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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]

Picea × darwyniana Björk & Goward was recently proposed to replace Picea albertiana S. Brown as the representative taxon for the hybrid offspring of Picea glauca (Moench) Voss and Picea engelmannii Parry ex Engelm., but should be considered invalid for three reasons. First, the proponents failed to document in any way that the P. albertiana isotypes were Picea glauca × mariana Little & Pauley hybrids, as they claimed. Secondly, the P. albertiana isotypes lacked pubescent twigs, purple markings on their seed cones, and denticulate cone-scale margins, which are characteristics of P. glauca × mariana. This indicates a misinterpretation of the P. albertiana parentage. Lastly, P. × darwyniana appears to be an example of P. albertiana ssp. ogilviei Strong & Hills. As the latter is an earlier and validly published name, P. × darwyniana is an illegitimate replacement for plants within the P. glauca × engelmannii (=P.a. ssp. albertiana) and P. engelmannii × glauca (=P.a. ssp. ogilviei) circumscriptions based on naming priority. [ABSTRACT FROM AUTHOR]

Rapid establishment of closed canopy cover is a key goal in land reclamation. While re-establishment of understory vegetation is likely to positively contribute to this goal, native herbaceous species are not typically planted on reclaimed sites though it is an alternative to reliance on natural regeneration. The objective of the present study was to test the principle of hitchhiker production of a tree species with native herbaceous species on recently reclaimed industrial sites with the aim of (1) understanding growth tradeoffs when mixing these species in the same container and (2) if hitchhiking had negative growth and survival outcomes for white spruce (Picea glauca) seedlings compared with singly grown white spruce seedlings. White spruce seeds were sown in nursery containers of contrasting cavity sizes, followed by sowing a forb species (Eurybia conspicua or Chamerion angustifolium) at different time intervals or grown singly. Larger cavities and earlier sow dates were associated with higher persistence and growth of the forbs, while later sow dates were associated with larger white spruce seedlings. We observed no evidence to suggest that hitchhiked seedlings were at a competitive disadvantage in terms of growth or survival compared with singly grown seedlings that were initially similar in size. [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]

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]

Herbivores affect plants via direct consumption, but also indirectly via changes in the vegetation. These indirect effects may only manifest after decades of exposure and, as a result, their impact is rarely accounted for. To better understand the direct and indirect impacts of a large herbivore, moose (Alces alces), on forests, we measured tree growth in areas that were both subjected to and excluded from herbivory for over 80 years. Growth data were gathered from five tree species, ranging from low to high palatability. We found that at small sizes, Betula papyrifera, a preferred species, benefited from herbivore exclusion. However, larger individuals grew more when exposed to herbivory, a response we attribute to lower competition in heavily browsed conditions. Populus tremuloides, a highly preferred tree, did not show any differences between levels of herbivory. Abies balsamea, a preferred winter browsed, was only marginally affected by browsing at smaller sizes. The two non-preferred species, Picea glauca and Picea mariana, did not show differential growth between herbivory levels. We conclude that herbivores can impact forests through both direct and indirect effects, that these effects are size specific, and that effects vary among species in ways not always predicted by consumption patterns. [ABSTRACT FROM AUTHOR]

Assisted gene flow according to expected climate gradients is considered as a forest management strategy to mitigate impacts of environmental change on forest growth. However, the effects of seed translocation on wood properties and lumber quality remain unknown. This study evaluated the effect of provenance origin on lumber production and quality at rotation age in two white spruce provenance trials established in contrasting environments in eastern Canada. Based on 108 sample trees, which resulted in 943 pieces of lumber, average volume production per tree at the southernmost site was twice that of the production at the northern site. Provenance had a significant influence on growth and lumber strength in the first sawlog but had no effect on lumber stiffness and wood density. Although visual grade yields of No. 2 and better were high in both trials (over 86%), the machine stress rated (MSR) grade potential and percentage of lumber that met the bending stiffness design values of the visual grades were generally low (12%–26%). Hence, plantation-grown lumber should preferably be machine stress rated to ensure its fitness for structural applications in buildings. Management strategies aiming to efficiently sequester carbon should primarily maximize volume productivity in northern sites, as moving seed sources north still reduces provenance productivity, while breeding programs should aim to prevent decrease in lumber stiffness due to augmented productivity and shortened rotation cycles. [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]

If the modern climate trend continues for an additional 20–40 years, forests along the southern border of Canadian taiga may shift into new ranges of annual temperature and precipitation that are similar to the modern climate of central New York State, United States. Here we investigate the possible future response of white spruce (Picea glauca (Moench) Voss) to a warmer climate by studying trees planted 90 years ago near the southern limit of their climate tolerance in central New York, 300 km south of the boreal forest where this species is prevalent. We employed high-frequency recording dendrometers to determine radial growth phenology of six mature white spruce trees during 2013–2017. The results demonstrate significant reductions in the length of radial growth periods inversely proportional to the number of hot days with air temperature exceeding 30 °C. During years with very hot summers, the start of radial growth began about 3 days earlier than the 2013–2017 average. However, in those same years, the end of radial growth was also about 17 days earlier resulting in a shorter (70 days versus 100 days) radial growth season. Abundant (350–500 mm) summer precipitation, which resulted in soil moisture values of 20%–30%, allowed us to dismiss drought as a factor. Instead, a likely cause of reduced radial growth was mean temperature that exceeded optimal growth ranges, exacerbated by temperature extremes in ranges that lead to photoinhibition. The latter case was evident in the two warmest growing seasons (2013 and 2017), marked by extended periods of high daytime temperature (>30 °C) followed by early cessation of radial growth independent of seasonal precipitation. [ABSTRACT FROM AUTHOR]

