Predicting productivity of trembling aspen in the Boreal Shield ecozone of Quebec using different sources of soil and site information

Site quality index (SQI) of trembling aspen (Populus tremuloides Michx) in the Boreal Shield of Quebec was predicted using two sources of information: (1) mappable permanent site variables derived from permanent sampling plots and other sources and (2) measured soil and site properties including both biological and permanent site variables. General mappable information did not produce reasonable relationships (R² < 0.25) with SQI while measured variables were able to explain much of trembling aspen SQI variability. For the two parent material types found in our study, i.e. fluvial and till, there was no difference between median SQI values between groups. However, different soil and site variables were better at predicting trembling aspen productivity for the individual parent material types. As much as 60% of the variability in trembling aspen productivity was explained when both biological and permanent site variables were considered in stepwise regression models. When treated individually, models developed for fluvial sites better explained trembling aspen productivity compared to models developed for till sites. Moreover, the ability of the model to predict trembling aspen productivity on till sites when using permanent site variables alone, e.g. soil texture, elemental chemistry and elevation was decreased (R² < 0.3). This indicates that the inclusion of biological site variables such as overstory species composition and forest floor properties provide a major contribution to SQI prediction and are necessary to yield high R². Overall, the data indicate that the traditional mapping of landscape attributes such as drainage and deposit as well as inferred soil geochemistry do not contribute much to explaining SQI. At present, field measurements are needed to predict SQI with a reasonable degree of precision within a forest management unit.     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.     

Climate and site characteristics affect juvenile trembling aspen development in conifer plantations across southern British Columbia

In southern British Columbia, juvenile trembling aspen is managed primarily as a competitor with conifers rather than for its ecological and economic value. As a result, brushing treatments have been applied on a widespread basis and this practice is likely to continue in the near future. Given the potential for climate change to affect our valuation of aspen, we require a better understanding of factors that affect its development, its competitive ability with conifers and its responses to brushing. We used data from 11 aspen management experiments to examine the influence of climate and site factors on aspen height, cover, and density in 17–24 year-old control stands and 9–16 years after manual cutting or girdling. Models explained 64% and 89% of the variation in aspen height in control and manually brushed stands, respectively, but were poor for girdling. Increasing length of the frost-free period was associated with increasing aspen height in control stands, whereas drier summer conditions on cool aspects favoured height growth of aspen suckers following manual cutting. We also examined the influence of climate and site factors on three simple competition indices that describe the height and density of aspen relative to conifer height, and then tested how well these indices predicted conifer growth. The density of aspen taller than conifers accounted for 39% of the variation in lodgepole pine diameter and the ratio of aspen/conifer height accounted for 33% of the variation in Douglas-fir height, suggesting that aspen competition was only moderately important to conifer growth. Our findings imply that aspen may become more productive with warmer summers provided it is not limited by summer moisture availability and that mixed stand management is a viable option in southern interior stands.     

Growth of white spruce underplanted beneath spaced and unspaced aspen stands in northeastern B.C.—10 year results

Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39 year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m² ha⁻¹. Thinning of overstory aspen to 1000 or 2000 stems ha⁻¹ did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.     

Corrigendum to: Growth of white spruce underplanted beneath spaced and unspaced aspen stands in northeastern B.C.—10 year results

Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39-year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m² ha⁻¹. Thinning of overstory aspen to 1000 or 2000 stems ha⁻¹ did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.     

Competitive interactions between juvenile trembling aspen and lodgepole pine: A comparison of two interior British Columbia ecosystems

To facilitate ecosystem-specific management of juvenile mixtures of lodgepole pine (Pinus contorta Dougl. Ex Loud. Var. latifolia Engelm.) and trembling aspen (Populus tremuloides Michx.) in south-central British Columbia, we compared the characteristics of pine–aspen competition between a moist sub-boreal spruce and a dry interior Douglas-fir ecosystem. A total of 252 lodgepole pine and their neighbourhoods were examined across four untreated stands, each of which was sampled three times between ages 12 and 24 years. Pine diameter and height decreased with increasing density of trembling aspen at least as tall as the target pine (tall aspen) in both ecosystems. Regression analysis was used to examine the ability of tall aspen density and four competition indices (CIs) to predict pine size. Tall aspen density, which is easily assessed in the field, accounted for 63% and 69% of the variation in pine diameter and height in 20–24 year-old stands, respectively. The most successful competition index, based on the basal diameter ratio (BDR) of trembling aspen to pine accounted for, respectively, 78% and 73% of the variation. In the same stands, R² values were 1–5% lower when tall aspen density and BDR at age 15–19 years were used to predict size of 20–24-year-old pine.     

