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Global change in the climate is affecting tree/forest growth. There have been many studies that analyzed climate effects on tree growth. Results presented in these studies showed that the climate had both positive and negative effects on tree growth. The nature (positive/negative) and magnitude of the effects and the climate variables affecting growth depended on tree species. Climate-sensitive diameter growth models are not available for white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss) plantations. These models are needed to project forest growth and yield and develop forest management plans. Therefore, diameter growth models were developed for white pine and white spruce plantations by incorporating climate variables. Four hundred white pine and white spruce trees (200 per species) were sampled from 80 (40 per species) even-aged monospecific plantations (five trees per plantation) across Ontario, Canada. Diameter–age pairs were obtained from these trees using stem analysis. A nonlinear mixed-effects modeling approach was used to develop diameter growth models. To make the models climate sensitive, model parameters were expressed in term of climate variables. Inclusion of climate variables significantly improved model fit statistics and predictive accuracy. For evaluation, diameters (inside bark) at breast height were estimated for three geographic locations (east, west, and south) across Ontario for an 80-year growth period (2021–2100) under three climate change (emissions) scenarios (representative concentration pathway or RCP 2.6, 4.5, and 8.5 watts m−2). For both species, the overall climate effects were negative. For white spruce, the maximum pronounced difference in projected diameters after the 80-year growth period was in the west. At this location, compared to the no climate change scenario, projected spruce diameters under RCPs 2.6 and 8.5 were thinner by 4.64 (15.99%) and 3.72 (12.80%) cm, respectively. For white pine, the maximum difference was in the south. Compared to the no climate change scenario, projected pine diameters at age 80 under RCPs 2.6 and 8.5 at this location were narrower by 4.54 (13.99%) and 7.60 (23.43%) cm, respectively. For both species, climate effects on diameter growth were less evident at other locations. If the values of climate variables are unavailable, models fitted without climate variables can be used to estimate these diameters for both species. [ABSTRACT FROM AUTHOR]

Stand tending using glyphosate to promote coniferous overstory trees has been a common practice in the boreal forests of Alberta. However, there are concerns about the impact of this practice on biodiversity of understory species. This study examined the impact of broadcast glyphosate application during the active reforestation phase, two to several years post-harvest, on forest plant diversity 25 to 40 years post-harvest. Herbicide treatments had the desired effect of shifting tree layer dominance from deciduous to coniferous species, driven by a 25-fold reduction in the density of trembling aspen (3927 vs. 154 stems·ha−1, untreated and treated, respectively). However, understory plant diversity was not significantly different between treated and untreated sites as examined by the Shannon–Wiener (H) and evenness (E) indices. Shared plant species (beta) across sites was high. Of the seven site-indicator species examined, three had significantly lower cover on treated sites: Wild sarsaparilla, low bush-cranberry and oak fern. Total understory plant cover was significantly greater in the treated portion (98.0%) versus untreated (71.4%); however, this difference was not significant when bryophytes were excluded in the analysis. The establishment, maintenance and monitoring of larger long-term trials is strongly recommended. [ABSTRACT FROM AUTHOR]

The timber producing forest land-base in Alberta, Canada, is decreasing due to the increase in allocation of land to other uses such as agriculture, infrastructure, energy development and conservation areas, while the demand for forest products continues to increase. Tree improvement, first implemented in Alberta in 1975, is one strategy used for increasing forest productivity to meet an increasing demand for fibre and forest products. To quantify the operational impact of tree improvement on white spruce productivity, we collected data from 15-year-old white spruce stands in Alberta: 11 stands planted with seedlings from improved (seed orchard) seedlots and seven stands planted with seedlings from wild (unimproved) seedlots. Eight plots (100 m2 circular plots) within each stand were randomly established for measurement (18 × 8 = 144 plots). Diameter at breast height (DBH), tree height, crown width, and branch diameter for each tree inside the plots were measured. Mixed effects models, with site and plot considered random, were used to evaluate the effect of tree improvement on the growth of white spruce. Basal area per ha of competitor species was used to develop a non-spatial competition index, and to compare the growth of improved and unimproved white spruce response to competition. Individual trees from improved stands showed 17% greater DBH and height compared to individual trees from unimproved stands. Individual trees of improved stands also had a greater height and smaller branch diameter for a given DBH than the trees grown in unimproved stands. Overall, tree improvement increased the site index of white spruce by 2 m while both improved and unimproved stands were similarly, and negatively, affected by competition. The application of tree improvement in white spruce resulted in an increase in growth rate, site index, and quality of timber, which will help offset the increasing demand for forest products. • Impact of tree improvement on white spruce was assessed operationally. • 17% greater growth in DBH and height was observed in improved white spruce. • Site index of improved white spruce was increased by 2 m. • Improved and unimproved stands were similarly affected by competition. [ABSTRACT FROM AUTHOR]