Tree establishment to restore degraded boreal post-mining lands is challenged by low soil productivity, a harsh microclimate, and potentially high contaminant levels. The use of mixed vegetation can facilitate the microclimate but increase competition for soil resources. A statistical accounting of plant–plant interactions and adaptation to multispecies conditions is hard to achieve in field experiments; trials under controlled conditions can distinguish effects of planting density and species interactions in the early stages of plant establishment. A greenhouse trial was established in containers ("mesocosms") with waste rock or fine tailings from gold mines. Pregerminated (1-week-old) seedlings (Alnus viridis subsp. crispa, Picea glauca, Populus tremuloides, Salix arbusculoides) were planted using a Nelder density gradient design, modified for species combinations. A relative competition effect was estimated as a competitiveness index for each species combination, calculated as a ratio of α coefficients in the Holliday growth equation. The specific leaf area (SLA) was measured to indicate plant water stress adaptation. All species grew better in monoculture on fine tailings, while only P. tremuloides grew better in all mixtures on waste rock. Although net positive effects of density on SLA increment during early growth suggested microclimate improvement on fine tailings, no mixture provided advantages for both species in paired combinations. [ABSTRACT FROM AUTHOR]

Populations of many tree species exhibit synchronous and highly temporally variable seed crops across years. This is called mast seeding, and there are two predominant hypotheses for this pattern of reproduction: pollination efficiency and seed-predator satiation. Mast seeding studies typically involve records of population-level reproduction, with less information on the characteristics of reproductive structures. Here, we use data across 6 years (2012–2017), spanning a range of population-level cone conditions, to characterize (i) white spruce (Picea glauca (Moench) Voss) cone lengths and seeds per cone, and (ii) levels of seed predation. We quantified population-level cone production and collected 1399 cones from a total of 38 trees in the Huron Mountains, Michigan, USA. Linear mixed models showed that mean and minimum cone lengths varied significantly across years; both being longest during the greatest cone production year. Larger cones had more seeds and the slopes of the relationships as well as the intercepts varied significantly across years. Generalized linear mixed models and Akaike's information criterion model selection showed that cones with insect predation damage was greatest when population-level reproduction was the lowest, with a mean proportion of cones damaged 0.82 in that year. Our findings show that white spruce cone characteristics and losses to insect seed predation vary temporally, and follow expectations based on mast seeding hypotheses. [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]

Long-term studies of vegetation succession can inform restoration of degraded forests. We examined resilience of a boreal mixedwood vegetation community, asking whether treatments employed to restore wood production in a degraded ecosystem could also restore diversity and composition of vegetation communities. The Inga Lake trial, established in 1987 in northeastern British Columbia, used mechanical, fire, and chemical and manual treatments, encompassing a gradient of restoration effort, and tree planting to restore a shrubland to white spruce (Picea glauca (Moench) Voss) forest. We monitored vascular plant, bryophyte, and macrolichen composition five times over 31 years on five to seven treatments replicated five times. We used mixed-effects models and nonmetric multidimensional scaling to compare diversity and composition among treatments and with mature reference forests. Low- to high-effort restoration created a gradient from broadleaf- to spruce-dominated overstories. Diversity increased with restoration effort. Four of 253 taxa occurred in mature forests only. There was no evidence that lower versus higher effort treatments followed divergent successional pathways toward broadleaved versus spruce reference communities. Our results suggest that these mixedwood vegetation communities lie within a broad domain of successional attraction that confers high ecological resilience to disturbance. Gap cuttings to stimulate understory re-initiation and provide woody debris are recommended to complete the restoration. [ABSTRACT FROM AUTHOR]

Understanding the magnitude and cause of variation in tree growth and forest productivity is central to sustainable forest management. Measurements of annual growth rings allow assessments of individual-tree, tree population, and forest ecosystem vulnerabilities to drought stress or other changing forest disturbance regimes (insects, diseases, fire), which can be used to identify areas at greatest risk of forest losses. Given a heightened demand for tree-ring data, we consolidated and synthesized tree-ring studies and datasets gathered over the past 30 years in Canada by scientists with the Canadian Forest Service and research partners. We incorporated these datasets into a data repository that currently contains tree-ring measurements from 40 206 tree samples from 4594 sites and 62 tree species from all Canadian provinces and territories. Through our synthesis, we demonstrate the value of such large ensembles of tree-ring data for identifying patterns in tree growth over large spatial scales by mapping pan-Canadian drought sensitivity. Overall, we found high coherence in the samples analyzed; low coherence was generally limited to data-poor regions and species. Drought sensitivity was widespread across species and regions: 34% of sampled trees displayed a significant positive relationship between annual growth increment and summer soil moisture index. Dependence upon water availability in species Picea mariana (Mill.) BSP, Pinus banksiana Lamb., Pinus contorta Douglas ex Loudon, and Pseudotsuga menziesii (Mirb.) Franco was more strongly expressed in the warmest regions of the species' range; for species Picea glauca (Moench) Voss and Populus tremuloides Michx., drought sensitivity was stronger in the driest regions. This unprecedented consolidation and synthesis of tree-ring data will enable new research initiatives (e.g., meta-analyses) aimed at improved understanding of the drivers, patterns, and implications of changes in tree growth, as well as facilitating new research collaborations in earth and environmental sciences. Among other things, there is a need for expanding the spatial distribution of sites across Canada's northern regions, increasing the number of samples collected from older stands and angiosperm species, and integrating datasets from studies that evaluate the effects of silvicultural experiments, including provenance and progeny trials, on tree growth. [ABSTRACT FROM AUTHOR]

Increasing the production of wood fibre from conifer species such as white spruce (Picea glauca (Moench) Voss) is one of many challenges in the management of boreal mixedwood forests. The effects of various competition measures on relative growth and relative growth rate variables were calculated for individual white spruce subject trees. Correlation analysis was used to explore relationships with competitor structural features, including the ratio of competitor basal area to subject tree basal area (CBAS), the ratio of competitor height to subject tree height (AHCS), and the proportion of softwood (FSW). Regression analysis was used to explore relationships with three distance-dependent competition indices. The ratio of subject tree height to diameter at breast height (DBH) (HDR), crown ratio (CR), and crown relative increment rate (CRIR) were significantly correlated with CBAS and AHCS. HDR, CR, CRIR, and DBH relative growth rate were all statistically significantly related to the competition indices. Results indicated that (i) relative growth and relative growth rate measures successfully captured a range of competition, (ii) crowns of trees with larger diameters used their horizontal growing space more efficiently to produce stemwood, and (iii) the proportion of softwood contributing to competition did not appear to influence subject tree production efficiency. Growth efficiency variables have the potential to improve our understanding of boreal mixedwood dynamics. [ABSTRACT FROM AUTHOR]

