies in the context of climate change. Working group discussions with local stakeholders enabled the structuring of a ratings table that helped practitioners and experts to characterize five alternative strategies stemming from the SFMP, based on its main goals and objectives. An Analytic Hierarchy Process (AHP) was then used to balance competing values and objectives and test the alternatives against each other. The strategy ‘Manage for multiple values and use’ had the highest AHP-score when compared to the four other alternatives (timber, wildlife, fire risk reduction and carbon), which were narrower in scope. It may represent the best balance for the ecological and socio-economic values listed in the SFMP. Although the multiple values (MV) alternative is closest to the current SFMP in terms of scope, this study highlights that there are viable alternatives such as ‘manage for wildlife’ that achieved similar scores to the MV approach. The current SFMP has great potential to serve as a basis and starting point for a continuous and adaptive planning process for forest management. Exploring the different/alternative strategic directions will help to better address uncertain futures, thereby leading to more sustainable approaches. [Copyright &y& Elsevier]

Ongoing warming at high latitudes is expected to lead to large changes in the structure and function of boreal forests. Our objective in this research is to determine the climatic controls over the growth of white spruce ( Picea glauca (Moench) Voss) at the warmest driest margins of its range in interior Alaska. We then use those relationships to determine the climate variables most likely to limit future growth. We collected tree cores from white spruce trees growing on steep, south-facing river bluffs at five sites in interior Alaska, and analyzed the relationship between ring widths and climate using boosted regression trees. Precipitation and temperature of the previous growing season are important controls over growth at most sites: trees grow best in the coolest, wettest years. We identify clear thresholds in growth response to a number of variables, including both temperature and precipitation variables. General circulation model (GCM) projections of future climate in this region suggest that optimum climatic conditions for white spruce growth will become increasingly rare in the future. This is likely to cause short-term declines in productivity and, over the longer term, probably lead to a contraction of white spruce to the cooler, moister parts of its range in Alaska. [ABSTRACT FROM AUTHOR]

Along with climate, multiple global or large-scale change agents shape forest ecosystem health. We present a case study where we attempted to elucidate the driving factors causing decline symptoms in white spruce ( Picea glauca (Moench) Voss) in young spruce-fir boreal stands. Tree defoliation rate in the studied areas was related to the foliage discoloration intensity of the 2-year old needles and the decreasing stem basal area increment from 1997 to 2008. The onset of this growth decline in 1997 coincided with the occurrence of extremes for four climatic indices. The foliage of affected trees was deficient in K. The relationship between tree decline and K deficiency was tested through a diagnostic fertilization trial using a two-level factorial combination of N, K, and Mg. The trial indicated that K was the single limiting nutrient among the three tested elements. A single K addition increased stem basal area by 43% on average after 11 years. It is hypothesized that the poor K status of trees can be attributed to a series of natural and anthropogenic disturbances along with past forest management activities. Climate change in the region since the last decades also may have contributed to exacerbate K deficiency in such forest ecosystems. [ABSTRACT FROM AUTHOR]

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest-tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest-tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest-tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest-tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest-tundra ecotone. [ABSTRACT FROM AUTHOR]

Abstract: Stable isotope ratios from tree rings and peatland mosses have become important proxies of past climate variations. We here compare recent stable carbon and oxygen isotope ratios in cellulose of tree rings from white spruce (Picea glauca), growing near the arctic tree line; and cellulose of Sphagnum fuscum stems, growing in a hummock of a subarctic peatland, in west-central Canada. Results show that carbon isotopes in S. fuscum correlate significantly with July temperatures over the past ~20yr. The oxygen isotopes correlate with both summer temperature and precipitation. Analyses of the tree-ring isotopes revealed summer temperatures to be the main controlling factor for carbon isotope variations, whereas tree-ring oxygen isotope ratios are controlled by a combination of spring temperatures and precipitation totals. We also explore the potential of combining high-frequency (annual) climate signals derived from long tree-ring series with low-frequency (decadal to centennial) climate signals derived from the moss remains in peat deposits. This cross-archive comparison revealed no association between the oxygen isotopes, which likely results from the varying sensitivity of the archives to different seasons. For the carbon isotopes, common variance could be achieved through adjustments of the Sphagnum age model within dating error. [Copyright &y& Elsevier]