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

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]

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]

The boreal forest of Alaska has experienced a small area of forest cuttings, amounting to 7137 ha out of a total of 256,284 ha of timberland in the Fairbanks and Kantishna area of state forest land. Low product values and high costs for management have resulted in a low-input type management with heavy reliance on natural regeneration. Because of increasing demand for wood biomass energy which may reduce rotation ages, understanding post-harvest regeneration is crucial. Harvested areas must meet stocking standards within seven years under the state Forest Resources & Practices Act (FRPA). We evaluated whether state forest harvest units are adequately regenerated up to 40 years following harvest based on FRPA standards in terms of stem density and biomass accumulation. We measured density of all tree size classes, and DBH and height of tree species in 726 plots from 30 representative harvest units, distributed according to harvest and treatment types, harvest year, unit size, and the geographical location of harvests. The majority of regenerated tree stems came from natural regeneration, even on planted units (77%). White spruce ( Picea glauca ) natural regeneration appears to continue for a few decades (seed crops) following harvest. Stem density was below the standard in most units surveyed during the FRPA 7-year period, but far exceeded the standard when resampled in this study (average 16 years later), suggesting either seven years is too early to evaluate tree regeneration, or that a different standard is needed for early surveys. We found a major peak in white spruce stem density (45,000 ha −1 ) in units harvested in 1987 (an historically large spruce seed crop year), suggesting that where possible, foresters need to adjust management plans according to spruce mast years. Post-harvest and post-fire successional patterns are similar, involving rapid establishment and growth of hardwoods and slow growth of white spruce, but post-harvest white spruce recruitment appears to continue longer than post-fire. By 2014 all measured harvest units met FRPA standard under low-input management, but some issues of uniformity of regeneration may remain. Although regeneration density varied among species and by management practices, biomass accumulated steadily over time (60 t ha −1 after 40 years), largely composed of hardwoods, indicating that short-rotation forest management must utilize hardwoods. Our results are based on relatively small harvest units within a matrix of natural forest, and similar results might not occur in landscapes dominated by stands originated from more extensive and intensive management. [ABSTRACT FROM AUTHOR]

Abstract: The commercial use of white spruce varieties produced by somatic embryogenesis (SE) permits increased forest productivity compared to other reproductive technologies. However, the use of SE in clonal forestry requires an accurate assessment of genetic parameters and the performance stability of clones in plantations. For these reasons, two clonal tests were established of 52 white spruce somatic clones. In each clonal tests, we measured survival, bud dormancy, stem form, growth and branching characteristics of clones 4years after outplanting. There was a large variability among clones for characteristics related to growth and branching. At this juvenile stage, the clonal heritability estimates for all characteristics remained low. Of all the characteristics studied, height had the highest heritability. The selection of the top 20 clones (38% of the clones) provided a genotypic gain in height of about 4% for the two planting sites, which is reasonable for such a low selection intensity. High genotypic correlations were observed between growth and branching characteristics. Although a significant site effect was observed for most characteristics, the genotype×site (G×E) interaction was low and consequently the correlation between the two sites for the same characteristic was high. The performance stability of the somatic clones at both sites indicates that opportunities exist for selection of clones that adapt and perform well over different ecological regions, permitting a tangible increase in forest productivity. [Copyright &y& Elsevier]