Defoliation level and site type are thought to influence tree response during spruce budworm (Choristoneura fumiferana (Clemens)) outbreaks. We determined the effects of four manual defoliation treatments (0%, 50%, 100%, and 100% + bud removal of current foliage) for 3 years on foliage production of balsam fir (Abies balsamea (L.) Mill.), black spruce (Picea mariana (Mill.) Britton, Sterns & Poggenb.), and white spruce (Picea glauca (Moench) Voss) trees on four site-quality classes. After 3 years of defoliation and 2 years of recovery, foliage biomass was reduced by 34%–98%. During defoliation, the number of shoots generally increased and shoot length of spruce generally decreased, especially on rich sites. During recovery, the number of shoots increased substantially, shoot length decreased, and bud destruction reduced the number of shoots by about 50% compared with that of trees that received the 100% defoliation treatment. Defoliation did not substantially affect needle length. Trees on rich sites had two- to fourfold greater foliage production than trees on poor sites. Effects of site and defoliation differed among species, but site quality, especially nutrition, played an important role in production of shoots and needles and the tree's ability to withstand defoliation. Black spruce had more limited ability to recover foliage biomass, only producing more shoots, whereas balsam fir and white spruce had stronger ability to recover needle and shoot length, respectively. [ABSTRACT FROM AUTHOR]

The onset of lifetime reproduction is central to processes of plant adaptation and population changes. The relative contributions of size and age to the onset of reproduction are important to predicting that key event in individuals. To estimate respective size and age effects on first reproduction in a conifer, I observed white spruce (Picea glauca (Moench) Voss) trees in forest stands in Alaska, United States, over ranges of cumulative growth rates. Age was estimated by counting annular growth rings, size was indicated by tree height, and reproductive status was indicated by seed cones. Multiple logistic regression showed the relative contributions of tree height and age, adjusted for each other, to statistical predictions of reproductive maturity. Trees selected for having little competition for light first produced cones at heights between 1 and 5 m with little additional effect of age. Tree height contributed approximately 50% to regression predictions of reproductive maturity. In contrast, tree age contributed ≤1.5% and bore little relation to reproductive maturity other than as a component of growth rate. This is among the first reports to partial the effect sizes of tree size and age on the commencement of lifetime reproduction in a conifer. [ABSTRACT FROM AUTHOR]

Understanding interactions between competition and climate in relation to their effects on individual tree growth is crucial to the development of climate-sensitive growth models required for modelling boreal forest succession in a changing climate. We used data from permanent growth and yield sample plots in western Canada and Alaska to investigate the impact of competition within a regional gradient of climatic conditions for lodgepole pine (Pinus contorta Douglas ex Loudon), jack pine (Pinus banksiana Lamb.), trembling aspen (Populus tremuloides Michx.), balsam poplar (Populus balsamifera L.), white spruce (Picea glauca (Moench) Voss), and black spruce (Picea mariana (Mill.) Britton, Sterns & Poggenb.). We characterized the effects of competition (basal area of spruce–fir, deciduous, and pine trees larger than the focal tree) and climate (mean annual temperature and precipitation) and their interactions on basal area growth of individual trees using linear mixed-effects models. Our results indicated that intraspecific competition had stronger effects on growth than interspecific competition and climate. Moreover, significant interactions between intraspecific competition and climate suggest that an increase in intraspecific competition will lead to a reduction in tree growth for warmer regions (lodgepole pine, trembling aspen, balsam poplar, and white spruce) and wetter regions (jack pine). The manner in which interspecific competition altered tree growth responses to climate was variable, depending on tree species and competition type. These results indicate that the relationships between growth and climate may differ according to the degree of competition and the structure of the stand. [ABSTRACT FROM AUTHOR]

Climate-sensitive height–age models were developed for top height trees of trembling aspen (Populus tremuloides Michx.), jack pine (Pinus banksiana Lamb.), and white spruce (Picea glauca (Moench) Voss) in natural and reclaimed oil sands stands. We used stem analysis data collected from the Athabasca oil sands region in northern Alberta, Canada, and climate data generated by the ClimateWNA model. Height–age trajectories differed between top height trees in natural and reclaimed stands for jack pine and white spruce, but not for trembling aspen. At a given age, white spruce top height trees were taller and jack pine top height trees were shorter in reclaimed stands than those in natural stands, suggesting that it is easier to achieve similar forest productivity for oil sands sites reclaimed with white spruce stands than for sites reclaimed with jack pine stands. The principal climate variables were growing season (May to September) precipitation averaged over the previous 10 years for trembling aspen and jack pine and summer (June to August) precipitation averaged over the previous 10 years for white spruce. These variables had positive effects on the height–age trajectories. [ABSTRACT FROM AUTHOR]

Regionally fitted allometric equations for individual trees and root-to-shoot ratio values are normally used to estimate local aboveground and belowground forest biomass, respectively. However, uncertainties arising from such applications are poorly understood. We developed equations for both aboveground and belowground biomass using destructive sampling for three dominant upland boreal tree species in northwestern Alberta, Canada. Compared with our equations, the diameter-based national equations derived for use across Canada underestimated aboveground biomass for Picea glauca (Moench) Voss but gave reasonable estimates for Populusbalsamifera L. and Populus tremuloides Michx. The national equations based on both tree diameter and height overestimated aboveground biomass for the Populus species but underestimated it for Picea glauca in our study area. The approach of root-to-shoot ratio proposed by the Intergovernmental Panel on Climate Change (IPCC) overestimated belowground biomass by 16%–41%, depending on forest cover type, in comparison with our values estimated directly on site, with the greatest bias in deciduous-dominated stands. When the general allometric equations for aboveground biomass and the root-to-shoot ratio for belowground biomass were combined to estimate stand biomass, overestimation could be as high as 18% in our study area. The results of our study support the development of improved regional allometric equations for more accurate local-scale estimations. Incorporating intraspecific variation of important traits such as tree taper may be especially helpful. [ABSTRACT FROM AUTHOR]

