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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]

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

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]

Merging robust statistical methods with complex simulation models is a frontier for improving ecological inference and forecasting. However, bringing these tools together is not always straightforward. Matching data with model output, determining starting conditions, and addressing high dimensionality are some of the complexities that arise when attempting to incorporate ecological field data with mechanistic models directly using sophisticated statistical methods. To illustrate these complexities and pragmatic paths forward, we present an analysis using tree‐ring basal area reconstructions in Denali National Park (DNPP) to constrain successional trajectories of two spruce species (Picea mariana and Picea glauca) simulated by a forest gap model, University of Virginia Forest Model Enhanced—UVAFME. Through this process, we provide preliminary ecological inference about the long‐term competitive dynamics between slow‐growing P. mariana and relatively faster‐growing P. glauca. Incorporating tree‐ring data into UVAFME allowed us to estimate a bias correction for stand age with improved parameter estimates. We found that higher parameter values for P. mariana minimum growth under stress and P. glauca maximum growth rate were key to improving simulations of coexistence, agreeing with recent research that faster‐growing P. glauca may outcompete P. mariana under climate change scenarios. The implementation challenges we highlight are a crucial part of the conversation for how to bring models together with data to improve ecological inference and forecasting. [ABSTRACT FROM AUTHOR]

Numerous recent studies have argued that moisture limitation is leading to growth declines and mortality of black spruce (Picea mariana) and white spruce (Picea glauca) in the North American boreal forest. A parallel line of research suggests that increasingly common severe wildfires are altering successional pathways and leading to long‐term replacement of spruce forests with those dominated by paper birch (Betula papyrifera, Betula neoalaskana) and trembling aspen (Populus tremuloides). When both conifers and deciduous trees establish after fire, this biome shift hypothesis implicitly assumes that deciduous species will outcompete the conifers, owing to their more rapid vertical growth and because they might be less sensitive to warm and dry conditions.We established a research site in a white spruce‐paper birch forest on an east‐facing slope in interior Alaska and tested the hypothesis that Alaska paper birch are better adapted to warm and dry conditions than white spruce. Over 6 years (2013–2018), we made hourly measurements of microclimate and xylem sap flux of both species. We also collected increment cores and conducted climate‐growth analyses for both species.During our 6‐year study, growing seasons with low volumetric soil water content (SWC) were those that followed shallow winter snowpacks and had limited summer rainfall, not necessarily those with warm air temperature. Both species were sensitive to moisture limitation. The tree‐ring data revealed significant positive effects of cumulative water year precipitation on radial growth, with a steeper slope for paper birch than for white spruce. Radial growth of both species was also positively related to mean water year air temperature. Sap flux density declined progressively for white spruce over the range of observed SWC and abruptly for paper birch when SWC fell below ~15%.Synthesis. Our results show that, while paper birch might be less sensitive to mild drought than white spruce, it may be more sensitive to severe drought, raising questions about the ability of paper birch to outcompete co‐occurring white spruce in a drier climate. [ABSTRACT FROM AUTHOR]

The use of wood ash as a soil amendment in afforestation and reforestation efforts is increasing. While most studies suggest benefits or neutral results on tree growth and survival, a few studies indicate adverse effects. Hybrid larch, jack pine and white spruce were studied at three northwestern Quebec plantation sites after they received wood ash at two application rates. Soil chemical properties, foliar nutrients and seedling growth and mortality were monitored over a period of eight years. The response of soil to ash application was mostly observed in the forest floor and was more pronounced in year 3 than year 8, likely due to the acidifying nature of the boreal soils studied. Jack pine growth increased linearly with wood ash application rates, white spruce growth showed an inconsistent and delayed positive response under the higher application rate, and hybrid larch growth and survival were either increased or decreased under the lower application rate depending of site but decreased at all sites under the higher application rate. The divergence in growth response between tree species underlines a trade-off between species with rapid acquisition of resources (e.g., pine, larch) to species that use more conservative strategies and store nutrients in their tissues for longer periods (e.g., spruce). In the case of hybrid larch, it accumulated larger amounts of Mn in its needles under the higher application rate and thus, the high bioavailabil - ity of Mn appears to have been detrimental to its survival and growth. Its higher sensitivity to Mn addition from ash is likely due to its highly acquisitive (nutrients) nature compared to other coniferous species as well as the initial levels of available Mn levels in the soil. The contrasted growth responses reported here under similar growing conditions highlight the importance of identifying suitable species, sites and application rates to maximize the benefits of wood ash amendments for future tree plantations in the boreal forest. [ABSTRACT FROM AUTHOR]