Site preparation and vegetation control can be used to mitigate climate change effects on early plantation growth in boreal forests. In this study, we explored growth of lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) and white spruce (Picea glauca (Moench) Voss) in relation to climate and site preparation using 20 years of data collected from studies in British Columbia. Results indicate that up to 45% of the variation in spruce growth and up to 37% of the variation in pine growth over this 20-year period can be explained by selected climatic variables. Monthly climate variables showed a stronger relationship to conifer growth than seasonal and annual variables. Climate variables related to the preceding year accounted for more than half of the variables in the final equations, indicating a lagged response in conifer growth. Future projections indicated that height growth of young lodgepole pine plantations in the sub-boreal zone could benefit (in the short term) from longer growing seasons by up to 12% on untreated stands. Untreated young white spruce plantations in the boreal zone may suffer height growth decreases of up to 10% due to increased drought stress. Vegetation control and mechanical site preparation treatments appear to mitigate effects of climate change to some extent. La préparation de terrain et la maîtrise de la végétation peuvent être utilisées pour atténuer les effets des changements climatiques sur la croissance de jeunes plantations en forêt boréale. Dans cette étude, nous évaluons la croissance du pin tordu latifolié (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) et de l'épinette blanche (Picea glauca (Moench) Voss) en fonction du climat et de la préparation de terrain à l'aide de données récoltées sur une période de 20 ans dans le cadre d'études réalisées en Colombie-Britannique. Les résultats indiquent que pendant cette période de 20 ans, jusqu'à 45 % de la variation de la croissance de l'épinette blanche et jusqu'à 37 % de la variation de la croissance du pin tordu peuvent être expliqués par des variables climatiques sélectionnées. Les variables climatiques mensuelles étaient davantage reliées à la croissance des conifères que les variables saisonnières et annuelles. Les variables climatiques reliées aux conditions de l'année précédente représentaient plus de la moitié des variables des équations finales, ce qui indique un décalage de la réaction de la croissance chez les conifères. Des projections indiquent que la croissance en hauteur des jeunes plantations de pin tordu de la zone subboréale pourrait, à court terme, être jusqu'à 12 % supérieure à celle de peuplements non traités si elles bénéficiaient de saisons de croissance plus longues. La croissance en hauteur des jeunes plantations non traitées d'épinette blanche dans la zone boréale pourrait diminuer jusqu'à 10 % à cause de l'augmentation des stress hydriques. La maîtrise de la végétation et la préparation mécanique de terrain semblent atténuer les effets des changements climatiques jusqu'à un certain point. [ABSTRACT FROM AUTHOR]

This paper integrates dendrochronological, demographic, and experimental perspectives to improve understanding of the response of white spruce (Picea glauca (Moench) Voss) tree growth to climatic variability in interior Alaska. The dendrochronological analyses indicate that climate warming has led to widespread declines in white spruce growth throughout interior Alaska that have become more prevalent during the 20th century. Similarly, demographic studies show that white spruce tree growth is substantially limited by soil moisture availability in both mid- and late-successional stands. Interannual variability in tree growth among stands within a landscape exhibits greater synchrony than does growth of trees that occupy different landscapes, which agrees with dendrochronological findings that the responses depend on landscape position and prevailing climate. In contrast, the results from 18 years of a summer moisture limitation experiment showed that growth in midsuccessional upland stands was unaffected by moisture limitation and that moisture limitation decreased white spruce growth in floodplain stands where it was expected that growth would be less vulnerable because of tree access to river water. Taken together, the evidence from the different perspectives analyzed in this study clearly indicates that white spruce tree growth in interior Alaska is vulnerable to the effects of warming on plant water balance. [ABSTRACT FROM AUTHOR]

