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We assessed the impacts of three approaches to thinning from below with varying spatial patterns on several stand and individual tree variables for interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco), interior spruce (Picea glauca (Moench) Voss × Picea engelmannii Engelm.), and lodgepole pine (Pinus contorta Dougl. Ex Loud. var. latifolia Englem.) in central British Columbia, Canada. The three thinning treatments were two experimental "clumped" treatments (3 m Clumped and 5 m Clumped) and the Standard (more uniform spacing) thinning treatment that was employed operationally at that time. We used long-term data from 24 plots measured five times over 21 years. Thinning increased stand basal area increment, with the plots that received the 5 m Clumped treatment having significantly higher periodic annual relative basal area increment than the unthinned Control plots. The responses for the two clumped treatments were not any lower than the Standard. The 3 m Clumped treatment was best if one is concerned about fast recovery of the growing space; however, the 5 m Clumped spacing treatment may be preferable if higher individual tree vigour is needed for resistance and resilience to fire, insects, and disease. [ABSTRACT FROM AUTHOR]

Data collected over a 30-year period from an experiment replicated across 21 locations in western Canada are used to explore the effects of precommercial thinning of trembling aspen to a range of densities in combination with three initial white spruce densities on tree growth and stand dynamics. Increasing differentiation amongst the 15 treatments was observed with age after thinning for both spruce and aspen responses. Spruce height and diameter declined with increasing aspen density. At age 10 spruce diameter with no aspen was 1.5× that of spruce in unthinned while it was 2.6× that of spruce in unthinned at age 28. Following thinning aspen DBH and crown width declined with increasing density of retained aspen while slenderness and height to crown base of aspen increased. Thinning to 1500 aspen·ha−1 resulted in aspen DBH that was 22% larger relative to the unthinned at age 28. Spruce volume and stemwood biomass at age 90, estimated using the Mixedwood Growth Model (MGM21), declined with increasing initial aspen density, with the complete removal of aspen resulting in nearly double the spruce volume of unthinned plots. However, total stand volume, total stand biomass, and stemwood carbon at age 90 are predicted to be largest in mixed stands with aspen densities of 1500 stems·ha−1 or higher. [ABSTRACT FROM AUTHOR]

The European spruce bark beetle Ips typographus and the North American spruce beetle Dendroctonus rufipennis cause high mortality of spruces on their native continents. Both species have been inadvertently transported beyond their native ranges. With similar climates and the presence of congeneric spruce hosts in Europe and North America, there is a risk that one or both bark beetle species become established into the non-native continent. There are many challenges that an introduced population of bark beetles would face, but an important prerequisite for establishment is the presence of suitable host trees. We tested the suitability of non-native versus native hosts by exposing cut bolts of Norway spruce (Picea abies), black spruce (Picea mariana) and white spruce (Picea glauca) to beetle attacks in the field in Norway and Canada. We quantified attack density, brood density and reproductive success of I. typographus and D. rufipennis in the three host species. We found that I. typographus attacked white and black spruce at comparable densities to its native host, Norway spruce, and with similar reproductive success in all three host species. In contrast, D. rufipennis strongly preferred to attack white spruce (a native host) but performed better in the novel Norway spruce host than it did in black spruce, a suboptimal native host. Our results suggest that I. typographus will find abundant and highly suitable hosts in North America, while D. rufipennis in Europe may experience reduced reproductive success in Norway spruce. [ABSTRACT FROM AUTHOR]

The non-destructive testing of wood fibre properties is crucial for informing forest management decisions and achieving optimal resource utilization. Moisture content (MC) is an important indicator of wood freshness and may reveal the presence of wood degradation. However, efficient methods are still needed to better monitor this property along the forest–wood value chain. The objective of the study was to develop prediction models to evaluate log MC based on the propagation of ground penetrating radar (GPR) signals. A total of 165 trees representing four species (black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss), red spruce (Picea rubens Sarg.), and balsam fir (Abies balsamea (L.) Mill.)) were harvested in two regions of the province of Quebec. GPR signals were acquired in the green (fresh) state and at three subsequent drying stages. Partial least squares regression (PLSR) and locally weighted PLSR (LWPLSR) were employed to establish relationships between GPR signals (antenna frequency: 1.6 GHz) and log properties. The models were fitted on three calibration sets containing four drying stages and different species mixes. The LWPLSR models performed better than the PLSR models for predicting log MC, with a lower root mean square error (RMSEp range: 10.8%–20.2% vs. 13.0%–20.5%) and a higher R2p (0.63–0.87 vs. 0.62–0.82). Spruce-only models performed considerably better than fir-only models while multi-species models were in-between. Despite the complex anisotropy of wood and the physics of wave propagation, the GPR technology can be successfully used to estimate log moisture content, but the GPR-based MC models should be calibrated for each specific type of wood material. [ABSTRACT FROM AUTHOR]

As forest managers increasingly seek to implement partial-harvest silviculture systems in complex forests, more data are required on the potential development of the residual stand following treatment. In this study, we present 25-year data from a long-term silvicultural study of tree- and stand-level responses to differing levels of basal area following partial harvest in a complex spruce–fir forest in central British Columbia, Canada. Reducing stand basal area elicited stronger basal area increment response in fir than spruce; however, spruce productivity was overall higher than fir across all treatments. Smaller trees increased their size proportionally more than larger trees, and thinning resulted in increased recruitment of smaller trees into larger diameter classes. At the stand-level, basal area reductions to well below 20 m2/ha resulted in higher volume and basal area growth over the length of this study, and suggest that optimum stand-level growth may be achieved at lower basal areas than suggested in prior literature. We also found a positive relationship between stand-level residual basal area and mortality. In central British Columbia, stand structure, species composition, and growth rates in complex spruce–fir forests can be maintained through partial-harvest silviculture treatments. [ABSTRACT FROM AUTHOR]

Herbivores shape vegetative communities via numerous mechanisms, including browse. We used vegetation monitoring data from Isle Royale National Park to examine woody species change across a nine‐year interval, coinciding with herbivore escalation. Here, moose and snowshoe hare are the dominant herbivores, while the gray wolf is the apex predator. Our initial sampling period (2010) followed six years of low moose abundance, while our second sampling event (2019) followed a nine‐year escalation in moose density. We tested for change in both saplings and shrubs and compared diameter size distributions of common tree species in three island sections. We found a decline in large saplings of sugar maple, a species limited to the west section of the island. We also saw declines in small saplings of sugar maple, paper birch, and trembling aspen. For some species, including black spruce and white spruce, taxa that are unpalatable to moose, diameter distributions were proportionally larger (i.e., indicative of fewer small individuals) in 2019 than in 2010. In contrast, diameter distributions of black ash in the central section and trembling aspen in the central and east sections of the island were proportionally smaller (indicative of more small individuals) in 2019 than in 2010. We found that ~50% of common shrub taxa were more abundant during the second sampling event, while none declined. Our work here brings to light several unanticipated results. While we demonstrated partial recovery of trembling aspen, likely originating during the earlier period of low moose abundance, our results suggest further changes may be attributable to other island herbivores. The response of black ash, a hydric species, may be stemming from a sixfold increase in active beaver sites, while that of spruce may be attributable to a historic spike in snowshoe hare density. We currently have only a limited understanding of the impacts of the island's lesser studied herbivores on overstory regeneration. Further research on these linkages can inform decisions addressing controls on forest structure. [ABSTRACT FROM AUTHOR]