White spruce (Picea glauca (Moench) Voss) plantations have historically been an important source of high-quality forest products in the Great Lakes Region of North America. Thinning in spruce plantations is a common silvicultural practice for reducing competition and promoting resiliency to forest health threats such as eastern spruce budworm (Choristoneura fumiferana Clemens), a native forest pest of eastern North America. Spruce budworm larvae feed on the foliage of trees, which reduces growth and potentially causes mortality during an outbreak. There has been continual spruce budworm defoliation in northern Minnesota, U.S.A., since the mid-1950s, with higher levels of defoliation in the late 1990s. This research modeled the diameter growth response of white spruce 18 years after initial thinning in stands that presently range between 44 and 64 years old. Some stands received a second thinning in recent years. We used generalized nonlinear least squares and nonlinear mixed-effects models to estimate annual diameter growth using common tree and stand metrics. Growth model performance was improved by including thinning and frequency of spruce budworm defoliation as modifiers of diameter growth. Results of this study highlight how thinning in combination with insect disturbance affect diameter growth in white spruce plantations of northern Minnesota. [ABSTRACT FROM AUTHOR]

The increased pressure on timber supply due to a reduced forest land base urges the development of new approaches to fully capture the value of forest products. This paper investigates the effects of knowing the position of knots on lumber volume, value, and grade recoveries in curve sawing of 31 white spruce (Picea glauca (Moench) Voss) and 22 jack pine (Pinus banksiana Lamb.) trees. Internal knot position was evidenced by X-ray computed tomography (CT) imaging, followed by the application of a knot-detection algorithm allowing log reconstruction for use as input in the Optitek sawing simulation software. Comparisons of the three levels of sawing optimization (sweep up, shape optimized, and knot optimized) revealed that considering internal knots before log sawing (e.g., knot optimized) generated 23% more lumber value for jack pine and 15% more for white spruce compared with the traditional sweep-up sawing strategy. In terms of lumber quality, the knot-optimized strategy produced 38% more pieces of grade No. 2 and better in jack pine and 15% more such pieces in white spruce compared with the sweep-up strategy. These results indicate a great potential to increase manufacturing efficiency and profitability by implementing the CT scanning technology, which should aid in developing a strong bioeconomy based on an optimized use of wood. [ABSTRACT FROM AUTHOR]

This study was conducted to quantify growth responses of three major commercial conifer species (lodgepole pine (Pinus contorta Douglas ex Loudon var. latifolia Engelm. ex S. Watson), interior Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.) Franco), and spruce (white spruce (Picea glauca (Moench) Voss) and hybrid spruce (Picea engelmannii Parry ex. Engelm. × Picea glauca (Moench) Voss × Picea sitchensis (Bong.) Carrière))) to various fertilizer blends in interior British Columbia, Canada. Over 25 years, growth-response data were repeatedly collected across 46 installations. The fertilizer blends were classified into three groups: nitrogen only; nitrogen and sulfur combined; and nitrogen, sulfur, and boron combined. The growth responses for stand volume, basal area, and top height were calculated through absolute and relative growth rate ratios relative to a controlled group. Fertilizer blend, inverse years since fertilization, site index, stand density at fertilization, and their interactions with the fertilizer blend were used as explanatory variables. The magnitude and significance of volume and basal area growth responses to fertilization differed by species, fertilizer-blend groups, and stand-condition variables (i.e., site index and stand density). In contrast, the response in top height growth did not differ among fertilization blends, with the exception of the nitrogen and sulfur fertilizer subgroup for lodgepole pine. The models developed in this study will be incorporated into the current growth and yield fertilization module (i.e., Table Interpolation Program for Stand Yields (TIPSY)), thereby supporting guidance of fertilization applications in interior forests in British Columbia. [ABSTRACT FROM AUTHOR]

Mid-latitude forests commonly show increased productivity and ultimately produce larger trees on shaded, northern aspects compared with those on sunny, southern aspects. Little research has been conducted on this phenomenon at higher latitudes where solar radiation is less available. We examined patterns of canopy tree size in a set of 142 naturally regenerated, late-seral conifer stands in northwestern British Columbia, Canada, at latitudes of 54°N–56°N. The height, diameter, and total stem volume of three shade-tolerant conifers (western hemlock (Tsuga heterophylla (Raf.) Sarg.), subalpine fir (Abies lasiocarpa (Hook.) Nutt.), and hybrid spruce (Picea glauca × engelmannii (Moench) Voss)) and shade-intolerant lodgepole pine (Pinus contorta var. latifolia Engelm. ex S. Watson) were analyzed. In contrast to most previous studies, mean canopy tree height, diameter, and volume were greatest on southern aspects and lowest on northern aspects for the subset of sites with pronounced slopes (≥30%). There was some variation in patterns among species, with western hemlock responding most strongly to topography, and species-specific effects of topography on height–diameter allometry. We suggest that decreased levels of radiation at northern latitudes and increased light and warmer temperatures on steeper, southern aspects promote growth. In contrast, colder, shaded, northern aspects with increased snow accumulation, delayed snowmelt, and decreased nitrogen mineralization inhibit growth. Our results, together with published data, suggest that a latitudinal shift from higher forest productivity on northern aspects to higher productivity on southern aspects typically occurs between 40°N and 50°N. [ABSTRACT FROM AUTHOR]

Additions of fire residues in the form of charcoal and wood ash may better emulate natural disturbance processes in managed boreal forests. We examined the effects of a poplar (Populus) wood biochar and a high-carbon wood ash on soil and vegetation in a 3-year experiment in northwestern Ontario, Canada. Both soil amendments increased soil pH and soil Ca levels; high-carbon wood ash also increased soil Cu, Zn, B, S, and Pb. Amendments had large effects on plant community composition, favoring a subset of ruderal species including raspberry (Rubus idaeus L.) and goldenrod (Solidago canadensis L.). The addition of high-carbon wood ash resulted in declines in growth of planted white spruce (Picea glauca (Moench) Voss); a path analysis suggests that this was due to the effects of toxic elements rather than the indirect effects of competition. We conclude that high-carbon wood ash, while qualifying as a type of biochar and having some beneficial effects on soil properties, can enhance toxic metals in boreal forest soils, with negative consequences to early tree growth. Differences in plant species responses to biochars and the potential for toxicity effects and indirect effects mediated by plant competition will require screening and field trials of potential biochars prior to their use in operational forestry and forest restoration. [ABSTRACT FROM AUTHOR]

