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Alaska's boreal forests are experiencing rapid changes in climate that may favor deciduous-dominated systems, with important implications for global biogeochemical and energy cycles. However, aspen (Populus tremuloides Michx.) has experienced substantial defoliation from the aspen leaf miner (Phyllocnistis populiella Cham., hereafter ALM) in Alaska, resulting in significant growth reductions. We conducted a tree-ring and Δ13C study to test the hypothesis that moisture limitation may have predisposed aspen to leaf miner damage. Contrary to our hypothesis, differences in climate-growth correlations between relatively severely and lightly affected trees were negligible during the pre-outbreak decades. Stands with greater summer precipitation had more limited ALM impact, however differences among models were small and multiple climate variables were suitable predictors of ALM impact. The strong negative relationship we detected between tree-ring Δ13C and basal area increment (BAI) suggested that interannual variation in Δ13C was driven primarily by variation in photosynthesis, limiting the utility of Δ13C as a tool to detect stomatal responses to moisture-limitation. Instead, we found that larger, faster-growing individuals on gentler slopes showed a stronger absolute reduction in BAI (pre-ALM BAIffpost-ALM BAI), but were similar in relative BAI reduction (pre-ALM BAI/post-ALM BAI), with smaller, slower growing trees. Older trees and stands with greater relative abundance of white spruce [Picea glauca (Moench) Voss] had greater relative ALM impact whereas slower growing trees on steeper slopes were less affected. The significant effect of white spruce abundance on ALM impact was likely due to favorable leaf miner overwintering habitat provided beneath white spruce trees, which can lead to increased leaf miner survival and thus greater reductions in aspen growth. Our results illustrate the subtle but complex biotic interaction between microclimate and pest physiology in determining ALM-induced aspen growth reductions, adding important nuance to a hypothesized increase in deciduous tree cover in Alaska's boreal forest. [ABSTRACT FROM AUTHOR]

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]

Global change in the climate is affecting tree/forest growth. There have been many studies that analyzed climate effects on tree growth. Results presented in these studies showed that the climate had both positive and negative effects on tree growth. The nature (positive/negative) and magnitude of the effects and the climate variables affecting growth depended on tree species. Climate-sensitive diameter growth models are not available for white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss) plantations. These models are needed to project forest growth and yield and develop forest management plans. Therefore, diameter growth models were developed for white pine and white spruce plantations by incorporating climate variables. Four hundred white pine and white spruce trees (200 per species) were sampled from 80 (40 per species) even-aged monospecific plantations (five trees per plantation) across Ontario, Canada. Diameter–age pairs were obtained from these trees using stem analysis. A nonlinear mixed-effects modeling approach was used to develop diameter growth models. To make the models climate sensitive, model parameters were expressed in term of climate variables. Inclusion of climate variables significantly improved model fit statistics and predictive accuracy. For evaluation, diameters (inside bark) at breast height were estimated for three geographic locations (east, west, and south) across Ontario for an 80-year growth period (2021–2100) under three climate change (emissions) scenarios (representative concentration pathway or RCP 2.6, 4.5, and 8.5 watts m−2). For both species, the overall climate effects were negative. For white spruce, the maximum pronounced difference in projected diameters after the 80-year growth period was in the west. At this location, compared to the no climate change scenario, projected spruce diameters under RCPs 2.6 and 8.5 were thinner by 4.64 (15.99%) and 3.72 (12.80%) cm, respectively. For white pine, the maximum difference was in the south. Compared to the no climate change scenario, projected pine diameters at age 80 under RCPs 2.6 and 8.5 at this location were narrower by 4.54 (13.99%) and 7.60 (23.43%) cm, respectively. For both species, climate effects on diameter growth were less evident at other locations. If the values of climate variables are unavailable, models fitted without climate variables can be used to estimate these diameters for both species. [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]