A significant portion of the harvested land base in western Canada is becoming old enough or entering a phase where thinning is a legitimate forest management option. A comprehensive review of the existing knowledge of commercial thinning (CT) treatments applied to pine and spruce-dominated stands in Alberta was conducted, with particular regard to the intensity, timing of interventions, method, and impacts on crop tree growth responses. Although the geographical focus of this review is Alberta, information on this topic is more complete in other areas of North America and Europe, where there is a long history of density management. In areas of eastern North America, our review revealed that CT from below, with tree removal levels from 27 to 43% of the basal area, could increase total merchantable wood produced from 11 to 60 m3 ha−1 over a rotation, depending on stand age and intensity of thinning. For Alberta conditions, and considering the risks, we conclude that commercial thinning basal area removal should be in the range of 25 to 40%, depending on a variety of factors such as species, wind firmness, and insect or disease incidence and risk. Thinning too aggressively and/or too late will increase the blowdown risk but the literature is fairly consistent in suggesting that live crown ratios should be >40% to maximize the chance of growth response and minimize the blowdown risk. In cases where stands are also threatened by stressors such as drought, wind, and insect or disease outbreaks, CT treatments likely offer the potential at limiting the overall risk, but localized knowledge and experience are critical. It is intended that the information presented may support ongoing and future research trials and growth and yield (G&Y) model development about potential CT treatments to apply and the likely results of practical application to commercial forestry. [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]

Much is still unknown about the growth and physiological responses of trees to global change at the northern treeline. We combined tree‐ring width data with century‐long stable carbon and oxygen isotope records to investigate growth and physiological responses of white spruce at two treeline sites in the Canadian Arctic to concurrent increases in temperature, atmospheric CO2 concentration (ca), and decline in sea ice extent over the past century. The tree‐ring records were assessed during three periods with contrasting climatic conditions: (a) the early 20th century warming, (b) the 1940–1970 cooling period, and (c) the anthropogenic late 20th century warming period. We found opposing growth trends between the two sites, but similar carbon isotope discrimination (Δ13C) and intrinsic water‐use efficiency (iWUE) trajectories. While tree growth (defined as basal area increment) increased at the site nearer to the Arctic Ocean during the 20th century following the rise in temperature and sea ice loss, growth declined after 1950 at the more interior site. At both sites, Δ13C slightly increased over these periods. However, trees showed a nonlinear response to increased ca, shifting after 1970 from a passive stomatal response (i.e., no changes in iWUE) to an active response (i.e., a moderate ∼12% increase in iWUE). Further, our isotope‐based findings do not support the idea that temperature‐induced drought stress caused the divergent growth trends at our treeline sites. This study thus highlights nonlinear and complex physiological and growth adjustments to concomitant changes in temperature, sea ice extent, and ca over the last century at the northern treeline. Plain Language Summary: In the last decades, Arctic ecosystems have experienced unprecedented warmer temperatures, higher atmospheric carbon dioxide concentrations, and a decline in sea ice extent. However, it is still unknown how these changes in environmental factors have impacted tree populations at their northern distribution limits. Here, we examine how the growth and water‐use efficiency of white spruce trees have changed over the last century in response to warmer temperatures, higher carbon dioxide concentrations, and sea ice decline at two treeline sites in the Canadian Arctic. We found that tree growth increased at the site nearer to the Arctic Ocean during the 20th century following the rise in temperature and sea ice loss, while growth declined after 1950 at the more interior site. At both sites, the water‐use efficiency of trees was stable until 1970. Following the rapid temperature and carbon dioxide increase since the 1970s, trees have become more efficient in using water. The contrasting growth trends found at our study sites highlight complex growth and physiological adjustments of white spruce trees to rapid and simultaneous changes in temperature, sea ice extent, and carbon dioxide concentrations in the Arctic. Key Points: Complex growth and physiological responses of white spruce trees to changes in temperature and sea ice extent inferred from stable isotopesWhite spruce radial growth at Arctic latitudinal treeline decreased at an interior site and increased at a site closer to the Arctic OceanWhite spruce shifted from a passive to an active water‐use efficiency response to rising CO2 concentration over the 20th century [ABSTRACT FROM AUTHOR]

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SUMMARY: Spruces (Picea spp.) are coniferous trees widespread in boreal and mountainous forests of the northern hemisphere, with large economic significance and enormous contributions to global carbon sequestration. Spruces harbor very large genomes with high repetitiveness, hampering their comparative analysis. Here, we present and compare the genomes of four different North American spruces: the genome assemblies for Engelmann spruce (Picea engelmannii) and Sitka spruce (Picea sitchensis) together with improved and more contiguous genome assemblies for white spruce (Picea glauca) and for a naturally occurring introgress of these three species known as interior spruce (P. engelmannii × glauca × sitchensis). The genomes were structurally similar, and a large part of scaffolds could be anchored to a genetic map. The composition of the interior spruce genome indicated asymmetric contributions from the three ancestral genomes. Phylogenetic analysis of the nuclear and organelle genomes revealed a topology indicative of ancient reticulation. Different patterns of expansion of gene families among genomes were observed and related with presumed diversifying ecological adaptations. We identified rapidly evolving genes that harbored high rates of non‐synonymous polymorphisms relative to synonymous ones, indicative of positive selection and its hitchhiking effects. These gene sets were mostly distinct between the genomes of ecologically contrasted species, and signatures of convergent balancing selection were detected. Stress and stimulus response was identified as the most frequent function assigned to expanding gene families and rapidly evolving genes. These two aspects of genomic evolution were complementary in their contribution to divergent evolution of presumed adaptive nature. These more contiguous spruce giga‐genome sequences should strengthen our understanding of conifer genome structure and evolution, as their comparison offers clues into the genetic basis of adaptation and ecology of conifers at the genomic level. They will also provide tools to better monitor natural genetic diversity and improve the management of conifer forests. The genomes of four closely related North American spruces indicate that their high similarity at the morphological level is paralleled by the high conservation of their physical genome structure. Yet, the evidence of divergent evolution is apparent in their rapidly evolving genomes, supported by differential expansion of key gene families and large sets of genes under positive selection, largely in relation to stimulus and environmental stress response. Significance Statement: The genomes of four closely related North American spruces indicate that their high similarity at the morphological level is paralleled by the high conservation of their physical genome structure. Yet, the evidence of divergent evolution is apparent in their rapidly evolving genomes, supported by differential expansion of key gene families and large sets of genes under positive selection, largely in relation to stimulus and environmental stress response. [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]