The world's forests are diverse and serve myriad purposes; however, regardless of the management objective, reliable models of forest stand dynamics, growth and yield are required. The compilation begins with a paper on modeling tree growth using forest inventory data [[1]], which is followed by a presentation on modeling spatial structure changes in white spruce plantations after thinning [[2]]. The focus returns to forest growth modeling with a contribution on the response to mid-rotation treatments in loblolly pine plantations [[4]]. [Extracted from the article]

This study examines the long-term effects of experimental conifer monocultures on stem volume and soil chemistry. Replicated plots of black spruce (Picea mariana), white spruce (Picea glauca), and red pine (Pinus resinosa) were planted in 1950 at three spacings (1.8, 2.7, and 3.6 m) on a briefly cultivated agricultural field on glaciolacustrine sandy loam soil near Thunder Bay, Ontario, Canada. Stem volumes per hectare and per tree were measured in 2002 (52 years after planting). Surface organic (i.e., forest floor) and mineral soil fertility in terms of pH, total N and P, and extractable NH4-N and P were measured in 2000 (50 years after planting). Of the three species red pine had the highest volume per hectare at all the three spacing followed by white spruce and black spruce; volume for all three species peaked at the 1.8-m spacing. The effects of conifer species on soil physical and chemical properties were more pronounced than spacing effects and the changes were mainly confined to forest floor layer. Few changes in mineral soil properties occurred because of the species and spacing treatments. Per hectare forest floor nutrient pools were higher under white spruce and red pine than black spruce, a pattern likely driven by higher litterfall and forest floor accumulation. It appears that toward the end of first rotation the higher productivity of red pine compared with the other conifers did not come at the cost of reduced soil pools of available NH4, P04, or K, but it was associated with reduced Ca levels in the forest floor. [ABSTRACT FROM AUTHOR]

Alterations in the chemical properties of the forest floor following clear-cut harvesting may have implications for forest productivity in boreal stands. We used proximate analysis, carbon-13 (13C) isotopic determination, and cross-polarization, magic-angle spinning (CPMAS) 13C nuclear magnetic resonance (NMR) spectroscopy to examine differences in the characteristics of the forest floors from uncut stands and clear-cut stands dominated by white spruce (Picea glauca (Moench) Voss; SPRUCE) and trembling aspen (Populus tremuloides Michx.; ASPEN) in northern Alberta. Proximate analysis revealed no difference in the chemical properties of forest floors from clear-cut and uncut stands in either stand type, but the acid-insoluble residue of forest floors from clear-cut ASPEN stands was enriched in 13C compared with those from uncut ASPEN stands. CPMAS 13C NMR spectroscopy revealed that forest floors from clearcuts were enriched in total aromatic C, particularly in ASPEN stands, and depleted in phenolic C, particularly in SPRUCE stands. These patterns indicate that forest floors from the clearcuts have become more humified, which may reflect stand-type differences in the amount of labile C available to the forest-floor microbial community and reductions in above- and below-ground inputs to the forest floor following clear-cutting in both stand types. Changes in the chemical properties of forest floors from clear-cut SPRUCE and ASPEN stands could exacerbate C limitation in these soils and alter patterns of nutrient cycling. [ABSTRACT FROM AUTHOR]

In boreal mixedwood forests, aspen (Populus tremuloides) and white spruce (Picea glauca) commonly grow in mixture. These species may avoid competition through differential shade tolerance, physical separation of canopies, phenological differences, successional separation, and differences in soil resource utilization. Aspen may also be able to positively affect the growth of white spruce by improving litter decomposition and nutrient cycling rates, controlling grass and shrub competition, ameliorating environmental extremes, and reducing pest attack. These positive relationships likely make mixed-species stands more productive than pure stands of the same species. The evidence regarding the productivity of pure versus mixed aspen/white spruce stands in natural unmanaged forests is examined in this paper. [ABSTRACT FROM AUTHOR]