An increase in the frequency and magnitude of drought events threatens the health of forests and the economic, ecological, and societal services they provide. It has been widely demonstrated that trees undergoing a succession of stresses may accumulate lesions that in turn lead to a decrease in their vigor and eventually to death. However, recent studies have shown that a nonlethal stress should also initiate a stress memory, which triggers a faster and stronger plant defensive response when a new stress occurs. Although this mechanism is well understood in many herbaceous plants, a better understanding in trees is needed. The aim of our study was to explore the capacity of two forest tree species to develop a stress memory. A greenhouse experiment was conducted to evaluate the tree seedlings' vigor after one or two consecutive droughts separate from a rehydration period during the same growing season. No stress memory pattern was observed for the two tree species as, on the contrary, we even observed a stress accumulation pattern in sugar maple. It remains possible that some individuals in our study developed stress memory, but that we were not able to detect it. The fine‐tuning of experimental parameters and the conducting of longitudinal studies would be helpful to detect individual capacity in stress memory activation. [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]

Global warming is predicted to extend the growing season of trees and plants, and advance spring phenology. However, intensification of extreme climate events in midlatitude forests, from weakening of the jet stream and atmospheric blockings, may expose trees to increased risk associated with more frequent late-spring frosts. Still, little is known regarding the intraspecific variation in frost tolerance and how it may be shaped by local adaptation to the climate of seed origin. As part of an assisted migration trial located in different bioclimatic zones in the province of Quebec, Canada, and following an extensive late-spring frost that occurred at the end of May 2021, we evaluated the frost damages on various white spruce (Picea glauca) seed sources tested on three sites (south, central, and north). The severity of frost damages was assessed on 5,376 trees after the cold spell and an early spring warming which advanced bud flush by approximately 10 days on average. The frost damage rate was similar among sites and seed sources and averaged 99.8%. Frost damage severity was unrelated to the latitude of seed origin but was variable among sites. The proportion of severely damaged trees was higher in the northern site, followed by central and southern sites. The proportion of severely damaged trees was linearly and inversely related to tree height before the frost event. Apical growth cancelation was not significantly different among seed sources including local ones, and averaged 74, 46, and 22%, respectively, in central, northern, and southern plantation sites. This study provides recommendations to limit the loss of plantation productivity associated with such a succession of spring climate anomalies. Implications for seed transfer models in the context of climate change and productivity of spruce plantations are discussed in the light of lack of local adaptation to such pronounced climate instability and ensuing large-scale maladaptation [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]

Spruce–fir (Picea–Abies) forests of the North American Acadian Forest Region are at risk of disappearing from the northeastern United States and Canada due to climate change. Species distribution models (SDMs) have been used to predict changes in this critical transitional ecosystem in the past, but none have addressed how seasonal patterns of temperature and precipitation interact to influence tree species abundance. Inferences have also been limited by contemporary inventory data that could not fully characterize species ranges because they either, (1) only sampled species occurrence after large‐scale human disturbance and settlement, or (2) did not span critical geopolitical boundaries (e.g., the US–Canadian border) that intersect the focal species' range(s). Here, we built new SDM models to better assess the bioclimatic distribution of four spruce–fir species and to test the importance of seasonal climate interactions. We compiled an extensive database of tree occurrence and abundance from recent (~1955–2012) and historical time periods (1623–1869) to model current species distributions and to predict how these might change under future climate. We found that including historical tree data in our SDMs revealed previously unrecognized suitable habitat along the southern edge of species' contemporary ranges. Random forest models predicted occurrence with high accuracy (area under receiver operator curve >0.98), and the seasonal climate variables that emerged as most important for these cold‐adapted species all included interactions that reflected sensitivity to colder temperatures, and preferences for wet weather concentrated in the winter months. Under moderate climate warming (representative concentration pathway 6.0), the northeastern United States retained additional suitable habitat when historical data were included through 2060 for three of the four species: red spruce (Picea rubens), black spruce (Picea mariana), and balsam fir (Abies balsamea), while white spruce (Picea glauca) habitat contracted into Canada. In contrast, future predictions from models that used contemporary data alone forecast extirpation for all four species from the northeastern United States. Overall, these findings highlight that prediction of species ranges in transitional ecosystems that span geopolitical boundaries and gradients of intense land use are improved when historical data and seasonal climate interactions of both temperature and precipitation variables are incorporated. [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]

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]

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]