Most wilderness areas still lack accurate distribution information on tree species. We met this need with a predictive GIS modeling approach, using freely available digital data and computer programs to efficiently obtain high-quality species distribution maps. Here we present a digital map with the predicted distribution of white spruce ( Picea glauca) in Alaska (4 km resolution, accuracy over 90%). Our presented concept represents a role-model for predicting tree species distribution for remote areas world-wide. Although this model intends to be accurate in making predictions rather than to give detailed biological mechanistic explanations, it can also be used as a baseline for further research and testable hypothesis on the importance of the environmental variables used to build a generalizable model. Further, we emphasize that work like presented here is a pre-condition for assessing human impacts and impacts of climate change on species distribution in a quantitative and transparent fashion, allowing for improved sustainable decision-making world-wide. [ABSTRACT FROM AUTHOR]

Interactions between vegetation and fire have the potential to overshadow direct effects of climate change on fire regimes in boreal forests of North America. We develop methods to compare sediment-charcoal records with fire regimes simulated by an ecological model, ALFRESCO (Alaskan Frame-based Ecosystem Code) and apply these methods to evaluate potential causes of a mid-Holocene fire-regime shift in boreal forests of the south-central Brooks Range, Alaska, USA. Fire-return intervals (FRIs, number of years between fires) are estimated over the past 7000 calibrated 14C years (7-0 kyr BP [before present]) from short-term variations in charcoal accumulation rates (CHARs) at three lakes, and an index of area burned is inferred from long-term CHARs at these sites. ALFRESCO simulations of FRIs and annual area burned are based on prescribed vegetation and climate for 7-5 kyr BP and 5-0 kyr BP, inferred from pollen and stomata records and qualitative paleoclimate proxies. Two sets of experiments examine potential causes of increased burning between 7-5 and 5-0 kyr BP. (I) Static-vegetation scenarios: white spruce dominates with static mean temperature and total precipitation of the growing season for 7-0 kyr BP or with decreased temperature and/or increased precipitation for 5-0 kyr BP. (2) Changed-vegetation scenarios: black spruce dominates 5-0 kyr BP, with static temperature and precipitation or decreased temperature and/or increased precipitation. Median FRIs decreased between 7-5 and 5-0 kyr BP in empirical data and changed-vegetation scenarios but remained relatively constant in static-vegetation scenarios. Median empirical and simulated FRIs are not statistically different for 7-5 kyr BP and for two changed-vegetation scenarios (temperature decrease, precipitation increase) for 5-0 kyr BP. In these scenarios, cooler temperatures or increased precipitation dampened the effect of increased landscape flammability resulting from the increase in black spruce. CHAR records and all changed-vegetation scenarios indicate long-term increases in area burned between 7-5 and 5-0 kyr BP. The similarity of CHAR and ALFRESCO results demonstrates the compatibility of these independent data sets for investigating ecological mechanisms causing past fire-regime changes. The finding that vegetation flammability was a major driver of Holocene fire regimes is consistent with other investigations that suggest that landscape fuel characteristics will mediate the direct effects of future climate change on boreal fire regimes. [ABSTRACT FROM AUTHOR]

CO2–light interactions can influence the competition among boreal plants, but are poorly understood. We investigated the effect of such interactions on the growth and biomass of 1-year-old black spruce (Sb) (Picea mariana (Mill.) BSP) and white spruce (Sw) (Picea glauca (Moench) Voss) grown with CO2 concentrations ([CO2]) of 360 and 720 µmol·mol–1 under 30%, 50%, and 100% light, in greenhouses. There were significant two-way and three-way interactions. Root collar diameter (RCD) of Sw decreased with decreasing light, while in Sb, there was no significant difference in RCD for plants grown under 50% or 30% light. Height was greater for plants grown under 100% light than if the plants were shaded. Elevated [CO2] increased RCD by 33% and enhanced stem volume by 67%, 98%, and 84% under 100%, 50%, and 30% light, respectively. The CO2 enhancement of total biomass was relatively higher under lower light, and greater for Sb than Sw. Elevated [CO2] decreased specific leaf area under 50% light only. Root mass was generally higher under 100% light than when shaded. Elevated [CO2] increased the root mass of Sb under 100% light, but decreased it under 30% light. Elevated [CO2] decreased the shoot/root ratio under 100% light, but increased it under 30% light. Our data suggest that raising [CO2] will likely increase species competitiveness under low light conditions, and that the increase will be greater in species that are relatively shade tolerant. [ABSTRACT FROM AUTHOR]