Truffles are a staple food source for many forest small mammals yet the vegetation or soil conditions associated with truffle abundance are unknown. We examined the spatial distribution of forest structures, organic layer depth, root density, and two of the most common western North American truffles (Elaphomyces granulatus and Rhizopogon parksii), in managed-young, natural-mature and old-growth western hemlock (Tsuga heterophylla)/Douglas-fir (Pseudotsuga menziesii) stands in Washington State. Forest conditions, E. granulatus and R. parksii sporocarp locations were mapped and analyzed using ARC/ INFO. Spatial patterns were assessed with univariate and bivariate Ripley's K analysis, which measures the scale at which one and two sets of points, respectively, are 'attracted' or 'repelled'. R. parksii truffles were not associated with organic layer depth, root density or forest structure. E. granulatus truffles were distributed in widely-spaced, high-biomass clusters which are significantly associated with thick organic layers with a high density of fine roots. E. granulatus truffles were significantly distanced from trees at 1-2 m. No other associations were found between E. granulatus truffles, logs, ferns or shrubs. Although E. granulatus comprised more than 90% of the total truffle biomass in these unmanaged mature and old-growth stands, in managed-young stands, E. granulatus truffles were rare and total truffle biomass was low. In managed-young stands, organic layer depth and fine root density have been significantly reduced with the introduction of fire. Slash burning and soil scarification practices in these forests may have a strong affect on local food abundance and availability of the most common truffle for small mammal consumers. [ABSTRACT FROM AUTHOR]

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

• White spruce is more severely damaged by growing-season frosts than black spruce. • Tree height was supressed even 13 years after planting. • Microtopography impacted tree height, thus mound planting could favor growth. • Mixed plantations of white and black spruce would increase plantation success. Reforestation in the boreal forest is challenging; trees must survive to large daily temperature variations and to the cold environment. Even if local tree species are adapted to withstand these harsh environmental conditions, spruce plantation failure after artificial regeneration occurs frequently, with important impacts on sustainable forest management. We hypothesized that this regeneration problem is caused by recurrent frost events occurring during the growing season. These events would freeze the terminal bud and the newly formed needles of the planted trees, thus limiting photosynthesis capacity and height growth. Our goal was to identify key permanent physical attributes of the landscape (elevation, slope shape and angular slope) and of microsite conditions (hole vs. mound) that best predict tree height and frost damage to foliage. In summer 2016 and 2017, we sampled tree height of 2,943 white spruce (Picea glauca [Moench] Voss) and black spruce (Picea mariana [Mill.] B.S.P.) trees in 66 monoculture plantations aged between 6 and 13 years distributed in the Clay Belt region of Quebec (Canada), and environment prone to frequent growing-season frosts. Using linear and binomial mixed regression models, we analyzed the effects of the physical attributes of the landscape and of microsite conditions on tree height (linear) and on frost damage (binomial). Tree height increased with increasing elevation and when seedlings were planted on mounds compared to planted in holes. The impact of microsite conditions on tree height increased as plantations aged, but the importance of elevation on tree height decreased with age. The probability of frost damage to foliage decreased for trees planted on mounds compared to trees planted in holes and from concave to convex slopes. These relations were most important in young plantations, but trees showing growth problems were still shorter by 2 m, even 13 years after planting. We also observed differences between species: white spruce was significantly more damaged by frost and was smaller compared to black spruce. Therefore, growing-season frosts can cause growth suppression problems in white spruce plantations established in the boreal mixedwood region. Since microsite conditions also play a key role in driving plantation success, mechanical site preparation techniques should not only focus on reducing the competition between the planted trees and the competing vegetation but should also focus on limiting frost damage by planting trees on elevated microsites. Our results will support forestry practices limiting plantation failure in boreal mixedwoods. [ABSTRACT FROM AUTHOR]