The ranges of black and white spruce are largely sympatric, suggesting both species have similar climate requirements. The two species, however, are highly segregated across the landscape with black spruce most common on nutrient‐poor sites with cold, poorly drained soils and white spruce more common on productive sites with warmer, well‐drained soils. Because site conditions influence tree climate–growth responses, it is difficult to compare white and black spruce climate–growth responses as these responses are confounded by the differences in site conditions in which the two species naturally occur. As the climate warms dramatically in northern latitudes, it is critical to understand how a changing climate and associated changes in permafrost and fire regimes will interact to shape future species composition and ecosystem functioning in the boreal forest. In this study, we examined the climate–growth responses of black and white spruce growing in the same sites. This approach eliminates the confounding factor of site conditions and facilitates our understanding of how these two species respond to climate. We included standardized thaw depth of the active layer in our analysis as a representation of permafrost, which is a key factor delineating these two species' habitat preferences and is actively warming and thawing as the climate warms. Our most important finding was that the climate–growth responses of the two species, but especially white spruce, hinged on the thaw depth of the active layer. Specifically, with increasing June–July temperatures white spruce radial growth increased in areas with deep thaw or no near‐surface permafrost, but strongly decreased when growing in areas with near‐surface permafrost. Black spruce radial growth was less sensitive to June–July temperature than white spruce but had a consistent and more positive response to summer precipitation. These findings point to a primary mechanism potentially driving the positioning of these two tree species within the landscapes of boreal interior Alaska and imply widespread thawing of permafrost may foster expansion of white spruce in this region at the expense of black spruce, but that in a wetter climate, black spruce may gain competitive advantage over white spruce in some landscape positions. [ABSTRACT FROM AUTHOR]

Future warming may alter plant stress at high‐elevation treelines and forests, thereby changing plant–plant interactions. The relative importance of competition and facilitation may depend on the degree of resource or physical stress. According to the stress gradient hypothesis (SGH), physical stress on trees is more important at cold high elevations where facilitation predominates, and less important at low elevations where competition is the main interaction. Our goals were to investigate whether plant–plant interactions along elevational gradients corresponded to those predicted by the SGH, and to assess the effects of increasing temperatures on the growth of conifer seedlings in tundra–forest ecosystems in interior Alaska, USA. We established sites along two elevational gradients: one in tundra (four sites, 550–1170 m) and one in forest (three sites, 210–760 m). A field warming and neighbor removal experiment was conducted using transplanted seedlings of white spruce (Picea glauca). After three growing seasons, regardless of elevation, spruce seedling biomass was approximately 20% lower in seedlings with neighbor plants compared to those without neighbors. Therefore, there was no evidence for greater facilitation at high elevations across either tundra or forest elevational gradient. Seedlings in forest sites increased their shoot‐to‐root ratios and relative growth rates in height, suggesting competition for light, but this was not seen for seedlings growing in tundra sites. When warmed by greenhouses, seedling growth was stimulated in forests, especially at high elevations. In contrast, at tundra sites, warmed seedlings grew less than controls and the seedlings at high elevations showed water stress. Our results suggest that the resource stresses of low light and water availability are more important drivers of plant–plant interactions than the physical stress imposed by low temperature, possibly due to warming in the past 50 yr in this region. Further warming may increase growth of seedlings in forests, but is likely to decrease it in open tundra habitats, thereby slowing the upward movement of treeline in the near future. [ABSTRACT FROM AUTHOR]

Key message: Growth sensitivity to climate varies with soil moisture regime in spruce–fir forests in central British Columbia. Stands growing at their dry edaphic limits displayed especially strong and unique climatic sensitivities. Soil moisture regime is an important influence of productivity, process, and structure in forested ecosystems. In western North America, projected warming trends may result in decreasing available soil moisture; however, the potential effects on forest growth remain unclear. This study aimed to determine the influence of stand-level soil moisture regime on the climatic sensitivity of mature hybrid white spruce (Picea glauca (Moensch) Voss x Picea engelmannii Parry) and subalpine fir (Abies lasiocarpa (Hook.) Nutt.) forests in central British Columbia, Canada. We collected and analyzed tree-ring data from 51 stands spanning a range of soil moisture regimes. Dendroecological analyses of climate–growth relationships indicated that warm summer temperatures and drought limit growth for both species across all soil moisture regimes; however, responses were strongest on the driest sites. Spruce populations across the gradient of soil moisture regimes displayed unique climate-growth relationships; growth in populations on wetter sites was more correlated with summer climate from the year prior to growth. Radial growth responses to prior summer temperatures strengthened over the past ca. 80 years in both species and across most sites, suggesting that climate–growth relationships are shifting in this region. This study presents evidence of the importance of considering site-level ecological factors such as soil moisture regime when studying forest growth responses to climate. [ABSTRACT FROM AUTHOR]