Background: Picea species are distributed and planted world-wide due to their great ecological and economic values. It has been reported that Picea species vary widely in growth traits in a given environment, which reflects genetic and phenotypic differences among species. However, key physiological processes underlying tree growth and the influencing factors on them are still unknown. Results: Here, we examined needle structures, needle chemical components, physiological characteristics and growth traits across five Picea species in a common garden in Tianshui, Gansu province in China: Picea glauca, P. mariana, P. likiangensis, P. koraiensis, and P. crassifolia, among which P. glauca and P. mariana were introduced from North America, P. likiangensis was from Lijiang, Yunan province in China, P. koraiensis was from Yichun, Heilongjiang province in China, and P. crassifolia was native to the experimental site. It was found that nearly all traits varied significantly among species. Tissue-level anatomical characteristics and leaf mass per area (LMA) were affected by needle size, but the variations of them were not associated with the variations in photosynthetic and biochemical capacity among species. Variations in area-based maximum photosynthesis (Pnmax) were affected by stomatal conductance (gs), mesophyll conductance (gm) and biochemical parameters including maximum carboxylation rate (Vcmax), and maximum electron transport rate (Jmax). The fraction of N allocated to different photosynthetic apparatus displayed contrasting values among species, which contributed to the species variations in photosynthetic nitrogen use efficiency (PNUE) and Pnmax. Additionally, all growth traits were positively correlated with Pnmax and PNUE. Conclusion: Needle structures are less important than needle biochemical parameters in determining the variations in photosynthetic capacity across the five Picea species. Pnmax and PNUE are closedly associated with the fraction of N allocated to photosynthetic apparatus (Pphoto) compared with leaf N content per area (Narea). The tremendous growth differences among the five Picea species were substantially related to the interspecies variation in Pnmax and PNUE. [ABSTRACT FROM AUTHOR]

Precise delineation of individual tree crowns is critical for accurate forest biophysical parameter estimation, species classification, and ecosystem modelling. Multispectral optical remote sensors mounted on low-flying unmanned aerial vehicles (UAVs) can rapidly collect very-high-resolution (VHR) photogrammetric optical data that contain the spectral, spatial, and structural information of trees. State-of-the-art tree crown delineation approaches rely mostly on spectral information and underexploit the spatial-contextual and structural information in VHR photogrammetric multispectral data, resulting in crown delineation errors. Here, we propose the spectral, spatial-contextual, and structural information-based individual tree crown delineation (S3-ITD) method, which accurately delineates individual spruce crowns by minimizing the undesirable effects due to intracrown spectral variance and nonuniform illumination/shadowing in VHR multispectral data. We evaluate the performance of the S3-ITD crown delineation method over a white spruce forest in Quebec, Canada. The highest mean intersection over union (IoU) index of 0.83, and the lowest mean crown-area difference (CAD) of 0.14 m 2 , proves the superior crown delineation performance of the S3-ITD method over state-of-the-art methods. The reduction in error by 2.4 cm and 1.0 cm for the allometrically derived diameter at breast height (DBH) estimates compared with those from the WS-ITD and the BF-ITD approaches, respectively, demonstrates the effectiveness of the S3-ITD method to accurately estimate biophysical parameters. [ABSTRACT FROM AUTHOR]

While deployment of white spruce (Picea glauca (Moench) Voss) from seed orchards is increasing in Alberta, genetic gain is only considered in pure spruce stands due to uncertainties in measuring yields in mixedwood stands. To better understand the performance of improved spruce in mixedwood stands, we compared the effects of interspecific competition on growth of improved (1.9% height gain at a 100-year rotation) and unimproved spruce in northeastern Alberta. By age 8 years the improved spruce showed no advantage over the unimproved spruce in either height (1.36 ± 0.36 m vs. 1.42 ± 0.38 m) or diameter (23.68 ± 7.33 mm vs. 25.65 ± 6.85 mm), and the largest diameter trees were found in a nutrient-poor subxeric site. A distance-independent Lorimer's index including tree size ratio, combined with a power function, accounted for most of the growth variation in diameter and height from 2016 to 2017. The unimproved and improved spruce had different growth-competition curves across ecosites, and their height growth was less sensitive to competition than diameter growth. These results highlight several considerations for managing improved spruce, including (i) deploying higher genetic worth seedlots, (ii) developing realized gain trials with a good statistical design, and (iii) developing growth and yield functions for improved spruce in mixedwood stands. [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]

Understory protection harvesting is a form of partial cutting that can be used in aspen (Populus tremuloides Michx.)-dominated stands that have understories of white spruce (Picea glauca (Moench) Voss). This practice involves removing 75% to 85% of the merchantable aspen while minimizing damage to the advance spruce regeneration, in addition to leaving 15% to 25% of the aspen standing to reduce potential windthrow of the spruce understory. In this paper, we summarize results from 18 stands measured 10 to 12 years after understory protection harvest. Diameter growth of spruce increased during the first five years after harvest while height growth increased during the second five-year period (5 to 10 or 7 to 12 years after release). Consistent with other studies, mortality rates of aspen trees ≥7.1 cm DBH (diameter breast height, 1.3 m) averaged 45.0% over the 10–12 year period following harvesting. Spruce mortality averaged 27.5% over the same 10–12 year period. Substantial aspen regeneration was evident across most harvested blocks, with aspen sapling densities 10–12 years from harvest being higher in removal (14,637 stems⋅ha−1) than in buffer areas (6686 stems⋅ha−1) and in extraction trails (7654 stems⋅ha−1). Spruce sapling (>1.3 m height and <4 cm DBH) densities averaged 1140 stems⋅ha−1 in removal areas at ages 10–12, with these trees likely being present as seedlings at the time of harvest. Mixedwood Growth Model projections indicate merchantable volumes averaging 168 m3⋅ha−1 (conifer) and 106 m3⋅ha−1 (deciduous) 70 years from harvest, resulting in MAI (mean annual increment) for this period averaging 2.0 m3⋅ha−1⋅y−1 with MAI for a full 150-year rotation of approximately 2.5 m3⋅ha−1⋅y−1. [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]

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]