Increase in frost damage to trees due to earlier spring dehardening could outweigh the expected increase in forest productivity caused by climate warming. We quantified the impact of growing‐season frosts on the performance of three spruce species (white, black, and Norway spruce) and various seed sources with different frost tolerance in two plantations, established on both sides of the eastern Canadian boreal‐temperate forest ecotone. The objectives of this study were to determine (a) if spruce species and seed sources planted in sites far from their natural provenance would be less adapted to local site conditions, leading to increased frost damage and reduced height growth; (b) at which height above the ground growing‐season frosts ceased to damage apical meristems; and (c) if height growth was best predicted by extreme climatic events (growing‐season frosts) or by mean annual or summer temperature. At each site and for all spruce species and seed sources, we cross‐sectioned spruce trees at different heights above the ground. Tree rings were cross‐dated and screened for frost rings, which were then given a severity score based on cellular damage. Frost severity reduced height growth of all spruce species and provenances at both sites. Height growth of the non‐native Norway spruce was the most reduced by frost severity and was the smallest species at both sites. Frost caused the highest growth reduction in white spruce at the boreal mixedwood site and had the least effect on black spruce at both sites. For all spruce species, height growth was affected up to 2 m above the ground. Model selection based on corrected Akaike's information criteria (AICc) identified that minimum temperature in May was by far the best climate variable predicting tree growth (AICc weight = 1), highlighting the importance of considering extreme climatic events, which are likely to increase in the future. [ABSTRACT FROM AUTHOR]

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

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

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

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]

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

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

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

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

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

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

Long-term experiments provide a way to test presumed causes of successional or environmentally driven vegetation changes. Early-successional nitrogen (N)-fixing plants are widely thought to facilitate productivity and vegetation development on N-poor sites, thus accounting for observed vegetation patterns later in succession. We tested this facilitative impact on vegetation development in a 23-yr field experiment on an Interior Alaska ( USA) floodplain. On three replicate early-successional silt bars, we planted late-successional white spruce ( Picea glauca) seedlings in the presence and absence of planted seedlings of an early-successional N-fixing shrub, thinleaf alder ( Alnus incana). Alder initially facilitated survivorship and growth of white spruce. Within six years, however, after canopy closure, alder negatively affected spruce survivorship and growth. Our three replicate sites followed different successional trajectories. One site was eliminated by erosion and supported no vegetation development during our study. The other two sites, which differed in site moisture, diverged in vegetation composition. Structural equation modeling ( SEM) suggested that, in the drier of these two sites, alder inhibited spruce growth directly (presumably by competition) and indirectly through effects mediated by competition with other woody species. However, at the wetter site, alder had both positive and negative effects on spruce growth, with negative effects predominating. Snowshoe hares ( Lepus americanus) in alder thickets further reduced height growth of spruce in the wetter site. We conclude that net effects of alder on white spruce, the late-successional dominant, were primarily inhibitory and indirect, with the mechanisms depending on initial site moisture. Our results highlight the importance of long-term research showing that small differences among initial replicate sites can cause divergence in successional trajectories, consistent with individualistic distributions of species and communities along environmental gradients. This divergence was detectable only decades later. [ABSTRACT FROM AUTHOR]