Xylem anomalies that are caused by unusual climate events have long been used to aid cross-dating in tree ring research. Here, we analyzed a range of xylem anomalies in a 39-year-old common garden experiment of white spruce (Picea glauca [Moench] Voss) in central Alberta, Canada, designed to investigate local adaptation. We extracted wood cores from trees representing 24 provenances covering much of the species range across the Canadian boreal forest. Using a double stain and light microscopy analysis, four xylem anomalies and their causes could be distinguished: (1) frost rings indicate issues with synchronizing the onset of growth with the start of the growing season, and were prevalent in young trees; (2) light rings represent thin cell walls caused by an insufficient growing season length, most prevalent in southern sources; (3) blue rings were caused by a failure to complete lignification of new wood due to an early end of the growing season; and (4) double rings represent density fluctuations due to drier than normal summers. Local provenances showed the least amount of xylem anomalies, indicating that they are correctly adapted to the environment in which they occur. In contrast, trees moved to the test site from other climate regions showed various types of xylem anomalies depending on their origin. In particular, populations originating from warmer regions showed an increased presence of latewood anomalies, consistent with a more extensive use of the growing season in the fall. We conclude that xylem anomalies may serve as a sensitive early indicator of maladaptation to climate before populations experience tree dieback or mortality. They may therefore be useful to monitor the health of natural populations, or to evaluate the success of assisted migration in reforestation to address climate change.

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

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]

Xylem anomalies that are caused by unusual climate events have long been used to aid cross-dating in tree ring research. Here, we analyzed a range of xylem anomalies in a 39-year-old common garden experiment of white spruce (Picea glauca [Moench] Voss) in central Alberta, Canada, designed to investigate local adaptation. We extracted wood cores from trees representing 24 provenances covering much of the species range across the Canadian boreal forest. Using a double stain and light microscopy analysis, four xylem anomalies and their causes could be distinguished: (1) frost rings indicate issues with synchronizing the onset of growth with the start of the growing season, and were prevalent in young trees; (2) light rings represent thin cell walls caused by an insufficient growing season length, most prevalent in southern sources; (3) blue rings were caused by a failure to complete lignification of new wood due to an early end of the growing season; and (4) double rings represent density fluctuations due to drier than normal summers. Local provenances showed the least amount of xylem anomalies, indicating that they are correctly adapted to the environment in which they occur. In contrast, trees moved to the test site from other climate regions showed various types of xylem anomalies depending on their origin. In particular, populations originating from warmer regions showed an increased presence of latewood anomalies, consistent with a more extensive use of the growing season in the fall. We conclude that xylem anomalies may serve as a sensitive early indicator of maladaptation to climate before populations experience tree dieback or mortality. They may therefore be useful to monitor the health of natural populations, or to evaluate the success of assisted migration in reforestation to address climate change. [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]

• Statistical validation alone may yield unrealistic models; spatial validation is vital. • Weak genetic effects for interior spruce, hinting modest local adaptation. • Climate change, especially heat stress, threatens current interior spruce habitats. • Climate change may lead to upslope shift and lower interior spruce productivity. Ecological models, including various forms of population transfer and response functions, have been widely employed to predict the performance of forest trees under future climate conditions. However, the accuracy of model predictions remains a challenge due to the lack of systematic model optimization and validation processes. In this study, we explored a series of candidate universal response functions (URFs) based on comprehensive provenance trials for interior spruce, a tree hybrid complex of great ecological and economic importance in British Columbia. We first developed URFs with various numbers of climate variables and their combinations, then evaluated and validated these models with conventional statistical methods. Although these models showed high predictive power and performed well in statistical validation, the map visualizations of these models varied substantially. Therefore, we further conducted a pixel-wise spatial validation based on site productivity data to identify the model with the best performance. Finally, we analyzed the impact of climate change on interior spruce based on the final model. Our results revealed that: 1) a minimum number of independent climate variables is required to generate a reliable and informative ecological model; 2) spatial validation is critical to ensure the credibility of ecological models; 3) among-population variation resulting from local adaptation is relatively weak, indicating a potential for modest productivity gains from selecting well-adapted populations for stand establishment; and 4) most of the current interior spruce habitat is projected to be vulnerable to anthropogenic climate change, largely due to temperature rise, with a major upslope shift predicted. [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]