The spruce bark beetle Ips typographus is the most damaging pest in European spruce forests and has caused great ecological and economic disturbances in recent years. Although native to Eurasia, I. typographus has been intercepted more than 200 times in North America and could establish there as an exotic pest if it can find suitable host trees. Using in vitro bioassays, we compared the preference of I. typographus for its coevolved historical host Norway spruce (Picea abies) and two non-coevolved (naïve) North American hosts: black spruce (Picea mariana) and white spruce (Picea glauca). Additionally, we tested how I. typographus responded to its own fungal associates (conspecific fungi) and to fungi vectored by the North American spruce beetle Dendroctonus rufipennis (allospecific fungi). All tested fungi were grown on both historical and naïve host bark media. In a four-choice Petri dish bioassay, I. typographus readily tunneled into bark medium from each of the three spruce species and showed no preference for the historical host over the naïve hosts. Additionally, the beetles showed a clear preference for bark media colonized by fungi and made longer tunnels in fungus-colonized media compared to fungus-free media. The preference for fungus-colonized media did not depend on whether the medium was colonized by conspecific or allospecific fungi. Furthermore, olfactometer bioassays demonstrated that beetles were strongly attracted toward volatiles emitted by both con- and allospecific fungi. Collectively, these results suggest that I. typographus could thrive in evolutionary naïve spruce hosts if it becomes established in North America. Also, I. typographus could probably form and maintain new associations with local allospecific fungi that might increase beetle fitness in naïve host trees. [ABSTRACT FROM AUTHOR]

Norway spruce (Picea abies L. Karst) is one of the most important forest tree species with significant economic and ecological impact in Europe. For decades, genomic and genetic studies on Norway spruce have been challenging due to the large and repetitive genome (19.6 Gb with more than 70% being repetitive). To accelerate genomic studies, including population genetics, genome‐wide association studies (GWAS) and genomic selection (GS), in Norway spruce and related species, we here report on the design and performance of a 50K single nucleotide polymorphism (SNP) genotyping array for Norway spruce. The array is developed based on whole genome resequencing (WGS), making it the first WGS‐based SNP array in any conifer species so far. After identifying SNPs using genome resequencing data from 29 trees collected in northern Europe, we adopted a two‐step approach to design the array. First, we built a 450K screening array and used this to genotype a population of 480 trees sampled from both natural and breeding populations across the Norway spruce distribution range. These samples were then used to select high‐confidence probes that were put on the final 50K array. The SNPs selected are distributed over 45,552 scaffolds from the P. abies version 1.0 genome assembly and target 19,954 unique gene models with an even coverage of the 12 linkage groups in Norway spruce. We show that the array has a 99.5% probe specificity, >98% Mendelian allelic inheritance concordance, an average sample call rate of 96.30% and an SNP call rate of 98.90% in family trios and haploid tissues. We also observed that 23,797 probes (50%) could be identified with high confidence in three other spruce species (white spruce [Picea glauca], black spruce [P. mariana] and Sitka spruce [P. sitchensis]). The high‐quality genotyping array will be a valuable resource for genetic and genomic studies in Norway spruce as well as in other conifer species of the same genus. [ABSTRACT FROM AUTHOR]

With climate change, the pressure on tree breeding to provide varieties with improved resilience to biotic and abiotic stress is increasing. As such, pest resistance is of high priority but has been neglected in most tree breeding programs, given the complexity of phenotyping for these traits and delays to assess mature trees. In addition, the existing genetic variation of resistance and its relationship with productivity should be better understood for their consideration in multitrait breeding. In this study, we evaluated the prospects for genetic improvement of the levels of acetophenone aglycones (AAs) in white spruce needles, which have been shown to be tightly linked to resistance to spruce budworm. Furthermore, we estimated the accuracy of genomic selection (GS) for these traits, allowing selection at a very early stage to accelerate breeding. A total of 1,516 progeny trees established on five sites and belonging to 136 full‐sib families from a mature breeding population in New Brunswick were measured for height growth and genotyped for 4,148 high‐quality SNPs belonging to as many genes along the white spruce genome. In addition, 598 trees were assessed for levels of AAs piceol and pungenol in needles, and 578 for wood stiffness. GS models were developed with the phenotyped trees and then applied to predict the trait values of unphenotyped trees. AAs were under moderate‐to‐high genetic control (h2: 0.43–0.57) with null or marginally negative genetic correlations with other traits. The prediction accuracy of GS models (GBLUP) for AAs was high (PAAC: 0.63–0.67) and comparable or slightly higher than pedigree‐based (ABLUP) or BayesCπ models. We show that AA traits can be improved and that GS speeds up the selection of improved trees for insect resistance and for growth and wood quality traits. Various selection strategies were tested to optimize multitrait gains. [ABSTRACT FROM AUTHOR]

Understanding the factors influencing the current distribution of genetic diversity across a species range is one of the main questions of evolutionary biology, especially given the increasing threat to biodiversity posed by climate change. Historical demographic processes such as population expansion or bottlenecks and decline are known to exert a predominant influence on past and current levels of genetic diversity, and revealing this demo‐genetic history can have immediate conservation implications. We used a whole‐exome capture sequencing approach to analyze polymorphism across the gene space of red spruce (Picea rubens Sarg.), an endemic and emblematic tree species of eastern North America high‐elevation forests that are facing the combined threat of global warming and increasing human activities. We sampled a total of 340 individuals, including populations from the current core of the range in northeastern USA and southeastern Canada and from the southern portions of its range along the Appalachian Mountains, where populations occur as highly fragmented mountaintop "sky islands." Exome capture baits were designed from the closely relative white spruce (P. glauca Voss) transcriptome, and sequencing successfully captured most regions on or near our target genes, resulting in the generation of a new and expansive genomic resource for studying standing genetic variation in red spruce applicable to its conservation. Our results, based on over 2 million exome‐derived variants, indicate that red spruce is structured into three distinct ancestry groups that occupy different geographic regions of its highly fragmented range. Moreover, these groups show small Ne, with a temporal history of sustained population decline that has been ongoing for thousands (or even hundreds of thousands) of years. These results demonstrate the broad potential of genomic studies for revealing details of the demographic history that can inform management and conservation efforts of nonmodel species with active restoration programs, such as red spruce. [ABSTRACT FROM AUTHOR]

Background: The presupposition of genomic selection (GS) is that predictive accuracies should be based on population-wide linkage disequilibrium (LD). However, in species with large, highly complex genomes the limitation of marker density may preclude the ability to resolve LD accurately enough for GS. Here we investigate such an effect in two conifer species with ~ 20 Gbp genomes, Douglas-fir (Pseudotsuga menziesii Mirb. (Franco)) and Interior spruce (Picea glauca (Moench) Voss x Picea engelmannii Parry ex Engelm.). Random sampling of markers was performed to obtain SNP sets with totals in the range of 200–50,000, this was replicated 10 times. Ridge Regression Best Linear Unbiased Predictor (RR-BLUP) was deployed as the GS method to test these SNP sets, and 10-fold cross-validation was performed on 1,321 Douglas-fir trees, representing 37 full-sib F1 families and on 1,126 Interior spruce trees, representing 25 open-pollinated (half-sib) families. Both trials are located on 3 sites in British Columbia, Canada. Results: As marker number increased, so did GS predictive accuracy for both conifer species. However, a plateau in the gain of accuracy became apparent around 10,000–15,000 markers for both Douglas-fir and Interior spruce. Despite random marker selection, little variation in predictive accuracy was observed across replications. On average, Douglas-fir prediction accuracies were higher than those of Interior spruce, reflecting the difference between full- and half-sib families for Douglas-fir and Interior spruce populations, respectively, as well as their respective effective population size. Conclusions: Although possibly advantageous within an advanced breeding population, reducing marker density cannot be recommended for carrying out GS in conifers. Significant LD between markers and putative causal variants was not detected using 50,000 SNPS, and GS was enabled only through the tracking of relatedness in the populations studied. Dramatically increasing marker density would enable said markers to better track LD with causal variants in these large, genetically diverse genomes; as well as providing a model that could be used across populations, breeding programs, and traits. [ABSTRACT FROM AUTHOR]