Survival and growth of Siberian larch (SL, Larix sibirica Ledeb.) were compared with those of three conifer species native to Alberta, Canada: lodgepole pine (LP; Pinus contorta var. latifolia Engelm. ex S. Watson), white spruce (WS, Picea glauca (Moench) Voss), and jack pine (JP, Pinus banksiana Lamb.) at 12, 10, and 3 trial locations, respectively. The average data age was 18 years (range: 3 to 27 years). Survival of SL averaged 4.2% and 6.5% worse than LP and WS, respectively, while it was 5% better than JP. SL grew 25%, 94%, and 23% taller than LP, WS, and JP, respectively. Stem forking rates were similar between SL and LP, WS, and JP. The best seed sources for Alberta were mature trees established in Alberta and Saskatchewan of unknown initial provenances. The Russian Altai Mountain source grew well at high elevations, while the Finnish Raivola performed well in the northern, low-elevation area. Open-pollinated progeny tests of 58 families planted in five diverse locations yielded individual-tree narrow-sense heritabilities and family mean heritabilities for height at age 15 of 0.15 and 0.59, respectively. The type B between-site genetic correlation was 0.44, indicating a strong genotype × environment interaction. SL has performed well in Alberta, and its growth can be further improved by selection and breeding from appropriate seed sources. [ABSTRACT FROM AUTHOR]

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

Survival and growth of planted white spruce (Picea glauca (Moench) Voss) were assessed at year 15 in boreal mixedwood stands of northern Alberta, Canada, in stands that were deciduous-dominated prior to logging or were conifer-dominated. Three overstory retention levels (0%, 50%, and 75% retention) and four site preparation treatments (mound, high speed mix, scalp, and no treatment) were evaluated. In deciduous-dominated stands, planted spruce performed best in the 50% retention; here, stem volume was at least double that of any other retention treatment after 15 years. In contrast, spruce had reduced growth in coniferous-dominated stands in both 50% and 75% retention treatments compared with the 0% retention. Survival of planted spruce was unaffected by level of retention, but survival was lower in coniferous-dominated stands than in deciduous-dominated stands; in the coniferous-dominated stands, survival was better with mounding and mixing and lowest with scalp treatments. All height variables tended to be greater in the mix and mound site preparation treatments. Finally, the best estimates of future total growth (regenerated spruce and deciduous combined) in the coniferous-dominated stands were in the clearcut treatment. In terms of regenerated spruce growth, the best estimates occurred in the deciduous-dominated – 50% retention stand planted with soil mixing–mounding treatments, where projected growth of spruce was comparable with that of open-grown and tended stands in Alberta's boreal forests. [ABSTRACT FROM AUTHOR]

A two-year field study was carried out to determine whether inoculating white spruce, Picea glauca (Moench) Voss, with a native endophytic fungus, Phialocephala scopiformis DAOM 229536 Kowalski & Kehr (Helotiales, Ascomycota), decreased the performance of eastern spruce budworm, Choristoneura fumiferana Clemens, developing on these trees. Second instars were reared at three densities in the mid crown and at one density in the lower, mid, and upper crown. Larval survival (i.e., survival of larvae to pupation) was lower on endophyte-inoculated trees than on control trees in the mid crown and especially the upper crown but was similar in the lower crown, resulting in a significant interaction between endophyte and crown level. A similar but marginally insignificant interaction was observed for overall survival up to adult emergence (i.e., total survival). Larval survival and total survival were approximately 22% and 19% lower, respectively, when developing in the upper crown of endophyte-inoculated trees than in control trees. Larval survival remained relatively constant, with increased density on control trees but decreased with density on endophyte-inoculated trees, resulting in a significant interaction between endophyte and larval density. Sex ratios of emerged adults and wing lengths of emerged females were not influenced by the endophyte. Our results suggest that endophytic fungi could be useful additions to integrated pest management programs. [ABSTRACT FROM AUTHOR]

Many plant species reproduce by cloning if environmental conditions are unfavorable for sexual reproduction. To test the alternative hypotheses, whether cloning is an "exit strategy" or caused by selection, clonal growth in white spruce (Picea glauca (Moench) Voss) was investigated in three stands in Alaska, each consisting of a core (closed forest) plot and an edge (tree-line) plot. In total, 2571 trees were mapped and genotyped with 11 single sequence repeat (SSR) markers. The proportion of clonal trees follows a moisture gradient and was lowest in the dry Interior basin (4.5%), followed by the sites at the Alaska Range (9.0%) and the Brooks Range (21.7%). At the two latter sites, clonal growth was more frequent in the edge plot. A comparison among 960 aged trees revealed that clonal growth becomes more likely with increasing age and continues over the life span of a tree. Genetic data do not indicate any genetic predisposition for cloning. Clonal growth in white spruce most likely takes place via layering and depends on environmental conditions. Because performance of the trees, and therefore likely plant reproductive success, is lower in plots with a high proportion of clones, selection for clonal growth seems to be highly unlikely. [ABSTRACT FROM AUTHOR]

Retention harvests may leave retention in aggregations evenly dispersed or in various combinations. Although the relative efficacy of aggregated and dispersed retention for biodiversity conservation is widely debated, there has been little study of combinations. We studied this question relative to saproxylic beetles that initially colonize horizontal and upright coarse woody material (CWM) of white spruce (Piceaglauca (Moench) Voss) in boreal mixedwood stands at the Ecosystem Management Emulating Natural Disturbance (EMEND) in northern Alberta 12–13 years after retention prescription. Neither species richness nor emergence of beetles differed among freshly cut bolts exposed as "horizontal" or "upright" in retention patches of two sizes (0.20 ha and 0.46 ha) that were surrounded by different levels of dispersed retention (2%, 20%, and 50%). However, species composition in retention patches differed significantly from those in unharvested controls, except in the larger patches surrounded by 50% dispersed retention, although patterns differed among feeding guilds. Both mycetophage and predator assemblages in patches were similar to those in unharvested controls, suggesting that even relatively small patches retain these guilds regardless of the surrounding matrix quality. Because species composition differed between horizontal and upright bolts, an appropriate mix of horizontal and upright CWM may contribute to conservation of saproxylic beetle faunas. [ABSTRACT FROM AUTHOR]