Post-fire natural regeneration of young stands on clearcut and partial-cut and uncut sites of boreal mixedwoods

Boreal mixedwoods are an important element and the most productive forest type in the Canadian boreal forests. However, they experience frequent disturbances. In order to better understand the responses of boreal mixedwoods to different combinations of anthropogenic and natural disturbances, we investigated the natural regeneration of boreal mixedwoods that were previously subjected to three different harvesting treatments (clearcut, partial-cut and uncut control) and naturally regenerated, but subsequently burnt by a severe natural fire 6 years later. The study was conducted 8 years following the fire. Significant interactions were found among harvesting method, species and block in several regeneration variables. There were a total of 12 woody species (trees and shrubs) regenerated, but not all the species were present in all the sites. In general, the species richness and species diversity of the new stands were lowest on clearcut sites while the differences between partial-cut and control varied with blocks. However, the combined total density for all species was lowest on uncut control sites. Density and regeneration index data show that trembling aspen was the predominant tree species in all stands except at one uncut control site where jack pine was the dominant species. The density of trembling aspen generally declined from clearcut to partial to the uncut control. Pincherry, beaked hazel and mountain maple were the dominant shrub species in the new stands, but no general patterns were found in terms of variations in density with harvesting methods for any of the shrub species. Jack pine and white birch were the tallest tree species in the clearcut treatment while white birch was taller than jack pine in the partial-cut and control. The results suggest that active measures are necessary to restore the complex structure of the initial mixedwoods.     

Effects of thinning on growth of six tree species in north-temperate forests of Lithuania

This paper summarises the results from 35 years-observed thinning experiments on 256 permanent sample plots in 10–60 year-old stands of ash, aspen, birch, oak, pine and spruce in Lithuania. Thinning enhanced crown projection area increment of residual trees. The largest effect was observed in stands of aspen and birch (growth increase by 200%), followed by ash and oak (over 100%), and spruce and pine (about 80%). Thinning also promoted dbh increment, especially in younger stands, and the increase of dbh increment was positively correlated with the thinning intensity. The strongest reaction was exhibited by oak and aspen, while ash, birch and conifers reacted to a lower extent. Low and moderate intensities of thinning stimulated volume production in younger stands while the opposite was observed in older stands with increasing removals. Spruce stands exhibited relatively strongest increase of volume increment and pine, –the weakest, while the effect on deciduous species was intermediate. The results demonstrate that significant increase in volume increment is achievable with thinning of only young forest stands, e.g. 10–20 year-old pine, birch and ash, or 10–30 year-old oak, aspen and spruce.     

Response of aspen stands to forest tent caterpillar defoliation and subsequent overstory mortality in northeastern Ontario,Canada

Overstory mortality, understory tree recruitment, and vegetation development were assessed in trembling aspen (Populus tremuloides Michx.) stands following two recent episodes of forest tent caterpillar defoliation (Malacosoma disstria Hbn.) in northeastern Ontario. The results suggest that poplar (aspen and balsam poplar (Populus balsamifera L.)) mortality increased with consecutive years of insect defoliation occurring from the mid-1980s to mid-2000s and the proportion of poplars in the overstory, but decreased with improved pre-defoliation tree vigour (DBH increment). The first outbreak, which lasted from the mid-1980s to early 1990s, was more severe in terms of insect defoliation and contributed more to poplar mortality and decline. The decline began in the late 1990s and peaked in early 2000s. Poplar regeneration and understory shrubs responded rapidly to foliage loss to insect defoliation and mortality of overstory poplars. The regenerated poplars were able to maintain their growth under developing shrubs and residual overstory canopy and numbers were sufficient to compensate for the poplar trees lost to insect infestation. The defoliation-induced overstory decline will accelerate the transition of aspen stands to conifer dominance through enhanced conifer recruitment and growth, and reduced hardwood overstory in aspen-dominated stands, while hardwood dominance will persist in pure aspen stands. From a timber supply perspective, the decline caused by forest tent caterpillar defoliation could delay the availability of aspen stands for harvesting by 40–50 years.     

Influence of woody and herbaceous competition on microclimate and growth of eastern white pine (Pinus strobus L.) seedlings planted in a central Ontario clearcut

The influence of woody and herbaceous plant competition, either alone or in combination, on microclimate and growth of planted eastern white pine (Pinus strobus L.) seedlings was examined over four consecutive growing seasons in a central Ontario clearcut. Treatments that manipulated the comparative abundance of these two plant functional groups significantly affected light availability, soil moisture, and air and soil temperature regimes. These microclimate alterations, coupled with the relative competitiveness of herbaceous and woody vegetation, corresponded to temporal changes in vegetation cover and dominance. The more rapid colonization and growth of the herbaceous plant community, dominated by bracken fern (Pteridium aquilinum) and ericaceous shrubs (Kalmia sp., Vaccinium sp.), resulted in this form of vegetation being a comparatively important early competitor for soil moisture. As the woody plant community, dominated by naturally regenerated trembling aspen (Populus tremuloides Michx.), grew in height and leaf area, it became a comparatively strong competitor for both light and soil moisture. For all vegetation treatments combined, white pine seedling growth responses were strongly correlated with total cover of competing vegetation and its relative influence on above- and belowground microclimatic variables. Higher total cover of competing vegetation was generally associated with lower light and soil moisture availability and cooler soil temperatures. Multiple regression analyses indicated that pine seedling relative height growth increased with soil moisture content and growing season soil heat sum, while seedling relative diameter and relative volume growth increased with light availability.     

