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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]

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]

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]

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]

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]

There is a growing interest in improving knowledge on wood quality of forest resources to allow for an end-product oriented supply for the wood-processing industry. We investigated the differences of white spruce ( Picea glauca (Moench) Voss) wood traits from 13 ecological regions of the eastern Canadian province of Quebec and identified climatic and geographic variables and tree characteristics that may help predict wood traits in novel approaches to forest inventories. One hundred and eleven wood disks were used to determine wood traits that are related to wood anatomy and to mechanical use of wood. Our results indicate significant differences between ecological regions for most of the investigated wood traits. Samples from several coastal regions showed, for example, wood anatomy that is advantageous for some pulp and paper grades, but they offer comparatively weak estimates for mechanical applications. Ordination and projection to latent structures allowed us to identify maximum temperature, degree days, geographic location, tree height, and tree diameter as important predictor variables for wood traits. The quality of models was variable and R2 values ranged from 0.13 to 0.59. Prediction errors in an independent sample were mostly below 10%, and bias was below 5%. Although the prediction of wood mechanics was less precise, the presented results are an encouraging step toward obtaining a wood trait signature for different environments. [ABSTRACT FROM AUTHOR]

Radial patterns of modulus of elasticity (MOE) were examined for white spruce ( Picea glauca (Moench) Voss) and trembling aspen ( Populus tremuoides Michx.) from 19 mature, uneven-aged stands in the boreal mixedwood region of northern Alberta, Canada. The main objectives were to (1) evaluate the relationship between pith-to-bark changes in MOE and cambial age or distance from pith; (2) develop species-specific models to predict pith-to-bark changes in MOE; and (3) to test the influences of radial growth, relative vertical height, and tree slenderness (tree height/DBH) on MOE. For both species, cambial age was selected as the best explanatory variable with which to build pith-to-bark models of MOE. For white spruce and trembling aspen, the final nonlinear mixed-effect models indicated that an augmented rate of increase in MOE occurred with increasing vertical position within the tree. For white spruce trees, radial growth and slenderness were found to positively influence maximum estimated MOE. For trembling aspen, there was no apparent effect of vertical position or radial growth on maximum MOE. The results shed light on potential drivers of radial patterns of MOE and will be useful in guiding silvicultural prescriptions. [ABSTRACT FROM AUTHOR]

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The purpose of this study was to develop an understanding of the pattern of variation in longitudinal, radial, and tangential shrinkage in two commercially important Canadian wood species using multivariate nonlinear mixed-effects models. Shrinkage data were measured in each growth ring at different heights on trees harvested from a mixed-species Nelder tree-spacing plot located in New Brunswick, Canada. Live crown base was reconstructed based on stem analysis. Cambial age, sampling height, crown characteristics (crown length, crown ratio, and distance to live crown base), and ring width at time of wood formation were used as prediction variables. The results indicated that shrinkage magnitude and pattern were closely related to crown dimensions and stem position at time of wood formation. Trees from wider tree spacings with larger crown lengths and ratios tended to have smaller tangential and radial shrinkage. Longitudinal shrinkage decreased nonlinearly from pith outwards with slower rate changes near the bottom of the trees. The model fixed effects accounted for more variation in jack pine ( Pinus banksiana Lamb.) than in white spruce ( Picea glauca (Moench) Voss) for shrinkage in the same anatomical direction. After further validation, the model developed can be used to provide shrinkage predictions for lumber distortion models within the framework of an individual tree growth model. [ABSTRACT FROM AUTHOR]

This paper calibrates climate controls over radial growth of floodplain white spruce ( (Moench) Voss) and examines whether growth in these populations responds similarly to climate as upland trees in Interior Alaska. Floodplain white spruce trees hold previously unrecognized potential for long-term climate reconstruction because they are the source of driftwood that becomes frozen in coastal deposits, where archeological timbers and beach logs represent well-preserved datable material. We compared ring width chronologies for 135 trees in six stands on the Yukon Flats and Tanana River with temperature and precipitation at Fairbanks from 1912-2001. Our sample contains a stable common signal representing a strong negative relationship between summer temperature and tree growth. We developed a floodplain temperature index (FPTI), which explains half of the variability of the composite chronology, and a supplemental precipitation index (SPI) based on correlation of monthly precipitation with the residual of the temperature-based prediction of growth. We then combined FPTI and SPI into a climate favorability index (CFI) in which above-normal precipitation partially compensates for temperature-induced drought reduction of growth and vice versa. CFI and growth have been particularly low since 1969. Our results provide a basis for building longer chronologies based on archeological wood and for projecting future growth. [ABSTRACT FROM AUTHOR]