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]

Under a changing climate, tree improvement programs should revise breeding zones to improve climate change adaptation and program efficiency once information about adaptive capacity of breeding populations and genotype × environment (G × E) interaction becomes available. In this study, we evaluated the adaptation and growth, as well as G × E interactions, shown by white spruce (Picea glauca (Moench) Voss) open-pollinated (OP) families across two adjacent breeding zones (i.e., the BZ3300 and BZ4300, respectively) in northwestern Ontario using progeny test data, including 80 OP families (40 from each) that were cross-tested between the breeding zones. Results showed that the OP families from the southerly BZ4300 breeding zone averaged higher survival and significantly greater mean tree height growth than the local OP families in the more northern BZ3300 breeding zone at tree age of 14 years, with no notable difference in frost damage. Estimates of type-B genetic correlation from pairwise tests or combined-test analyses were almost all greater than 0.7, suggesting weak G × E interactions. Greater variation among families and higher heritability estimates in tree height were observed for families from the BZ3300 breeding zone or when families from the two breeding zones were combined. The weak G × E interactions, in addition to better survival and growth of the southerly families in the more northern breeding zone, suggested benefits from deploying seed from current seed orchard in the BZ4300 breeding zone to the BZ3300 breeding zone and merging the two breeding zones when starting the second generation breeding to increase tree improvement efficiency and likely higher genetic gain under future climate. [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]

Long hydroclimate records are essential elements for the assessment and management of changing freshwater resources. These records are especially important in transboundary watersheds where international cooperation is required in the joint planning and management process of shared basins. Dendrochronological techniques were used to develop a multicentury record of April 1 snow water equivalent (SWE) for the Stikine River basin in northern British Columbia, Canada, from moisture‐sensitive white spruce (Picea glauca) tree rings. Explaining 43% of the instrumental SWE variability, to our knowledge, this research represents the first attempt to develop long‐term snowpack reconstructions in northern British Columbia. The results indicated that 15 extreme low April 1 SWE events occurred from 1789 to the beginning of the instrumental record in 1974. The reconstruction record also shows that the occurrence of hydrological extremes in the Stikine River basin is characterized by persistent below‐average periods in SWE consistent with phase shifts of the Pacific Decadal Oscillation (PDO). Spectral analyses indicate a very distinct in‐phase (positive) relationship between the multidecadal frequencies of variability (~40 years) extracted from the SWE tree‐ring reconstruction and other reconstructed winter and spring PDO indices. Comparison of the reconstructed SWE record with other tree‐ring‐derived PDO proxy records shows coherence at multidecadal frequencies of variability. The research has significant implications for regional watershed management by highlighting the hydrological response of the Stikine River basin to prior climate changes. [ABSTRACT FROM AUTHOR]

Highlights • We evaluate risk of frost damage in assisted migration to address climate change. • Planting stock moved north shows tradeoffs between growth and cold hardiness. • Day length regime is the main environmental variable regulating this tradeoff. • Limiting northward transfers to 500 km maintained hardiness with good growth. Abstract Human assisted movement of reforestation stock poleward or upward in elevation has been proposed as a tool to address climate change in regular reforestation programs. However, moving warm adapted seed sources to colder environments could carry the risk of frost damage if seed sources are moved too far. Here, we assess genetic differentiation in growth potential, survival and cold hardiness of a wide-ranging tree species, white spruce (Picea glauca [Moench] Voss). We use data from a 34-year-old common garden experiment planted in the approximate center of the species range. Cold hardiness was negatively related to growth and positively to survival. Generalized additive models identified mean coldest month temperature and latitude (as a proxy for the day length regime) as an exceptionally good predictor for the onset of cold hardiness (R2 adj = 0.91). The results suggest that day length, an environmental factor that is not influenced by climate change, is an important factor controlling the timing of the onset of hardiness. Survival was only moderately well predicted, primarily by precipitation of the provenance origin (R2 adj = 0.36) indicating that other adaptive traits besides cold hardiness should be considered in assisted migration prescriptions. Survival of seed sources appears primarily compromised when transplanting sources from wet origins to a dry location. Nevertheless, acceptable migration distances without significant tradeoffs were up to 500 km north and 1500 km west towards a central planting location. [ABSTRACT FROM AUTHOR]