The spruce budworm, Choristoneura fumiferana, is presumed to be panmictic across vast regions of North America. We examined the extent of panmixia by genotyping 3,650 single nucleotide polymorphism (SNP) loci in 1975 individuals from 128 collections across the continent. We found three spatially structured subpopulations: Western (Alaska, Yukon), Central (southeastern Yukon to the Manitoba–Ontario border), and Eastern (Manitoba–Ontario border to the Atlantic). Additionally, the most diagnostic genetic differentiation between the Central and Eastern subpopulations was chromosomally restricted to a single block of SNPs that may constitute an island of differentiation within the species. Geographic differentiation in the spruce budworm parallels that of its principal larval host, white spruce (Picea glauca), providing evidence that spruce budworm and spruce trees survived in the Beringian refugium through the Last Glacial Maximum and that at least two isolated spruce budworm populations diverged with spruce/fir south of the ice sheets. Gene flow in the spruce budworm may also be affected by mountains in western North America, habitat isolation in West Virginia, regional adaptations, factors related to dispersal, and proximity of other species in the spruce budworm species complex. The central and eastern geographic regions contain individuals that assign to Eastern and Central subpopulations, respectively, indicating that these barriers are not complete. Our discovery of previously undetected geographic and genomic structure in the spruce budworm suggests that further population modelling of this ecologically important insect should consider regional differentiation, potentially co‐adapted blocks of genes, and gene flow between subpopulations. [ABSTRACT FROM AUTHOR]

Analyses of maximum early tree ring growth within a regional curve standardization (RCS) framework were used to construct ring width chronologies in mm year−1 to estimate historical growth rates in southwest Yukon (Canada), without the necessity of correcting for age related changes. The analysis relied on the use of the 11th–50th year growth rings to standardize chronology comparisons. This age range represents the period of peak radial growth for Picea albertiana and P. glauca (white spruces), which are common trees on upland boreal sites in the study area. Of 1,460 available Picea ring series, 1,029 fit the criteria for inclusion in the study and they represented five commonly occurring RCS curve types. When converted to chronologies, all curve types had year-to-year fluctuations in average ring widths (RW11–50), but nine of the 12 RCS types (four types in three study areas) also had long-term growth rate increases of 15–132% between ad 1850 and 2000, based on regression modeling (R/r = 0.54–0.76, p < 0.001). Associated with these increases were changes through time in RCS curve type and the timing of peak growth. Among studied areas, RW11–50 values on average increased 47% from 1.59 to 2.35 mm year−1 between 1850 and 2000. The greatest increase (71% vs 35–38%) occurred in the coldest study area. RW11–50 values were significantly (p < 0.001) and strongly correlated (r > 0.94, n > 100) with conventional tree ring width index values derived from multiple RCS curves. This indicates that the two chronology construction approaches have very similar temporal patterns despite the maximum early growth analysis technique representing ring widths in mm. [ABSTRACT FROM AUTHOR]

An emerging trend in tree-ring research is use of multiple species for reconstructing paleoclimates, but the possible simultaneous use of boreal-cordilleran species is untested. In this study, ring-width chronologies of sympatric Pinus contorta (lodgepole pine, n = 116 series) and Picea albertiana (western white spruce, n = 348 series) were constructed to assess their temporal (dis)similarities and correlative relationships with meteorological variables. Chronology construction was based on multiple Regional Curve Standardization. Most (97.5%) Pinus chronology values occurred within ±1 SD of their Picea counterparts, but Picea values tended to be slightly less (sign-test, p < 0.001, n = 201). Pinus ring widths were more frequently (16 versus 9) and more strongly correlated (U-test, p < 0.009) with 1942-2013 meteorological variables than Picea. Both species were correlated with moisture variables, but Picea was not correlated with those of temperature. Pinus and Picea ring-width variation was best explained by summer (r = -0.434) and annual (r = -0.426) heat-moisture index values (p < 0.001, n = 72) among tested variables, respectively. Although seldom used, P. contorta appears as suitable as conventionally-used P. albertiana based on chronology similarity, but a greater diversity of significant correlation outcomes suggest Pinus is potentially more useful for boreal-cordilleran paleoclimate reconstruction. [ABSTRACT FROM AUTHOR]

• Introducing spruce in plantations promoted the establishment of bryophytes. • Poplar plantations provided habitat for lichens living on deciduous tree trunks. • Mixed plantations were composed of terrestrial bryophytes and epiphytic lichens. • Vascular plant composition was sensitive to the land-use history. Although monocultures are important for timber production, they are often associated with lower biological diversity than mixtures. Thus, mixed plantations have been suggested as a way to enhance biodiversity because of their inherent compositional diversity. However, the effects of monocultures versus mixtures on understory diversity and composition can vary in different ecosystems. The objective of this study was to assess how monocultures and mixed plantations influence understory vegetation diversity and composition in the boreal forest region of southern Quebec. We sampled plantations established with deciduous Populus trichocarpa Torrey & A. Gray × balsamifera L. and P. maximowiczii Henry × balsamifera L. and coniferous Picea abies (L.) Karst. and Picea glauca (Moench) species planted in monocultures and in mixed plantations on abandoned farmlands and a forest site. We assessed understory vegetation diversity and composition in each canopy type (coniferous, deciduous, mixed) and in each plantation type. We evaluated bryophyte and lichen diversity and composition specifically in tree microhabitats: soil, tree bases, and tree trunks. We found that vascular plant and lichen species richness was similar in all plantation types, while bryophyte species richness was higher in spruce monocultures and in mixed plantations compared to poplar monocultures. Our results also highlight how land-use history influenced vascular plant composition as abandoned farmland sites were composed of more ruderal vascular plants, while the previously forested site was composed of species found in natural forests. In the context of reforestation and plantations, our study suggests mixing spruce with poplars to maximize understory vegetation diversity as the addition of spruce in mixed plantations promoted the establishment of terrestrial bryophytes, while poplars favored the establishment of epiphytic lichens. [ABSTRACT FROM AUTHOR]

