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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]

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]

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]

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

Ongoing climate change is likely to result in shifts in successional dynamics in boreal mixedwood stands. Using data from provincial forest inventory databases, we examined the occurrence and abundance of the regeneration of various coniferous species (white spruce, black spruce and balsam fir) along an east-west Canadian gradient in aspen-dominated stands. The interpretation of the results was based on environmental conditions, including climate, natural fire regime and human impacts. We found that conifer regeneration was present in aspen stands along the entire gradient, despite differences in climatic conditions and fire regimes between the west (warmer and drier, with large recurrent fires) and east (more humid with relatively long fire cycles). However, abundance and distribution varied from one conifer species to the next. The abundance of white spruce decreased towards the eastern end of the longitudinal gradient, while balsam fir and black spruce abundance decreased towards the west. Although abundance decreased, balsam fir and black spruce regeneration was still present in western Canada. This study shows that it is difficult to interpret the effects of climate change on conifer recruitment without accounting for the superimposed effects of human activities. [ABSTRACT FROM AUTHOR]

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

Abstract: This paper examines the variability in the relationship between climate and radial growth of black spruce (Picea mariana (Mill.) B.S.P.) and white spruce (Picea glauca (Moench) Voss) trees across central Labrador, Eastern Canada. Using climate-sensitive trees, an 11-year running Pearson correlation is applied to local records to examine the relationship between radial tree growth and climate over the last 50 years and the spatial pattern in this relationship with increasing distance inland from the Labrador Sea. Results indicate that there is a high degree of instability in the climate/tree-ring sensitivity despite an overall statistically significant relationship throughout the instrumental time period (1942 to present). Although some periods of reduced climate sensitivity are coincident with insect outbreaks, others cannot be explained by forest disturbance factors. Spatially, the two sites that are most representative of higher elevation areas have more time-stable climate-growth relationships than those inland or along the coast. The results also suggest that the stability of the relationship may be the result of moisture availability, rapid changes in precipitation and temperature, and site-specificity. [Copyright &y& Elsevier]

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

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

Models of forest growth and yield (G&Y) are a key component in long-term strategic forest management plans. Models leveraging the industry-standard "empirical" approach to G&Y are frequently underpinned by an assumption of historical consistency in climatic growing conditions. This assumption is problematic as forest managers look to obtain reliable growth predictions under the changing climate of the 21st century. Consequently, there is a pressing need for G&Y modelling approaches that can be more robustly applied under the influence of climate change. In this study we utilized an established forest gap model (JABOWA-3) to simulate G&Y between 2020 and 2100 under Representative Concentration Pathways (RCP) 2.6, 4.5, and 8.5 in the Canadian province of Newfoundland and Labrador (NL). Simulations were completed using the province's permanent sample plot data and surface-fitted climatic datasets. Through model validation, we found simulated basal area (BA) aligned with observed BA for the major conifer species components of NL's forests, including black spruce [Picea mariana (Mill.) Britton et al.] and balsam fir [Abies balsamea (L.) Mill]. Model validation was not as robust for the less abundant species components of NL (e.g., Acer rubrum L. 1753, Populus tremuloides Michx., and Picea glauca (Moench) Voss). Our simulations generally indicate that projected climatic changes may modestly increase black spruce and balsam fir productivity in the more northerly growing environments within NL. In contrast, we found productivity of these same species to only be maintained, and in some instances even decline, toward NL's southerly extents. These generalizations are moderated by species, RCP, and geographic parameters. Growth modifiers were also prepared to render empirical G&Y projections more robust for use under periods of climate change. [ABSTRACT FROM AUTHOR]

• Both nutrient supply and root interaction between seedlings influenced responses to CO 2 elevation. • CO 2 stimulation to photosynthesis was much greater under low than high nutrient supply. • CO 2 elevation substantially reduced low nutrient limitation to photosynthesis. • The degrees of responses were very different between black spruce and white spruce. The carbon dioxide concentration ([CO 2) in the atmosphere has been increasing since the start of the industrialization. The impact of increasing [CO 2 on the physiology and growth of plants and on the composition, dynamics and productivity of plant communities have been studied extensively in the past few decades. However, the responses to CO 2 elevation can be altered by other environmental and biological factors. Such interactions, particularly those involving multiple factors, are complex and not well understood. In this study, we investigated the interactive effects of CO 2 elevation, nutrient availability and root interaction between plants on the total seedling photosynthesis, growth, and biomass allocation of two boreal conifer tree species, black spruce and white spruce. Seedlings were exposed to two [CO 2 (380 vs. 720 µmol mol−1) and two nutrient levels (low vs. high) with or without root interaction. We found that each of the three treatments influenced the effects of others and the two species also responded differently to those interactions. Root interactions reduced seedling growth only when nutrient supply was high. White spruce was more sensitive root interaction than black spruce while black spruce was more sensitive to the negative impact of low nutrient supply than white spruce. Surprisingly, CO 2 elevation and its interactions did not significantly affect seedling growth. The CO 2 elevation significantly increased total seedling photosynthesis while root interaction reduced it and the reduction was greater under the elevated CO 2. The photosynthetic response to CO 2 elevation was more sensitive to nutrient availability in white spruce than in black spruce. Seedling biomass was more sensitive to the negative impact of root interaction and low nutrient supply in black spruce than white spruce. Root interaction reduced biomass allocation to leaf and the impact was reduced by low nutrient. Biomass allocation to leaf was more sensitive to low nutrient supply in black spruce than white spruce. CO 2 elevation reduced but root interaction increased root/leaf mass ratio, partially offsetting each other when both treatments were applied. The root/leaf ratio of white spruce was more sensitive to CO 2 elevation while that of black spruce was more sensitive to root interaction and nutrient supply. Low nutrient supply and root interaction both had positive effect on root/leaf ratio and their effects were confounded when both were applied simultaneously. Therefore, their effects were observed only when one of them was absent. [ABSTRACT FROM AUTHOR]