The western Canadian mixedwood boreal forests were projected to be significantly affected by regional drought. However, drought degrees were spatially different across elevations, longitudes and latitudes, which might cause different tree growth responses to climate change in different sub-regions within western Canada. In this way, regional classification of western Canadian boreal forests and understanding spatial tree growth responses to climate might be necessary for future forest management and monitoring. In this paper, tree-ring chronologies of two dominant tree species, trembling aspen ( Populus tremuloides Michx.) and white spruce ( Picea glauca (Moench.) Voss), were obtained from mixed forest stands distributed across western Canada to study spatial tree growth response to climate based on three regional classification schemes (a phytogeographic sub-region classification, a natural sub-region classification and non-classification). Phytogeographic sub-region classification was estimated based on tree ring samples we collected in this study, while natural sub-region classification was previously developed based on analysis of regional differences in vegetation, soil, site and climate conditions. Results showed that air temperature did not significantly increase, while drought stress became more severe between 1985 to 2010. Relationships between trembling aspen growth and temperature differed between north and south parts of the study area, resulting from spatial difference in water supply. Trembling aspen growth was influenced by temperature or moisture variables of the previous years. White spruce growth was influenced primarily by moisture variables (current or previous year), and response coefficients between white spruce and drought conditions (represented by drought code) were negative in all phytogeographic sub-regions, suggesting that white spruce was more sensitive to drought stress under climate change. As a late-successional dominant species, increasingly drought stress on white spruce might cause significant alteration in forest composition of western Canadian boreal forest. [ABSTRACT FROM AUTHOR]

[Display omitted] • Boreal forests evaporate considerably more than higher elevation shrub ecosystems. • Forests exist in a growing season water deficit relying on snowmelt recharge. • ET variability declines with increased vegetation cover. • Majority of growing season water for streamflow is generated at higher elevations. • We propose treeline advance will result in drying of northern catchments. As a result of altitude and latitude amplified climate change, widespread changes in vegetation composition, density and distribution have been observed across northern regions. Despite wide documentation of shrub proliferation and treeline advance, few field-based studies have evaluated the hydrological implications of these changes. Quantification of total evapotranspiration (ET) across a range of vegetation gradients is essential for predicting water yield, yet challenging in cold alpine catchments due to heterogeneous land cover, including both boreal forest and shrub taiga ecosystems. Here, we present six years of surface energy balance components and ET dynamics at three sites along an elevational gradient in a subarctic, alpine catchment near Whitehorse, Yukon Territory, Canada. These sites span a gradient of thermal and vegetation regimes, providing a space-for-time comparison for future ecosystem shifts: 1) a low-elevation boreal white spruce forest (~12–20 m), 2) a mid-elevation subalpine taiga comprised of tall, dense willow (Salix) and birch (Betula) shrubs (~1–3 m) and 3) a high-elevation subalpine taiga with short, sparse shrub cover (<0.75 m) and moss, lichen, and bare rock. Eddy covariance instrumentation ran year-round at the forest and during the growing season at the two shrub sites. Total ET decreased and interannual variability increased with elevation, with mean May to September ET totals of 349 (±3) mm at the forest, 249 (±10) mm at the tall, dense shrub site, and 240 (±26) mm at the short, sparse shrub site. Comparatively, over the same period, ET:R ratios were the highest and most variable at the forest (2.19 ± 0.37) and similar at the tall, dense shrub (1.22 ± 0.09) and short, sparse shrub (1.14 ± 0.05) sites. Our results suggest that advances in treeline will increase overall ET and lower interannual variability; however, the large growing season water deficit at the forest indicates strong reliance on soil moisture from late fall and snowmelt recharge. In contrast, ET was considerably less at the cooler higher elevation shrub sites , which exhibited similar ET losses over 6 years despite differences in shrub height and abundance. ET rates between the two shrub sites were similar throughout the year, except during the peak growing season. Greater interannual variability in ET at the short, sparse shrub site indicates the reduced influence of vegetation controls on ET. Results suggest that predicted changes in vegetation type and structure in northern regions will have a considerable impact on water partitioning and will vary in a complex way in response to changing precipitation timing, phase and magnitude, growing season length, and vegetation snow and rain interactions. [ABSTRACT FROM AUTHOR]