Given a seed source, the quality of available substrates is a key factor in determining the success of white spruce ( Picea glauca (Moench) Voss) natural regeneration. We examined the influence of substrate and competing vegetation on survival and growth of natural regeneration of white spruce up to 4 years following harvesting in deciduous-dominated upland boreal mixedwood sites. Feather moss, thick soil surface organic layers, litter, and solid wood were poor substrates for establishment. Early successional mosses establishing on mineral soil, thin organics, and rotten wood were generally favourable microsites but were not highly available on postharvest sites. Mineral soil substrates were not as suitable as expected, likely because on a postlogged site, they are associated with unfavourable environmental characteristics (e.g., low nutrient availability, exposure). There was some evidence that survival and growth of seedlings were improved by surrounding vegetation in the first years, but heavy competing vegetation had a negative impact on older seedlings. Burial by aspen litter greatly increased seedling mortality, especially when combined with a brief period of submergence due to heavy spring snowmelt. The results provide insight into conditions under which natural regeneration could be an option for establishing white spruce following harvesting of deciduous-dominated boreal mixedwood forests. [ABSTRACT FROM AUTHOR]

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

We investigated relationships between phosphorus (P) availability and fractionation and growth of lodgepole pine ( Pinus contorta var. latifolia Engelm. ex S. Watson) and white spruce ( Picea glauca (Moench) Voss) in the Athabasca oil sands region. The study sites had peat - mineral soil mix (PMM) as cover soil over a tailings sand substrate planted to pine or an overburden substrate planted to spruce. Available P was determined using a modified Kelowna extraction and sequentially extracted to obtain exchangeable, Fe- and Al-bound, Ca- and Mg-bound, organic, and residual P. The organic P content in PMM explained 83% and 65% of the variations in available P in the pine and spruce sites, respectively. Phosphorus bound to Fe and Al in PMM was also positively related to available P on pine sites ( p < 0.001) but not on spruce sites. Available P in PMM was correlated with aboveground biomass increment of spruce ( p < 0.05) and explained 78% of the variation in foliar P. Foliar P in both pine and spruce was lower than the adequate level for tree growth. Low foliar P and available P in PMM affect the successful establishment of pine and spruce trees on the reclaimed sites, and proper P nutrition management is important to overcome P limitation in reclaimed soils. [ABSTRACT FROM AUTHOR]

Site trees used to estimate site index are selected based on characteristics that ensure that the tree reflects the potential productivity of the site. Hidden leader damage can make it difficult to identify site trees. Using these trees as site trees could lead to erroneous estimates of site index and height growth trajectories. One hundred and fifteen white spruce ( Picea glauca (Moench) Voss) trees were selected, harvested, and split open to identify hidden damage and to quantify the effect of the damage on height growth and site index. A mixed-effects height growth model based on the Chapman-Richards function was formulated. A height growth modifier was included in the model to estimate the effect of leader damage on height growth. It was found that height growth was reduced by 28% in the year that the damage occurred and by 6% and 3% in the following two years. This results in a reduction of about 0.16 m in site index per incidence of damage on average, although this will depend on the age when the damage occurred and the timing between damage events. Since the damage is not outwardly visible, this creates problems when developing and applying site index models. [ABSTRACT FROM AUTHOR]

Conifer winter damage results primarily from loss of cold hardiness during unseasonably warm days in late winter and early spring, and such damage may increase in frequency and severity under a warming climate. In this study, the dehardening dynamics of lodgepole pine ( Pinus contorta Dougl. ex. Loud), jack pine ( Pinus banksiana Lamb.), white spruce ( Picea glauca (Moench) Voss), and black spruce ( Picea mariana (Mill.) B.S.P.) were examined in relation to thermal accumulation during artificial dehardening in winter (December) and spring (March) using relative electrolyte leakage and visual assessment of pine needles and spruce shoots. Results indicated that all four species dehardened at a similar rate and to a similar extent, despite considerably different thermal accumulation requirements. Spring dehardening was comparatively faster, with black spruce slightly hardier than the other conifers at the late stage of spring dehardening. The difference, however, was relatively small and did not afford black spruce significant protection during seedling freezing tests prior to budbreak in late March and early May. The dehardening curves and models developed in this study may serve as a tool to predict cold hardiness by temperature and to understand the potential risks of conifer cold injury during warming-freezing events prior to budbreak. [ABSTRACT FROM AUTHOR]

X-ray computed tomography (CT) of logs means possibilities for optimizing breakdown in sawmills. This depends on accurate detection of knots to assess internal quality. However, as logs are stored, they dry to some extent, and this drying affects the density variation in the log and therefore the X-ray images. For this reason, it is hypothetically difficult to detect log features in partially dried logs using X-ray CT. This paper investigates the effect of improper heartwood-sapwood border detection, possibly due to partial drying, on knot detection in jack pine ( Pinus banksiana Lamb.) and white spruce ( Picea glauca (Moench) Voss) logs from New Brunswick, Canada. An automatic knot detection algorithm was compared to manual reference knot measurements, and the results showed that knot detection was affected by detected heartwood shape. It was also shown that logs can be sorted into two groups based on how well the heartwood-sapwood border is detected to separate logs with a high knot detection rate from those with a low detection rate. In that way, a decision can be made whether or not to trust the knot models obtained from CT scanning. This can potentially aid both sawmills and researchers working with log models based on CT. [ABSTRACT FROM AUTHOR]