• We exposed seedlings of conifer species to three realistic simulated future climates. • Seedlings of conifer species experienced high mortality (75%) induced by drought. • Conifer species are highly vulnerable at the seedling stage to future drought. • Hydraulic safety margins are good indicators of mortality risk for conifer species. The future climate of northern temperate forests is projected to be drier and warmer by the end of this century. As a result, more drought-induced forest dieback events are anticipated in northeastern North America, and assessing the vulnerability of dominant tree species to drought is critical for understanding the future composition of these forests. In a greenhouse experiment, we exposed two-year-old seedlings of Picea glauca (Moench) Voss, Picea mariana (Mill.) B.S.P. and Pinus strobus L. to three future climate treatments for southern Quebec, Canada, and evaluated their mortality, growth, and foliage water status responses to soil water availability and atmospheric drought. Using a unique approach, climate treatments emulated droughts of different frequencies, durations, and intensities. Treatments closely simulated one growing season, with changes in air temperature and relative humidity every six hours and daily adjustment in the amount of water delivered to the seedlings. The three species experienced high mortality (75%) in all water-limited treatments compared to a control treatment that provided non-limiting soil moisture (0% mortality). The biomass of the seedlings that survived was 40% lower than that of control seedlings. Our results confirmed that the hydraulic safety margins, defined as the difference between seasonal minimum water potential and xylem water potential leading to 12, 50 and 88% of hydraulic conductivity loss, were good predictors of probability of tree mortality. Therefore, hydraulic safety margins are useful functional traits that can be used to compare the vulnerability of various species to drought and then provide crucial information to practitioners and policymakers to adjust forest management to climate change. We showed that three dominant conifer species of northern temperate forests were highly vulnerable to drought in future climates. Because drought is projected to be a significant threat to forests, understanding potentially adaptive physiological responses to drought, such as hydraulic safety margins of tree seedlings, is important for predicting the response of forest regeneration and composition in warmer and drier climates. [ABSTRACT FROM AUTHOR]

With climate change, favorable growing conditions for tree species are shifting northwards and to higher altitudes. Therefore, local populations are becoming less adapted to their environment. Assisted migration is one of the proposed adaptive measures to reduce the vulnerability of natural populations and maintain forest productivity. It consists of moving genetic material to a territory where future climate conditions correspond to those of its current location. Eight white spruce (Picea glauca [Moench] Voss) seed sources representing as many seed orchards were planted in 2013 at three forest sites simulating a south-north climatic gradient of 1.7°C in Québec, Canada. The objectives were to (1) evaluate the morpho-physiological responses of the different seed sources and (2) determine the role of genetic adaptation and physiological plasticity on the observed variation in morpho-physiological traits. Various seedling characteristics were measured, notably height growth from nursery to the fourth year on plantation. Other traits such as biomass and carbon allocation, nutritional status, and various photosynthetic traits before bud break, were evaluated during the fourth growing season. No interaction between sites and seed sources was observed for any traits, suggesting similar plasticity between seed sources. There was no change in the rank of seed sources and sites between years for height growth. Moreover, a significant positive correlation was observed between the height from the nursery and that after 4 years in the plantation. Southern seed sources showed the best height growth, while optimum growth was observed at the central site. Juvenile height growth seems to be a good indicator of the juvenile carbon sequestration and could serve as a selection criterion for the best genetics sources for carbon sequestration. Vector analysis showed no nitrogen deficiency 4 years after planting. Neither seed sources nor planting sites had a significant effect on photosynthesis before bud break. The observed results during the establishment phase under different site conditions indicate that southern seed sources may already benefit from assisted migration to cooler climatic conditions further north. While northern seed sources are likely to benefit from anticipated local global warming, they would not match the growth performance of seedlings from southern sources.