We investigated habitat attributes related to the occupancy of the globally rare and endangered epiphytic lichen, Erioderma pedicellatum, in a newly discovered (2009) population center in Denali National Park and Preserve (DNPP), Alaska. We measured forest, tree and epiphytic lichen community characteristics on eighty-five systematically selected plots in four study areas. We aggregated these data at three spatial scales (tree, plot and study area) at which to compare E. pedicellatum occupancy (probability of occurrence) and abundance to environmental covariates. We observed 2,035 E. pedicellatum thalli on 278 individual Picea glauca stems. The species occurred in 61% of the plots measured. Occupancy of E. pedicellatum at the individual tree-scale was influenced by stem diameter, study area, live crown length, plot P. glauca basal area, plot canopy cover and distance to open water. Our models for E. pedicellatum occupancy at the plot-scale identified study area, P. glauca density, deciduous basal area (Betula neoalaskana and Populus trichocarpa), and tall shrub cover (≥ 200 cm in height) as significant covariates. Our estimates of the DNPP population size increase the world population by at least tenfold to approximately 100,000 thalli. We suggest important topics for further research on E. pedicellatum. [ABSTRACT FROM AUTHOR]

White spruce (Picea glauca [Moench] Voss) is a valuable commercial species found in interior and southceniral Alaska. Numerous regional and local volume tables or equations exist; however, no statewide model exists or has been tested for accuracy. There is a demand for an accurate model to determine the cubic-foot volume of white spruce trees in Alaska. Multiple models were developed for white spruce to estimate total and merchantable cubic-foot volume to a 2-, 4-, and 6-in, top. These multiple-entry (diameter and height) models were developed for both inside and outside bark volume from a 6-in, stump. The models were tested on a regional basis at various geographic locations and were shown to be highly accurate. The Alaska models chosen have R2 at or near 0.99 and mean square error from 0 to 0.16 for all models. These models are shown to be superior to other white spruce models in Alaska. [ABSTRACT FROM AUTHOR]

Abstract: Landscape level factors such as overstory canopy composition can have a profound effect on the ecology of microbial communities in boreal forest floors. However, factors influencing community composition at the microsite scale are still poorly described and understood. Here we explored moisture effects on microbial communities in forest floor derived from undisturbed trembling aspen and white spruce stands, two of the dominant trees in the boreal forest of western Canada. Forest floor samples were incubated in a laboratory experiment for a period of one month under a moisture regime ranging from moist to dry (field capacity, 60% of field capacity and wilting point). As in previous studies we found that the origin of the forest floor material had a strong effect on the microbial community. Additionally, we found that moisture manipulation did not alter the microbial communities of the white spruce forest floor. On the other hand, the moisture had a profound effect on the aspen forest floor, and resulted in structurally and functionally distinct microbial communities. This different response to moisture could be linked to the adaptation of microbial groups to the physical environment inherent to the aspen and spruce forest floors and provides an avenue to further work into microbial mediated biogeochemical processes in the boreal forest. [Copyright &y& Elsevier]

* Premise of the study: In ecosystems where seed production is low and masting years are sporadic, or with species that have short-lived seeds, regeneration is assured by seedling banks rather than seed banks. Seedling establishment and survival play a critical role in determining the composition of these plant communities by supplying new individuals for their maintenance. Seedling emergence and mortality were investigated to test the hypothesis that recruitment into the seedling bank is periodic. * Materials and methods: Seed production and seedling emergence and survival was monitored during 1994-2007 in balsam fir (Abies balsamea) and white spruce (Picea glauca) in four pristine stands of the boreal forest of Quebec, Canada. Measurements were collected twice per month by sampling one permanent plot of 20 x 20 m per stand. * Key results: Seed-rain abundance reached 9 x 10³ seeds m-2 year-1, and was characterized by synchronous sequences of low and high seed production. New seedlings appeared only during the year following a seed production of at least 1 x 10³ and 1.5 x 10² seeds m-2 year-1 for balsam fir and white spruce, respectively. Seedlings emerged in July and survived 34-52 d on average, with balsam fir showing a longer lifespan and lower mortality, although 85-99% of seedlings died before completing one year of life. * Conclusions: The emergence of young seedlings was coupled with massive seed rains, which allowed synchronous replenish ment of the seedling banks among stands and species, and generated different cohorts, yielding a discontinuous age structure. [ABSTRACT FROM AUTHOR]