Eastern dwarf mistletoe ( Arceuthobium pusillum Peck) parasitizes the branches of white spruce ( Picea glauca [Moench] Voss) and ultimately kills this host species. Previous studies of host needle physiology generally focused on trees experiencing infections of moderate severity; here, we expanded upon previous research to investigate the morphological, physiological, and biochemical properties of needles from white spruce trees experiencing the full range of infection severities. We found that the impact of parasitism on host needle parameters did not scale uniformly with infection severity. Rather, parasitism reduced host needle size distal to infection and resulted in more negative carbon isotope ratios across all infection severities. Reduced foliar nitrogen and chlorophyll content were only evident in trees with moderate or severe infection, while photosynthetic capacity was depressed in only severely infected trees. Furthermore, while some impacts were confined to infected host branches, others, such as smaller needles and reduced photosynthetic capacity, were also evident in branches showing no visible signs of infection. We conclude that symptoms of infection appearing only after infection worsened in severity or appearing in uninfected branches may in whole or in part also be manifestations of a decline in resource availability and vigor of the whole tree. [ABSTRACT FROM AUTHOR]

Losses in forest productivity through poor soil management are typically evaluated by changes in crop production, but conflicting growth responses among co-occurring species can challenge criteria for sustainability. In this study, we evaluate species response to compaction and organic matter removal by contrasting the growth and foliar attributes at age 20 of lodgepole pine ( Pinus contorta Dougl. ex Loud.) and hybrid white spruce ( Picea glauca × engelmannii (Moench) Voss) in sub-boreal forests of central British Columbia. Bole volume increment was stable for lodgepole pine, averaging a 6 % difference across treatments. White spruce, in contrast, often had large growth increases on forest floor retained - compacted plots (average of 63 % gain) but reductions in productivity on the forest floor removed - compacted plots (40 % decline). Foliar nitrogen concentrations converged across treatments for both species, despite the removal of over 50 % of the site N capital. Some key differences in species traits were suggested by foliar δ15N, attributed to deeper rooting of pine, and foliar δ13C, indicating possibly higher CO2 assimilation potential for spruce. The implication is that standard metrics of sustainable forestry are contingent upon tree autecology and that comprehensive assessments of land management require a measure of collective species response. [ABSTRACT FROM AUTHOR]

White spruce ( Picea glauca (Moench) Voss) cone crops were measured at five regional centers in southern and central Yukon for 30 years at one site from 1986 to 2015 and at four other sites during 9 to 11 years to select the best climatic model that uses cues from growing season temperature and rainfall to predict the size of cone crops. We evaluated six climatic models that use summer temperature and rainfall of years t - 1 and t - 2 to predict cone crops in year t. July temperatures provided the best predictors of white spruce cone crops, and no rainfall variable was related to the size of cone crops. We explored three variants of July temperatures: mean temperature, degree-days > 5 °C, and maximum temperatures. For each of these, we used the Δ T model that uses the difference in the July temperature measures of years t - 1 and t - 2. We compared the resulting six models with corrected Akaike's information criterion (AICc) to determine their relative predictive performance. The best model combined Δ T measures of degree-days > 5 °C and the four highest daily maximum July temperatures with R2 = 0.65. By comparison, the Δ T model involving only mean July temperatures was less successful ( R2 = 0.49). There was good regional synchrony ( rp = 0.7 to 0.8) in high cone crops over southern and central Yukon during 1986 to 2015. [ABSTRACT FROM AUTHOR]

Fluorapatite-solubilizing bacteria were isolated from the hyphosphere of the ectomycorrhizal (ECM) fungus Wilcoxina sp., a dominant species in the Picea glauca (Moench) Voss rhizosphere. Hundreds of strains isolated from the ascomycete Wilcoxina sp. could dissolve tricalcium phosphate, while only 27 of them could produce clarification halos on fluorapatite-amended solid medium. Most of the fluorapatite-solubilizing strains belonged to the Burkholderia genus. Scanning electron microscopy observations have shown that these efficient phosphate-solubilizing bacteria (PSB) were able to completely solubilize fluorapatite crystals within 22 h. The efficient PSB Burkholderia sp. strain 205 and Curtobacterium sp. strain 168 were tested for their ability to associate with a genetically distant fungal host while fulfilling their phosphate-solubilizing function. Burkholderia sp. strain 205 successfully associated with the basidiomycete Laccaria bicolor (Maire) P.D.Orton when hydroxyapatite was the only phosphorus source available to the fungus, while there was no bacterial development when Laccaria bicolor could access soluble phosphorus as well. Optical microscopic observation of Laccaria bicolor associated with Burkholderia sp. revealed extensive colonization of fungal hyphae by the bacterium. These results suggest an important role of bacteria - ECM fungi associations in white spruce phosphate nutrition. [ABSTRACT FROM AUTHOR]

Carbon isotope ratio ( δ13C) has been used as an indicator of water stress because plants discriminate less against 13C when under stress. The depth-to-water (DTW) topographic index provides an estimation of soil moisture based on topographic position and other characteristics of a site. To evaluate whether DTW and carbon isotope discrimination were related and to determine if these relationships are influenced by climate, we sampled three time periods, which differed in the amount of annual precipitation (MAP), from tree cores collected from 42 trembling aspen and 43 white spruce trees growing along DTW gradients at two locations in central Alberta, Canada. Increasing MAP led to lower δ13C, indicating less drought stress as water availability increases, while δ13C increased with DTW up to a threshold value, after which the relationship levelled off, suggesting that higher DTW values represent stress-inducing soil conditions. DTW and MAP were then combined into models (aspen, R2 = 0.72; spruce, R2 = 0.44) that could be used to delineate drought-prone areas during periods of low MAP. Tree height and diameter were also related to DTW, suggesting a functional relationship between an index capturing soil properties and tree size. Our results demonstrate the potential to use the DTW index as a measure of site conditions and to predict stand-level responses. [ABSTRACT FROM AUTHOR]