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]

With climate change, favorable growing conditions for tree species are shifting northwards and to higher altitudes. Therefore, local populations are becoming less adapted to their environment. Assisted migration is one of the proposed adaptive measures to reduce the vulnerability of natural populations and maintain forest productivity. It consists of moving genetic material to a territory where future climate conditions correspond to those of its current location. Eight white spruce (Picea glauca [Moench] Voss) seed sources representing as many seed orchards were planted in 2013 at three forest sites simulating a south-north climatic gradient of 1.7°C in Québec, Canada. The objectives were to (1) evaluate the morpho-physiological responses of the different seed sources and (2) determine the role of genetic adaptation and physiological plasticity on the observed variation in morpho-physiological traits. Various seedling characteristics were measured, notably height growth from nursery to the fourth year on plantation. Other traits such as biomass and carbon allocation, nutritional status, and various photosynthetic traits before bud break, were evaluated during the fourth growing season. No interaction between sites and seed sources was observed for any traits, suggesting similar plasticity between seed sources. There was no change in the rank of seed sources and sites between years for height growth. Moreover, a significant positive correlation was observed between the height from the nursery and that after 4 years in the plantation. Southern seed sources showed the best height growth, while optimum growth was observed at the central site. Juvenile height growth seems to be a good indicator of the juvenile carbon sequestration and could serve as a selection criterion for the best genetics sources for carbon sequestration. Vector analysis showed no nitrogen deficiency 4 years after planting. Neither seed sources nor planting sites had a significant effect on photosynthesis before bud break. The observed results during the establishment phase under different site conditions indicate that southern seed sources may already benefit from assisted migration to cooler climatic conditions further north. While northern seed sources are likely to benefit from anticipated local global warming, they would not match the growth performance of seedlings from southern sources. [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]

In the absence of large-scale stand replacing disturbances, boreal forests can remain in the old-growth stage over time because of a dynamic equilibrium between small-scale mortality and regeneration processes. Although this gap paradigm has been a cornerstone of forest dynamics theory and practice for decades, evidence suggests that it could be disrupted, threatening the integrity and sustainability of continuous forest cover. The objective of this study was to evaluate the gap dynamics in old-growth boreal forests across a large landscape where deer populations currently exist at high abundance. We hypothesized that chronic deer browsing is limiting recruitment, particularly of palatable species, creating a demographic disequilibrium between canopy mortality and recruitment. We analysed understory regeneration density and distribution in relation to canopy gap size and condition on multiple sample areas within a 360 km 2 area of old-growth balsam fir ( Abies balsamea [L.] Miller) forest on Anticosti Island, Canada. The combined effect of accelerating canopy gap expansion and recruitment failure created a demographic disequilibrium important enough to cause a loss of forest cover. The forest is now at risk of shifting to alternative successional pathways that seem to be dependent upon gaps size. Rather than sustaining historic balsam fir composition, sucession in 57% of gap area was more susceptible to following a pathway leading toward white spruce parklands, while sucession in the other 43% was more susceptible to following a pathway toward white spruce forests. The occurrence of these novel ecosystems represents a threat to biodiversity and ecosystem services that are provided by preindustrial forests. Climate change could exacerbate these threats by allowing deer to go into as yet unoccupied boreal forests that are driven by gap dynamics. Novel management issues will arise in these boreal ecosystems and challenge forest managers. When the traditional approaches of identifying gaps will not work because the forest itself is losing cover, the method we have developed will help forest managers recognize demographic disequilibrium threatening maintenance of forests. [ABSTRACT FROM AUTHOR]