The northernmost balsam fir forest in eastern Canada forms disjunct stands far beyond the extensive balsam fir forest zone of southern Canada. The northern balsam fir stands are distributed in the subalpine belt of high plateaus and coexist locally with white spruce stands. These subalpine stands contrast greatly with black spruce forest stands located in lowlands. Given that subalpine stands are remnants of an earlier northern expansion of the balsam fir forest, the main objective of this study is to assess whether white spruce stands are distinct communities having diverged from the balsam fir forest community earlier in the Holocene or if they rather correspond to a different stage of the chronosequence within the subalpine belt. Macrofossil analysis of charcoal in mineral soils was used to compare the stand-scale fire histories and taxonomic fossil composition of subalpine, old-growth balsam fir stands and white spruce stands. No significant differences of mean number of observed fires (mean = 6.35 fires per site), Holocene fire recurrence at the landscape scale and mean fire-return interval (mean = 580 years) were found between white spruce stands and balsam fir stands. The botanical composition of charcoal fragments from mineral soils showed that Abies, Betula and Picea were present throughout the fire period from 5600 cal. BP to present, and no difference was found in the fossil composition of the balsam fir and white spruce stands. No historical change in the botanical composition of charcoal from soils of both stand types was observed indicating that the initial floristic composition remained through the period of recurrent fires. Charcoal data suggest that white spruce stands are not divergent community types. Rather, the two community types are arranged along a chronosequence of different successional stages within the subalpine relict flora. [ABSTRACT FROM AUTHOR]

Aim The main objective of this study was to investigate the origin of white spruce ( Picea glauca) range discontinuities in eastern North America. A first analysis aimed at uncovering the glacial origin and post-glacial migration route of white spruce in the eastern part of the range. A second analysis aimed at evaluating whether disjunct white spruce populations in central Québec result from a recent expansion or are remnant stands following a northern range contraction. Location The first analysis covered the North American transcontinental boreal distribution of white spruce, and the second focused on the eastern part of its range. Methods A total of 589 P. glauca individuals from 51 populations, 24 of which were from a previous published source, were analysed for variation in three chloroplast DNA regions ( trnT/L, trnL/F and ndhK/C). At the continental scale, haplotypic diversity, zones of genetic discontinuities, clustering analysis and analysis of molecular variance (AMOVA) were employed. In the eastern part of the range, haplotypic diversity was assessed with a permutation procedure, demographic history with mismatch analyses, and genetic structure with AMOVA. Results At the continental scale, various lines of evidence (genetic-diversity hotspots, endemic haplotypes, zones of genetic discontinuities, and pattern of molecular variation) show that different lineages migrated from three glacial refugia (Beringian, Mississippian, east Appalachian). Strong genetic discontinuities, low diversity and lack of evidence for demographic expansion were found among stands from isolated high hills in central Québec. Main conclusions White spruce in the eastern part of its range has an east Appalachian origin, a lineage genetically distinct from populations west of the Great Lakes. Previous studies have shown that the northward migration of white spruce is still ongoing as the northernmost populations expand into the tundra. In contrast, mixed white spruce and balsam fir ( Abies balsamea) stands from isolated high hills in central Québec have already reached their Holocene maxima and are currently declining. This range contraction is probably a result of increased wildfire disturbance, which gradually creates a pattern of regional white spruce isolation and replacement by fire-prone black spruce ( Picea mariana) forest. [ABSTRACT FROM AUTHOR]