Vertical and horizontal patterns of light within gaps and expansion of trees into gaps were examined in young stands dominated by trembling aspen (Populus tremuloides Michx.) in northeastern British Columbia and northwestern Alberta. Lateral growth of branches of aspen edge trees averaged 15.2 cm/year and was similar to crown expansion rates reported for edge trees in eastern North America. Branches growing into gaps were significantly longer than those growing away from gaps and are likely to lead to gap shrinkage and reductions in light within gaps. However, height growth of edge trees was not significantly different from that of trees within the surrounding stands. Models describing relationships between diffuse transmittance and locations within gaps and surrounding stand height were developed in this paper. Sky view angle (SVA) was found to be effective for predicting transmittance to different locations within gaps. When SVA values exceed 1.2 (at the B.C. sites), white spruce (Picea glauca (Moench) Voss) will receive <40% of full sunlight, which is less than optimal for spruce height growth. [ABSTRACT FROM AUTHOR]

In this study, compatible height and diameter increment models were fitted for lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.), trembling aspen (Populus tremuloides Michx.), and white spruce (Picea glauca (Moench) Voss), using the relationship between diameter and height growth. It was assumed that tree diameter increment is directly proportional to height increment, and the proportionality constant is a function of competition and site productivity. The results showed that the fit statistics are comparable with results of other studies, with adjusted R2 ranging from 30% to 50%. A validation test of the models, using independent permanent sample plots data, showed that the short-term predictions of the models for both pure and mixedwood stands are fairly unbiased. The models also gave reasonable average height growth and diameter growth trajectories for pure stands of the three species and also projected long-term mixedwood (aspen – white spruce mixture) volume growth dynamics reasonably well. The models also projected reasonably well (i) the effect of increasing initial stem density on average diameter and height, and (ii) the stand volume compared with an older version the Mixedwood Growth Model (ver. 2000A). It was concluded that explicitly linking tree height and diameter increment models does not only have a solid ecological basis, but it also results in a compatible prediction of tree growth and stand dynamics. [ABSTRACT FROM AUTHOR]

This study investigated the genetic variation in veneer quality of white spruce (Picea glauca (Moench) Voss) and its correlation with tree growth and wood density. A total of 270 sample trees from 35 families were harvested from 36-year-old provenance-progeny trials at two sites through a thinning operation. A 36-cm long bolt was collected from each sample tree at an 8-foot (or 2.45 m) height for this veneer quality study. The results indicate that conversion of fast-grown white spruce into veneer and plywood may present some problems. Low wood density, numerous knots, and possibly a high proportion of juvenile wood appear to be major factors contributing to low veneer stress grading, resulting in a production of low-quality veneer. This study also showed that the environmental factors at the two sites played an important role in determining veneer quality and tree radial growth. The narrow-sense heritability for veneer density, veneer modulus of elasticity, and veneer roughness were 0.62, 0.13, and 0.14, respectively. The results revealed considerable phenotypic variation and relatively high additive genetic variation in the veneer modulus of elasticity. The phenotypic and genetic correlations between ring width and veneer density or veneer modulus of elasticity were negative. A positive phenotypic and genetic correlation was found between veneer density and veneer modulus of elasticity. This suggests that selection for tree volume growth in white spruce would lead to a decrease in wood density and veneer stiffness. [ABSTRACT FROM AUTHOR]