Insufficient chilling resulting from rising winter temperatures associated with climate warming has been an area of particular interest in boreal and temperate regions where a period of cool temperatures in fall and winter is required to break plant dormancy. In this study, we examined the budburst and growth of trembling aspen (Populus tremuloides Michx.), balsam poplar (Populus balsamifera L.), white birch (Betula papyrifera Marsh.), black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss), jack pine (Pinus banksiana Lamb.), and lodgepole pine (Pinus contorta Dougl. ex. Loud.) seedlings subjected to typical northern Ontario, Canada, spring conditions in climate chambers after different exposures to natural chilling. Results indicate that chilling requirements (cumulative weighted chilling hours) differed substantially among the seven species, ranging from 300 to 500 h for spruce seedlings to more than 1100 h for trembling aspen and lodgepole pine. Only spruce seedlings had fulfilled their chilling requirements before December 31, whereas the other species continued chilling well into March and April. Species with lower chilling requirements needed more heat accumulation for budburst and vice versa. Insufficient chilling delayed budburst but only extremely restricted chilling hours (<400) resulted in abnormal budburst and growth, including reduced needle and shoot expansion, early budburst in lower crowns, and erratic budburst on lower stems and roots. Effects, however, depended on both the species' chilling requirements and the chilling-heat relationship. Among the seven tree species examined, trembling aspen is most likely to be affected by reduced chilling accumulation possible under future climate scenarios, followed by balsam poplar, white birch, lodgepole pine, and jack pine. Black and white spruce are least likely to be affected by changes in chilling hours. [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]

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]

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]

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]

• 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]

Because of changes in climatic conditions, tree seeds originating from breeding programs may no longer be suited to sites where they are currently sent. As a consequence, new seed zones may have to be delineated. Assisted migration consists of transferring seed sources that match the future climatic conditions to which they are currently adapted. It represents a strategy that could be used to mitigate the potential negative consequences of climate change on forest productivity. Decisions with regard to the choice of the most appropriate seed sources have to rely on appropriate knowledge of morpho-physiological responses of trees. To meet this goal, white spruce (Picea glauca [Moench] Voss) seedlings from eight seed orchards were evaluated during two years in a forest nursery, and at the end of the first growing season on three plantation sites located in different bioclimatic domains in Quebec.The morpho-physiological responses obtained at the end of the second growing season (2+0) in the nursery made it possible to cluster the orchards into three distinct groups. Modelling growth curves of these different groups showed that the height growth of seedlings from the second-generation and southern first-generation seed orchards was significantly higher than that of those from other orchards, by at least 6%. A multiple regression model with three climatic variables (average growing season temperature, average July temperature, length of the growing season) showed that the final height of seedlings (2+0) from the first-generation seed orchards was significantly related to the local climatic conditions at the orchard sites of origin where parental trees from surrounding natural populations were sampled to provide grafts for orchard establishment. Seedling height growth was significantly affected by both seed source origins and planting sites, but the relative ranking of the different seed sources was maintained regardless of reforestation site. This knowledge could be used, in conjunction with transfer models, to refine operational seed transfer rules and select the most suitable sites in an assisted migration strategy.

• We assess spring phenology in white spruce to support climate change adaptation. • A range-wide provenance trial revealed a latitudinal cline in heatsum requirements. • Chilling requirements were moderate and genetically undifferentiated. • The results suggest that assisted migration of southern sources poleward is safe. • Climate warming should not de-synchronize bud break from the growing season. The timing of spring dormancy release and bud break in trees is an adaptive trait with potentially important management implications. Depending on how spring phenology is controlled, climate warming may disrupt the synchronization of bud break with the available growing season. Further, addressing climate change through human assisted migration in reforestation programs could cause additional problems if phenology triggers of source and target locations differ. Here, we assess how phenology is controlled by heat sum and chilling requirements for a widespread and commercially important boreal tree species, white spruce (Picea glauca) in a range-wide common garden experiment. We find significant genetic population differentiation in heat sum requirements ranging from 390 to 450 degree days among regions (±12 average SE), and from 375 to 500 (±16) degree days among populations within regions. The most northern populations showed the lowest heat sum requirements, which could be explained by frost avoidance or growth optimization strategies. Our data favors a frost avoidance hypothesis, although the explanations are not mutually exclusive. Chilling requirements in white spruce were also found (approx. 15 degree days), but they showed no geographic population differentiation. Due to generally low chilling requirements, we do not anticipate de-synchronization of spring phenology with the growing season under climate warming in spruce. Synthesizing results from this study and other research on additional growth and adaptive traits, we conclude that assisted migration northward, compensating for approximately 2° C warming, is safe and yields higher growth rates for white spruce. [ABSTRACT FROM AUTHOR]