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

Mixed-species stands present a number of opportunities for and challenges to forest managers. Boreal mixedwood stands in eastern Canada are often characterized by a dominant canopy of shade-intolerant aspen ( Populus tremuloides Michx.) with more shade-tolerant conifers in the mid- to sub-canopy layers. Because the aspen and conifer components often attain optimal merchantable sizes at different moments in stand development, there is an interest in developing silvicultural practices that allow partial or total removal of aspen and favour accelerated growth of residual conifers. We tested four partial harvesting treatments in mixed aspen - white spruce ( Picea glauca (Moench.) Voss) stands in which different proportions of aspen (0%, 50%, 65%, and 100% basal area) were removed. Ten years after treatments, 72 spruce stems representing dominant, co-dominant, and suppressed social classes were destructively sampled for stem analysis. Using linear mixed effect models, we analyzed growth as a function of treatment intensity, time since treatment, social status, pretreatment growth rate, and neighbourhood competition. Relative to control stands, radial and volume growth responses were detected only in the extreme treatment of 100% aspen removal. In relative terms, suppressed trees showed the greatest magnitude of cumulative growth increase. Compared with control trees, average annual radial and volume increments were, respectively, 23.5% and 7.1% higher for dominant trees, 67.7% and 24.1% higher for co-dominant trees, and 115.8% and 65.6% higher for suppressed trees over the 10 years after treatment. Growth response was proportional to pretreatment growth rate, and among neighbouring trees, only coniferous neighbours had a negative effect on white spruce growth. Our results suggest that in similar mixed-stand conditions, relatively heavy removal of overstory aspen accompanied by thinning of crowded conifers would result in greatest growth response of residual spruce stems. [ABSTRACT FROM AUTHOR]

Planting white spruce ( Picea glauca (Moench) Voss) under established aspen ( Populus tremuloides Michx.) stands has substantial potential for regenerating mixedwood ecosystems in the western Canadian boreal forest. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. Under future climatic regimes with more frequent and severe drought episodes, underplanting may be a cost-effective strategy for lowering the risk of mortality in mixedwood regeneration. We examine the growth of white spruce during the first 18 years after being planted beneath a 39-year-old stand of trembling aspen. Treatments included thinning from over 6000 stems·ha−1 to 3000, 2000, and 1000 stems·ha−1 and fertilization. Initial stimulation of understory vegetation by fertilization had no measureable effect on spruce heights or diameters at year 18. Aspen thinning treatments did not have a significant effect on spruce height growth rates after spruce crowns had emerged above the understory shrub layer due to rapid aspen basal area increases after thinning. Small, but significant, increases for spruce height and diameter were present in the 1000 and 2000 stem·ha−1 aspen thinnings. A much wider range of aspen stand conditions may be suitable for planting spruce to create mixedwood ecosystems than has been previously considered. [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]

White spruce ( Picea glauca (Moench) Voss) and aspen ( Populus tremuloides Michx.) from unmanaged stands in the boreal forest of Alberta, Canada, were examined for two of the main structural assumptions in the process-based model CROBAS: ( i) a constant allometric relationship between foliage mass and crown length and ( ii) a constant relationship between foliage mass and sapwood area. We evaluated these relationships at both at the whole-crown and within-crown levels. Results indicated that for both species, a constant allometric relationship between foliage mass and crown length was maintained at the whole-crown level over a period exceeding the peak mean annual increment of each species. Within the crowns of spruce, foliage mass accumulated faster near the tree apex as total crown length increased. For aspen, the increase in foliage mass per unit crown length for any section within the crown showed greater similarity to the relationship observed at the whole-crown level. The assumption of a constant relationship between foliage mass and sapwood area at the crown base generally held for spruce but showed considerable variation for any given diameter class. For aspen, this assumption did not appear to be appropriate. For both species, there was more foliage mass per unit sapwood area with increasing height from the ground for nearly all tree size classes. This latter finding was in conflict with the pipe model theory but could not be explained by the hydraulic theory of crown architecture, which predicts a decrease in the ratio of foliage mass to sapwood area with increasing path length. [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]

In Isle Royale National Park, heavy moose browsing over the past 100 years has suppressed the regeneration of many tree species, gradually resulting in a shift towards more open forests and savannas. By 1996, 16% of the forests at the southwestern end of Isle Royale had become savanna and another 20% of forests were starting to have canopy breakup. The changes in understory vegetation brought about by savanna formation have received little attention, even though the future of moose and wolf populations on Isle Royale is tied to these vegetation changes. This study examined the vegetation of savannas ranging in age (date since formation) from <10 years to 80 years to examine how the ground flora changes over time from initial forest canopy breakup, to extensive grassland, and finally to a zootic subclimax dubbed 'moose-spruce savanna.' Ordination techniques were used to describe plant communities and to identify environmental variables that influence vegetation development. Nonparametric multiplicative regression was used to predict how these variables influenced individual plants and structure within the communities. Plant communities had a notable shift from forest herbs to ruderal species, especially non-native plants. This succession was influenced strongly by the underlying bedrock and hydrology-altering moisture regimes and plant communities. Picea glauca (Moench) Voss. and Poa pratensis L. competed in a dynamic inhibitory relationship. The former facilitated forest plants while the latter out-competed other plants and promoted open swards. These interactions are dramatically changing the character of Isle Royale's upland plant communities and will have important trophic consequences for the island. [ABSTRACT FROM AUTHOR]

Exponential fertilization has been shown to be a useful technique for improving seedling quality during nursery production. In this study, we evaluated ( i) the impact of exponential vs. conventional fertilization on trembling aspen ( Populus tremuloides Michx.) and white spruce ( Picea glauca (Moench) Voss) seedlings during nursery production and ( ii) the growth performance and nitrogen (N) utilization of those seedlings in response to foxtail barley ( Hordeum jubatum L.) competition after the transplantation of seedlings in a greenhouse experiment. Exponential fertilization with an application rate higher than the conventional fertilization increased the internal N reserve in trembling aspen and white spruce seedlings during nursery production and also increased new leaf, new stem, and old stem biomass and N retranslocation rates in trembling aspen seedlings but not in white spruce seedlings after transplantation. Foxtail barley competition decreased N concentrations in seedlings and soil N uptake by the seedlings; however, increased N retranslocation rates with foxtail barley competition were observed in white spruce seedlings but not in trembling aspen seedlings. Our results suggest that the growth performance of seedlings was improved by N loading, whereas the impact of vegetation management was species specific. [ABSTRACT FROM AUTHOR]