A portable freezer was developed to apply frost to branches of large trees to study their growth and recuperation after frost application under natural conditions. The freezer measures 37.5 63.5 31.5 cm and weighs approximately 3 kg. It consists of two compartments, a freezing compartment and a dry ice compartment. The portable freezer provides a ramp-and-soak freezing pattern using a programmable controller. The nonfreezing temperature plateaus can be set from 1 to 6 C and maintained for 0 to 12 h. The cooling and warming rates can be programmed from 1 to 12 C h?1. Test temperatures can be maintained for a period of time ranging from 0 to 12 h at set temperatures. Freezers were tested without samples under controlled conditions at ambient temperatures of 0, ?5, ?10, 5, 15, 20, and 25 C. Under these conditions, the cooling and warming rates showed a deviation of less than -1 C h?1 at a set rate of 2 C h?1. The freezer provides test temperatures as low as ?38 C and ?47 C at ambient temperatures of 20 and ?10 C, respectively. Freezers were also tested under field conditions on attached branches in mature white spruce (Picea glauca (Moench) Voss) trees under hardening conditions. [ABSTRACT FROM AUTHOR]

Seed orchards for the production of conifer seed in British Columbiaare usually located in areas favorable (warm and dry) for flowering and seed development, often considerably south of the source locationof the parent trees. Differences in environmental conditions betweenseed orchard location and location of origin can affect progeny performance. It is suspected that this is caused by environmental factorsthat affect reproductive processes of parent trees and lead to altered physiological traits (aftereffects). This study examined if aftereffects are present in white spruce (Picea glauca (Moench) Voss). Control pollinations were made at two locations, Red Rock, near Prince George (530N) in central British Columbia, and Kalamalka, near Vernon (500N) in southern British Columbia, on five female trees using a four-male polymix. Identical genotypes through grafting were present at the two locations. Pollen maturing at each site was only used in the polymix at that particular location. Progeny were raised and germination traits, number of needle primordia, greenhouse and field heights, and frost hardiness were evaluated in a common environment. The location of seed development significantly affected all traits evaluated. Height growth aftereffects in the second field season were much less than those observed in the first field season. These results suggest that aftereffects are detectable in white spruce progeny performance in British Columbia. [ABSTRACT FROM AUTHOR]

Three experiments were conducted to examine the effects of Calamagrostis canadensis (Michx.) Beauv. on the growth of Populus tremuloides Michx. Containerized seedlings of P. tremuloides were transplanted into large pots and subjected to (1) three soil temperatures (20, 12, and 60C), (2) root competition with C. canadensis, or (3) presence or absence of C. canadensis litter, each at two nutrient regimes. In thefirst experiment, significant differences (p = 0.0001) in pre- and neo-formed leaf area and root and shoot dry weights were observed for plants subjected to the three different soil temperatures. In experiment two, the presence of C. canadensis significantly (p < 0.001) reduced total biomass, plant height, and root collar calliper of P. tremuloides. In the third experiment, a significant interaction (p = 0.027) between C. canadensis litter and nutrient regime was observed, withthe greatest inhibitory effect on P. tremuloides growth resulting from litter at the low nutrient regime. Low soil temperature had the strongest negative effect on P. tremuloides of the four factors investigated. However, direct competition with C. canadensis, low nutrient conditions, and C. canadensis litter likely would add to the growth suppression of P. tremuloides by this grass. The results of this study have significant implications for the management of P. tremuloides after harvesting. [ABSTRACT FROM AUTHOR]

Seedlings of lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) and white spruce (Picea glauca (Moench) Voss) were grown for 8 years under five different levels of deep tillage. These were, in increasing order of disturbance: control, vibration ripping, deep plowing, deep plowing plus residue, and Marttiini plowing. Five-year growth curves of each tree were partitioned into three orthogonalcomponents: mean, linear, and quadratic trends. Nitrogen and phosphorus concentrations and mass of the current-year needles were analyzed. Deep plowing plus residue significantly reduced pine heights relative to both vibration ripping and the control, whereas spruce heights were less in the control treatment than in the vibration-ripped treatment. Tillage treatments did not affect nitrogen or phosphorus concentrations of current-year pine or spruce needles. In pine, deep plowing plus residue treatment resulted in the lightest needles, and Marttiini plowing, deep plowing and control treatments resulted in the heaviest needles. In spruce, needle mass was lowest for the deep plowing plus residue treatment and similar among the other treatments. Vibration ripping of medium-textured, well to imperfectly drained Gray Luvisols may be recommended to increase spruce seedling height growth, but any of the deep-tillage treatments used cannot be recommended for pine. [ABSTRACT FROM AUTHOR]