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Highlights • Four pines and four spruces in a 2 × 2 factorial of CO 2 and soil moisture. • Overall, pines respond better to eCO 2 and soil moisture stress. • Shade-tolerant spruces perform better in eCO 2 than shade-intolerant spruces. • Greater the mass, greater the response to eCO 2 and soil moisture. • Supports shift toward increased use of pines in NE N. America. Abstract Growth, components of growth, and biomass allocation were quantified for eight species in two commercially important genera, Pinus and Picea , grown in a 2 × 2 factorial of atmospheric CO 2 and soil moisture stress. Four of the pines and three of the spruces are native to eastern North America; a fourth spruce, Norway spruce (NS: P. abies), is native to Europe but is used for reforestation in northeastern North America. Height, basal diameter (BD), and total biomass response of pines were often more than two times greater than that of spruces under elevated CO 2 (eCO 2). A significant species × CO 2 interaction for total biomass was a result of species' differential response to eCO 2 : Pinus rigida had the greatest biomass stimulation (59%), followed by P. resinosa (39%), P. strobus (26%), and P. banksiana (19%). Among spruces, Picea glauca showed the greatest response (30%), and P. mariana the least response under eCO 2 (5%). Overall, soil moisture stress reduced total productivity by 12%. Most pines did have greater growth under moisture stress, and NS and BS grew well. Percent needle mass was lower under eCO 2 , but this was not due directly to eCO 2 but to ontological changes. Controlling for size, pines had 20% greater needle biomass than spruces, while having a negative relationship to total biomass. A comparison of total biomass under eCO 2 in relation to aCO 2 by species showed that overall, the greater the species mass, the greater the mass gain under eCO 2, and the greater the mass loss under drought conditions. In addition, our results for spruces lend strong support to the theory that late-successional species have greater growth response under eCO 2 than early to mid-successional species. A diverse portfolio of tree species for artificial reforestation would help forest management adapt to the many uncertainties over future environments and markets, but our results on responses in spruces and pines to eCO 2 and soil moisture stress support a shift toward increased use of pines in forest management and artificial reforestation. [ABSTRACT FROM AUTHOR]

Fundamental to the success of forest restoration following major disturbances such as mining is development of a functioning soil, including the amount, properties and rates of accumulation of soil organic matter (SOM). SOM enters mineral soil through leaching of dissolved organic matter from the forest floor; macrofaunal processing of above-ground litter and mixing with soil (bioturbation); and direct deposition of root litter and rhizodeposits. Our study focused on how SOM accumulation in reclaimed mineral soils was affected following re-vegetation with three vegetation treatments: deciduous trees (Populus tremuloides Mitchx. and Populus balsamifera L.), spruce trees (Picea glauca (Moench) Voss.) and grasses (Festuca sp., Bromus inermis), compared to the natural boreal forest. Seventeen sites were studied: 4 reclaimed Deciduous, 5 reclaimed Spruce, 4 reclaimed Grass, and 4 Natural forest. Concentrations of soil organic matter in the upper 30 cm of soil were highest at the Deciduous sites and lowest at Natural sites. SOM concentrations in the top 10 cm of soil were elevated relative to 10–30 cm soil at the Deciduous and Grass sites. The elevated SOM in the upper soil at Deciduous and Grass sites was associated with greater macrofauanal activity and higher root biomass at these sites. The higher macrofaunal activity in the surface organic layer and greater amounts of faunal fecal material in the uppermost cm of soil at Deciduous and Grass sites indicate greater enrichment of upper soil by macrofauna at these sites. SOM concentrations were significantly positively related to root abundance at Grass sites (indicating a contribution of roots to SOM), and a similar trend was apparent at the Deciduous sites. The elevated organic matter concentrations in the upper 10 cm of soil at the Deciduous and Grass sites suggest that planting of aspen following reclamation would hasten C sequestration into soil organic matter. [ABSTRACT FROM AUTHOR]

Motivation In the modern genomics era, genome sequence assemblies are routine practice. However, depending on the methodology, resulting drafts may contain considerable base errors. Although utilities exist for genome base polishing, they work best with high read coverage and do not scale well. We developed ntEdit, a Bloom filter-based genome sequence editing utility that scales to large mammalian and conifer genomes. Results We first tested ntEdit and the state-of-the-art assembly improvement tools GATK, Pilon and Racon on controlled Escherichia coli and Caenorhabditis elegans sequence data. Generally, ntEdit performs well at low sequence depths (<20×), fixing the majority (>97%) of base substitutions and indels, and its performance is largely constant with increased coverage. In all experiments conducted using a single CPU, the ntEdit pipeline executed in <14 s and <3 m, on average, on E.coli and C.elegans , respectively. We performed similar benchmarks on a sub-20× coverage human genome sequence dataset, inspecting accuracy and resource usage in editing chromosomes 1 and 21, and whole genome. ntEdit scaled linearly, executing in 30–40 m on those sequences. We show how ntEdit ran in <2 h 20 m to improve upon long and linked read human genome assemblies of NA12878, using high-coverage (54×) Illumina sequence data from the same individual, fixing frame shifts in coding sequences. We also generated 17-fold coverage spruce sequence data from haploid sequence sources (seed megagametophyte), and used it to edit our pseudo haploid assemblies of the 20 Gb interior and white spruce genomes in <4 and <5 h, respectively, making roughly 50M edits at a (substitution+indel) rate of 0.0024. Availability and implementation https://github.com/bcgsc/ntedit Supplementary information Supplementary data are available at Bioinformatics online. [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]

• 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]

We examine the extent to which foliar monoterpenes are trapped in the epicuticular waxes as part of an investigation into their role in natural defense against folivores. We monitored concentrations in white spruce ( Picea glauca (Moench) Voss) previous-year foliage and expanding foliage and their epicuticular waxes over the 2010 (14 trees) and 2011 (25 trees) growing seasons. In 2010, concentrations were low in the spring and increased over the summer; in 2011, they stayed low. The monoterpene profile of individual trees was similar between years and showed a consistent pattern over the growing season; in expanding foliage, δ-3-carene was only present in spring, whereas bornyl acetate increased over the growing season. Individual wax monoterpene profiles correlated with those of foliage, but the total concentration showed a different phenological pattern. Total content remained constant throughout the growing season on previous-year foliage but decreased on expanding foliage. Electron microscopy suggests this is due to changes in stomatal wax plugs and their role in blocking evaporation from the stomata. These findings suggest that insects contacting the leaf surface will receive accurate information from the wax chemical composition about the monoterpene mix but not about overall monoterpene levels. [ABSTRACT FROM AUTHOR]

The relationships between white spruce radial increment and wood properties were investigated in relation to tree and stand attributes using data from mature white spruce stands in the boreal forest of western Canada that experienced a range of shelterwood treatments. The model with the highest predictive ability was radial increment (adj-R² = 67%) and included crown attributes, diameter at breast height (DBH), average height of competitors, and a climate index. Radial growth was positively related to live crown ratio, whereas wood density and modulus of elasticity were negatively correlated to the crown attribute. Tree slenderness had a significant negative effect on wood density and modulus of elasticity, as it reflects the mechanical stability requirement of the tree. The models consistently improved when using annual averages calculated over longer periods of time. However, when the annual averages were calculated using time periods of 5-10 and 10-20 years prior to sampling, the predictive ability of the models decreased, which indicated that the current tree and stand conditions were the best predictors of growth and wood properties up to five years prior to sampling. This study suggests that crown length equal to 2/3 of the tree height might represent an optimal balance between radial growth and wood quality. [ABSTRACT FROM AUTHOR]