1. Mast seeding is the synchronous and highly variable production of seed by a population of plants. Mast seeding results from the behaviour of individuals; however, little is known about the synchrony of individuals at local scales. 2. We address two primary questions at a within-population (17–36 ha study plots) and individual level: (i) How variable is seed production between and within years? (ii) How synchronized is seed production between individuals? 3. We monitored annual cone production of 356 Picea glauca (white spruce) from 1990 to 2005 within four plots spanning a total distance of 5.3 km in the Yukon Territory, Canada. 4. Spearman correlations ( rs) were conducted to test for synchrony. Overall, the trees were moderately synchronous (mean rs (± SE) of 0.52 ± 0.14), and synchrony was statistically detectable ( rs > 0) over all distances. Individuals < 75 m apart were highly synchronous (0.64 ± 0.18), and correlations dropped to 0.33 ± 0.07 for trees > 3 km apart. There was considerable variation in cone production patterns among pairs of individuals. 5. The number of mast years per plot varied from one to three. During a mast year, many individuals within plots produced large cone crops, with more variability between individuals in low mean cone years. Individual trees had dominant endogenous cycles varying from none to 1–5 years. Forty-four per cent of trees had no significant lag, 23% a negative 1-year lag, and 20% a positive 3-year lag. Basal area did not influence lags, but trees with higher mean cone production throughout the study were more likely to have a 3-year lag compared with no lag. 6. The scale of highest synchrony coincided with the scale at which the dominant seed predator in the area, the territorial red squirrel ( Tamiasciurus hudsonicus), operates. This may be the scale at... [ABSTRACT FROM AUTHOR]

The northern Québec-Labrador tree lines are the most climatically stressed tree ecosystems of eastern North America. In particular, white spruce (Picea glauca) tree line populations distributed between 56° N and 58° N and 61° W and 66° W show contrasted responses to recent changes in climate according to their geographic position relative to the Labrador Sea. Along the coast, the northernmost latitudinal and altitudinal tree lines responded positively to warming over the last 50 years with invading spruce several tens of meters above the current tree line. In contrast, white spruce tree lines across the wind-exposed Labrador plateau are located much higher in altitude and have receded a few tens of meters beginning around AD 1740-1750 and have not yet recovered. Whereas no field evidence of recent fire and insect damage was found, all inland tree line stands were progressively damaged likely due to mechanical defoliation of wind-exposed trees. Massive tree death in the 19th century caused a reduction in the number of seed-bearing trees, and declining tree lines were not replenished by seedlings. Recent warming reported for northern latitudes has not been strong enough to change the regressive tree line trajectory in interior Labrador. However, white spruce expansion above coastal tree line in the northernmost forest site in Labrador is in line with current climatic trends. It is hypothesized that the species is still advancing toward its potential tree line higher in altitude due to delayed postglacial migration. The slow advance of white spruce in northernmost coastal Labrador is likely caused by the rugged topography of the Torngat-Kaumajet-Kiglapait mountains. [ABSTRACT FROM AUTHOR]

An index of tree growth based on the mean growth curve of black spruce shows that tree growth in the boreal forest of central Canada decreases linearly as the forest-tundra boundary is approached. A theoretical tree line is defined by black spruce growth data. The distribution of tree growth and the theoretical tree line indicate that the forest-tundra boundary is the mean summer position of the arctic front. The growth of white spruce is briefly considered, but because of a small sample size, no conclusions are drawn. [ABSTRACT FROM AUTHOR]

A white spruce seedling, Picea glauca (Moench) Voss, was found at the northern edge of the Brooks Range in Alaska, more than 50 km north of the latitudinal tree line. The seedling, 19 cm tall and about nine years old, was growing at the side of the Dalton Highway to Prudhoe Bay. It most likely sprouted from a seed transported across the Brooks Range on a vehicle and has survived on the well-drained gravel road berm, where site conditions are more favorable for germination and survival than in the surrounding tundra. This spruce has survived for about a decade under current climatic conditions. Even with a warming climate, natural seed dispersal is severely hampered by the rugged topographic barrier of the Brooks Range. Considering the amount of vehicle traffic on the Dalton Highway, however, it is likely that more pioneering spruce seedlings will turn up along this corridor. Once over the Brooks Range, a spruce population can potentially develop and expand. [ABSTRACT FROM AUTHOR]