Assisted migration of forest tree populations through reforestation and restoration is a climate change adaptation strategy under consideration in many jurisdictions. Matching climates in which seed sources evolved with near future climates projected for plantation sites should help reduce maladaptation and increase plantation health and productivity. For threatened tree species, assisted migration outside of the species range could help avert extinction. Here, we examine lessons, limitations, and challenges of assisted migration through the lens of three assisted migration field trials of conifers in Canada and Mexico: Pinus albicaulis Engelm., an endangered subalpine tree species in the mountains of western North America; the Picea glauca (Moench) Voss × P. engelmannii Parry ex Engelm hybrid complex, of great economic and ecological importance in western Canada, and Abies religiosa (Kunth) Schltdl. & Cham., a tree species that provides overwintering sites for the monarch butterfly. We conclude that: (a) negative impacts of climate change on productivity of Picea glauca × P. engelmannii may be mitigated by planting seed sources from locations that are 3 °C mean coldest month temperature warmer than the plantation; (b) it is possible to establish Pinus albicaulis outside of its current natural distribution at sites that have climates that are within the species' modelled historic climatic niche, although developing disease-resistant trees through selective breeding is a higher priority in the short term; (c) Abies religiosa performs well when moved 400 m upward in elevation and local shrubs (such as Baccharis conferta Kunth) are used as nurse plants; (d) new assisted migration field trials that contain populations from a wide range of climates tested in multiple disparate climates are needed, despite the costs; and (e) where naturalization of a migrated tree species in recipient ecosystem is viewed as undesirable, the invasive potential of the tree species should be assessed prior to large scale establishment, and stands should be monitored regularly following establishment. [ABSTRACT FROM AUTHOR]

• Climate change may cause reduced forest productivity and higher tree mortality. • We infer white spruce growth limitations from 9795 tree cores from 227 sample sites. • Water deficits make trees vulnerable to climate change in its west-central range. • Future climate projections imply water deficits in the southwest and central boreal. • The north-east may be a climate change refugium due to lack of water deficits. Climate change may cause reduced forest productivity and higher tree mortality due to water deficits that result from increased evapotranspiration. Such limitations may occur in some areas of the North American boreal forest where precipitation is low and warming trends are high. This paper analyzes climatic factors that limit the growth of a commercially important and widespread boreal tree species, white spruce (Picea glauca (Moench) Voss), based on a range-wide dendroclimatological analysis of 9795 trees from 227 sample sites across the North American boreal forest. A bootstrapped response function analysis was conducted for monthly temperature and precipitation variables, and a multivariate regression tree analysis was used to group white spruce populations with similar response coefficients where climatic factors explained 46% of the total variance in response coefficients. The results of this study show that white spruce populations in the west-central boreal forest of North America are the most precipitation-limited group and therefore likely to be most susceptible to climate change. Populations both to the north and south of this group appear less vulnerable, and eastern populations generally do not experience growth-limiting moisture deficits. Regional climate change projections suggest that moisture limitations will become more pronounced in the future, especially for southwest and west-central boreal tree populations. [ABSTRACT FROM AUTHOR]

Trees growing near the Arctic treeline have long been used to reconstruct past climates. However, recent studies have shown deterioration of historically strong positive correlations between air temperature and tree growth (known as "divergence"). Divergence has important implications for confidence in paleoclimate reconstructions and ecosystem–atmosphere carbon exchange. Studies in the Brooks Range of northern Alaska showed that white spruce in the west increased growth in response to late 20th century warming, whereas those in the east failed to show a growth increase. In an earlier study across four watersheds in the Brooks Range, we tested and rejected the hypothesis that divergence in the easternmost watershed reflects moisture limitation of growth. Here, using 16 sites distributed across the same four watersheds, we tested an alternative hypothesis, that greater nutrient limitation in the east may have impeded positive growth responses to warming. Climate comparison across the four Brooks Range study watersheds revealed that, although the easternmost watershed generally had a drier growing‐season climate, the most consistent difference was that winter air temperature and both winter and summer soil temperatures were much colder in the central and eastern watersheds. Soil nutrient availability, foliar nutrient concentrations, and tree growth were all generally lower in the central and eastern than in the western watersheds. Foliar phosphorus concentration was the best predictor of spatial variation in branch extension growth—a finding that is somewhat inconsistent with the theory that forest productivity on young, glacially derived soils should be strongly nitrogen limited. Experimental fertilization yielded the greatest growth increase in the eastern, an intermediate response in the central, and the smallest growth increase in the western watershed, generally mirroring trends in soil temperature, soil nutrient availability, foliar nutrient concentrations, and growth of control trees. Our results confirm that growth in the easternmost watershed is more nutrient limited and suggest that phosphorus limitation may be at least as important as nitrogen limitation of growth. We hypothesize that cold soil effects on tree access to nutrients might explain divergence in the eastern Brooks Range and elsewhere near the Arctic treeline, particularly in areas with cold winters and widespread permafrost. [ABSTRACT FROM AUTHOR]