Intimate mixtures of trembling aspen (Populus tremuloides Michx.) and white spruce (Picea glauca (Moench) Voss) are a key feature of western Canadian boreal forests. These mixtures have the potential to produce high yields of merchantable fibre and provide numerous ecological services. Achievement of this potential has been difficult, and often expensive, to realize as a regeneration goal in managed forests. We report 21 to 25 year results of managed mixtures on two study sites where the white spruce was planted, and the density of aspen natural regeneration manipulated within five years of the stand initiation disturbance. On both sites, white spruce mortality did not increase with increasing aspen density. While height and diameter growth of white spruce declined with increasing aspen density, the effect was not entirely consistent across the two sites. Abrasion from aspen branches was the most common source of damage to spruce crowns. Mixed stands had greater merchantable volume production than pure spruce stands based on model projections. Application of aspen harvest at year 60, while protecting the spruce component for a second harvest entry at year 90, was projected to optimize combined yield for the mixedwood stands. [ABSTRACT FROM AUTHOR]

We assessed whether commercially thinning (CT) spruce ( Picea spp.) plantations (40% basal area removal) can cause structural changes in fine woody debris (FWD) and understory vegetation to improve habitat quality for small rodents in northwestern New Brunswick, Canada. We created three contrasting environments (FWD rich - understory vegetation rich, FWD poor - understory vegetation rich, and FWD poor - understory vegetation poor) by establishing CT with debris retention (merchantable trunk removal), CT with all debris removed (full tree removal), and non-CT (plantation without CT) in six midrotation white spruce ( Picea glauca (Moench) Voss) plantations. We live-trapped small mammals in each plantation during spring and summer of 2011 and 2012 and estimated animal density and survival with capture-recapture models. Southern red-backed vole ( Myodes gapperi (Vigors, 1830)) density was two to three times greater in CT with debris retention than in either non-CT or CT with debris removal, and their survival rate was twice that in CT with debris removal. Woodland jumping mouse ( Napaeozapus insignis (Miller, 1891)) density was two to five times greater in non-CT than in either CT treatment. Deer mice ( Peromyscus maniculatus (Wagner, 1845)) did not show any treatment effect. Productivity and demographics were not affected by CT treatment for any species. We found evidence that midrotation spruce plantations are used differently by small mammal species based on stand condition and recommend that managers maintain plantations in CT and non-CT states. [ABSTRACT FROM AUTHOR]

The contribution of hyphae to water transport in ectomycorrhizal ( ECM) white spruce ( Picea glauca) seedlings was examined by altering expression of a major water-transporting aquaporin in Laccaria bicolor., Picea glauca was inoculated with wild-type ( WT), mock transgenic or L. bicolor aquaporin -overexpressing ( OE) strains and exposed to root temperatures ranging from 5 to 20°C to examine the root water transport properties, physiological responses and plasma membrane intrinsic protein ( PIP) expression in colonized plants., Mycorrhization increased shoot water potential, transpiration, net photosynthetic rates, root hydraulic conductivity and root cortical cell hydraulic conductivity in seedlings. At 20°C, OE plants had higher root hydraulic conductivity compared with WT plants and the increases were accompanied by higher expression of P. glauca PIP GQ03401_ M18.1 in roots. In contrast to WT L. bicolor, the effects of OE fungi on root and root cortical cell hydraulic conductivities were abolished at 10 and 5°C in the absence of major changes in the examined transcript levels of P. glauca root PIPs., The results provide evidence for the importance of fungal aquaporins in root water transport of mycorrhizal plants. They also demonstrate links between hyphal water transport, root aquaporin expression and root water transport in ECM plants. [ABSTRACT FROM AUTHOR]

Background Genomic selection (GS) may improve selection response over conventional pedigree-based selection if markers capture more detailed information than pedigrees in recently domesticated tree species and/or make it more cost effective. Genomic prediction accuracies using 1748 trees and 6932 SNPs representative of as many distinct gene loci were determined for growth and wood traits in white spruce, within and between environments and breeding groups (BG), each with an effective size of Ne ≈ 20. Marker subsets were also tested. Results Model fits and/or cross-validation (CV) prediction accuracies for ridge regression (RR) and the least absolute shrinkage and selection operator models approached those of pedigreebased models. With strong relatedness between CV sets, prediction accuracies for RR within environment and BG were high for wood (r = 0.71-0.79) and moderately high for growth (r = 0.52-0.69) traits, in line with trends in heritabilities. For both classes of traits, these accuracies achieved between 83% and 92% of those obtained with phenotypes and pedigree information. Prediction into untested environments remained moderately high for wood (r ⩾ 0.61) but dropped significantly for growth (r ⩾ 0.24) traits, emphasizing the need to phenotype in all test environments and model genotype-by-environment interactions for growth traits. Removing relatedness between CV sets sharply decreased prediction accuracies for all traits and subpopulations, falling near zero between BGs with no known shared ancestry. For marker subsets, similar patterns were observed but with lower prediction accuracies. Conclusions Given the need for high relatedness between CV sets to obtain good prediction accuracies, we recommend to build GS models for prediction within the same breeding population only. Breeding groups could be merged to build genomic prediction models as long as the total effective population size does not exceed 50 individuals in order to obtain high prediction accuracy such as that obtained in the present study. A number of markers limited to a few hundred would not negatively impact prediction accuracies, but these could decrease more rapidly over generations. The most promising short-term approach for genomic selection would likely be the selection of superior individuals within large full-sib families vegetatively propagated to implement multiclonal forestry. [ABSTRACT FROM AUTHOR]

Background and Aims MADS-box genes comprise a gene family coding for transcription factors. This gene family expanded greatly during land plant evolution such that the number of MADS-box genes ranges from one or two in green algae to around 100 in angiosperms. Given the crucial functions of MADS-box genes for nearly all aspects of plant development, the expansion of this gene family probably contributed to the increasing complexity of plants. However, the expansion of MADS-box genes during one important step of land plant evolution, namely the origin of seed plants, remains poorly understood due to the previous lack of whole-genome data for gymnosperms. Methods The newly available genome sequences of Picea abies, Picea glauca and Pinus taeda were used to identify the complete set of MADS-box genes in these conifers. In addition, MADS-box genes were identified in the growing number of transcriptomes available for gymnosperms. With these datasets, phylogenies were constructed to determine the ancestral set of MADS-box genes of seed plants and to infer the ancestral functions of these genes. Key Results Type I MADS-box genes are under-represented in gymnosperms and only a minimum of two Type I MADS-box genes have been present in the most recent common ancestor (MRCA) of seed plants. In contrast, a large number of Type II MADS-box genes were found in gymnosperms. The MRCA of extant seed plants probably possessed at least 11–14 Type II MADS-box genes. In gymnosperms two duplications of Type II MADS-box genes were found, such that the MRCA of extant gymnosperms had at least 14–16 Type II MADS-box genes. Conclusions The implied ancestral set of MADS-box genes for seed plants shows simplicity for Type I MADS-box genes and remarkable complexity for Type II MADS-box genes in terms of phylogeny and putative functions. The analysis of transcriptome data reveals that gymnosperm MADS-box genes are expressed in a great variety of tissues, indicating diverse roles of MADS-box genes for the development of gymnosperms. This study is the first that provides a comprehensive overview of MADS-box genes in conifers and thus will provide a framework for future work on MADS-box genes in seed plants. [ABSTRACT FROM AUTHOR]

Leaf life span is an important plant trait associated with interspecific variation in leaf, organismal, and ecosystem processes. We hypothesized that intraspecific variation in gymnosperm needle traits with latitude reflects both selection and acclimation for traits adaptive to the associated temperature and moisture gradient. This hypothesis was supported, because across 127 sites along a 2,160-km gradient in North America individuals of Picea glauca Picea mariana, Pinus banksiana, and Abies balsamea had longer needle life span and lower tissue nitrogen concentration with decreasing mean annual temperature. Similar patterns were noted for Pinus syivestris across a north-south gradient in Europe. These differences highlight needle longevity as an adaptive feature important to ecological success of boreal conifers across broad climatic ranges. Additionally, differences in leaf life span directly affect annual foliage turnover rate, which along with needle physiology partially regulates carbon cycling through effects on gross primary production and net canopy carbon export. However, most, if not all, global land surface models parameterize needle longevity of boreal evergreen forests as if it were a constant. We incorporated temperature-dependent needle longevity and %nitrogen, and bio-mass allocation, into a land surface model. Community Atmosphere Biosphere Land Exchange, to assess their impacts on carbon cycling processes. Incorporating realistic parameterization of these variables improved predictions of canopy leaf area index and gross primary production compared with observations from flux sites. Finally, increasingly low foliage turnover and biomass fraction toward the cold far north indicate that a surprisingly small fraction of new biomass is allocated to foliage under such conditions. [ABSTRACT FROM AUTHOR]

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Areas of geographic overlap between potentially hybridizing species provide the opportunity to study interspecific gene flow and reproductive barriers. Here we identified hybrids between Picea engelmannii and P. glauca by their genetic composition at 17 microsatellite markers, and determined the broad-scale geographic distribution of hybrids in the central Rocky Mountains of North America, a geographic region where hybrids and isolation between species had not previously been studied. Parameter estimates from admixture models revealed considerable variation in ancestry within and among collection sites, suggesting that within this area of geographic overlap, the interaction of the two species varies extensively. The results document a previously unrecognized patchy distribution of hybrids between P. engelmannii and P. glauca, including locations where hybrids were not known or expected to exist. Further, the ancestry of many hybrids was consistent with multiple generations of hybridization, with probable directional backcrossing to P. engelmannii, suggesting a relatively porous species boundary. The identification and characterization of hybridization between these spruce in this region raises the question of what factors maintain barriers to gene flow in these long-lived forest trees. The current research lays the groundwork for future study of the ecological and evolutionary contexts of their hybridization, as well as of differential introgression and permeability of species boundaries. [ABSTRACT FROM AUTHOR]

High-density SNP genotyping arrays can be designed for any species given sufficient sequence information of high quality. Two high-density SNP arrays relying on the Infinium iSelect technology (Illumina) were designed for use in the conifer white spruce ( Picea glauca). One array contained 7338 segregating SNPs representative of 2814 genes of various molecular functional classes for main uses in genetic association and population genetics studies. The other one contained 9559 segregating SNPs representative of 9543 genes for main uses in population genetics, linkage mapping of the genome and genomic prediction. The SNPs assayed were discovered from various sources of gene resequencing data. SNPs predicted from high-quality sequences derived from genomic DNA reached a genotyping success rate of 64.7%. Nonsingleton in silico SNPs (i.e. a sequence polymorphism present in at least two reads) predicted from expressed sequenced tags obtained with the Roche 454 technology and Illumina GAII analyser resulted in a similar genotyping success rate of 71.6% when the deepest alignment was used and the most favourable SNP probe per gene was selected. A variable proportion of these SNPs was shared by other nordic and subtropical spruce species from North America and Europe. The number of shared SNPs was inversely proportional to phylogenetic divergence and standing genetic variation in the recipient species, but positively related to allele frequency in P. glauca natural populations. These validated SNP resources should open up new avenues for population genetics and comparative genetic mapping at a genomic scale in spruce species. [ABSTRACT FROM AUTHOR]

Mixedwood forests of white spruce ( Picea glauca (Moench) Voss) and trembling aspen ( Populus tremuloides Michx.) may possess ecological advantages over monospecific white spruce stands. Belowground competition may be reduced through vertically stratified roots; facilitation of growth may occur in upper soil layers through nutrient-rich trembling aspen litterfall. These effects may incentivize white spruce to preferentially exploit upper soil layers in mixedwood stands, resulting in wider root systems. This research contrasted white spruce fine root (diameter <2 mm) distributions in organic layers of white spruce and mixedwood stands. Research occurred at the Fallingsnow Ecosystem Project site in northwestern Ontario. Eighteen plots represented mixedwood and pure white spruce stands. Trees were mapped; foliage and root samples were collected. Roots were separated by species and scanned to determine length. Simple sequence repeat DNA profiles were determined for all white spruce trees and for 45 white spruce root fragments per plot. Root and tree DNA profiles were matched; corresponding distances were calculated. Most (80%) root fragments were within 3.2 m of tree stems. Root prevalence decreased rapidly with distance. Organic layer pH was significantly less acidic in mixedwood plots, but only in one block. A subtle significant trend towards wider root distributions occurred in mixedwood stands. [ABSTRACT FROM AUTHOR]

Predator satiation resulting from inter-annual reproductive synchrony has been widely documented in masting plants, but how reproductive synchrony within a year influences seed escape is poorly understood. We evaluated whether the intra-annual reproductive synchrony of individual white spruce trees (Picea glauca) increased seed escape from their primary predispersal seed predator, North American red squirrels (Tamiasciurus hudsonicus). Trees with cones that matured synchronously relative to those of other trees within red squirrel territories were significantly more likely to escape squirrel predation in years with both low and superabundant levels of cone production, generating a significantly positive linear selection differential for increasing intra-annual reproductive synchrony. Thus, this masting plant escapes seed predation in numbers through inter-annual synchrony in seed production and in time through intra-annual synchrony of seed availability. [ABSTRACT FROM AUTHOR]

Copyright of Forestry Chronicle is the property of Canadian Institute of Forestry and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)