Spatial distribution,architecture, and development of the root system of Pinus banksiana Lamb. in natural and planted stands

In the province of Québec, Canada, the majority of planted jack pine (Pinus banksiana Lamb.) seedlings are produced in rigid wall containers. More than 95% of them exhibit deformations of the root system which may induce stem instability. Studies of the root architecture of planted jack pine have been limited to a 30 cm radius from the stem, as barely any studies have been devoted to naturally regenerated stands. Moreover, only a few researches have focused on temporal evolution of root systems. The aim of the present study was to characterize the architectural, spatial, and temporal development of jack pine roots in natural and planted stands. Study sites were located in the continuous boreal forest of Quebec. The plantation was done in 1987, so that the trees were 15 years old at the time of sampling. Trees from natural stand had regenerated after a fire in 1983 and were 13–16 years old. The root systems of 14 jack pine trees per site were manually excavated up to a <5 mm diameter, without regard to their distance from the stem. The number, length, diameter, and the spatial and temporal development of roots were analyzed according to three scales of root architecture: the root system, axes, and segments. Overall, the numbers and lengths of roots were higher with planted pines. However, naturally regenerated trees displayed a better distribution of their roots around the stem and at depth, combined with more rapid length growth during the first years. In natural stands, all the trees had a taproot and 30% of the main roots originated at a depth of more than 20 cm, and they are regularly distributed around the stems. Planted trees did not present a taproot and 97% of the main roots originated in the first 20 cm beneath the soil surface. Moreover, 50% of root length was located in one-third of the area surrounding the stems, an area that corresponded to the furrow. Finally, the annual development of lateral roots in planted stand displayed a 5-year delay when compared with natural stand, which also affected maximum growth length and development of the branching pattern. Root distribution and temporal development are known to play a major role in the stability of aerial parts. Seedling production methods, container type, site preparation and planting techniques need to be examined in greater detail in order to assess their effect throughout the development of the root system. It is necessary to compare different sylvicultural practices and with natural/planted stands to gain a clearer understanding of this problem.     

Height–diameter equations for boreal tree species in Ontario using a mixed-effects modeling approach

Height–diameter relationships based on stand characteristics (trees/ha, basal area, and dominant stand height) were investigated for balsam fir, balsam poplar, black spruce, jack pine, red pine, trembling aspen, white birch, and white spruce using data from permanent growth study plots in northern Ontario, Canada. Approximately half the data were used to estimate model parameters with the rest used for model evaluation. Multiple Chapman–Richards functions with parameters expressed in terms of various stand characteristics were fit to determine the best models for predicting height.     

The effects of species mixture on the growth and yield of mid-rotation mixed stands of Scots pine and silver birch

The effect of tree species mixture on stand volume yield and on tree-species-specific diameter and height growth rates were analysed in managed mixed stands of Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Ehrn.).Data were obtained from 14 repeatedly measured stands located in Southern Finland on mineral soil sites with varying admixture of Scots pine and silver birch. Statistical analysis was carried out for studying the effect of species mixture on the development of stand characteristics. For the analysis, the plots were categorised into three groups (plot types) according to the species dominance. In order to analyse species-specific growth rates, individual-tree mixed linear growth models for tree diameter and height growth were developed for both tree species.The results clearly show that the yield of the managed mid-rotation, mixed stands was greater for stands dominated by Scots pine than for stands dominated by birch, and the stand volume increment decreased with an increasing proportion of silver birch. Analysis of diameter and height growth by tree species revealed that the main reason for this pattern is the negative impact of birch competition on the growth of pine trees. The increase in diameter of pine was clearly hampered if the proportion of birch was high. An abundance of birch also slightly decreased the growth in height of Scots pine, although the effect was less than on diameter growth. Species mixture did not affect the diameter growth of birch but did have a significant effect on height development. Height growth of birch was considerably greater in pine-dominated stands than in birch-dominated stands. In pine-dominated mixed stands, the height growth of birch was quite close to that of dominant pine trees, and birches can endure in competition with pines for light.The results apply for even-aged and single-storey managed stands, where stocking density and structure are controlled with pre-commercial and commercial thinnings. The results are not applicable to unmanaged mixed stands undergoing self-thinning. This study provides new information on mixed stands from a silvicultural perspective, which can be applied in decisions involving the management of mixed stands.     

The effects of herbaceous and woody competition on planted white pine in a clearcut site

We investigated the effects of herbaceous and woody vegetation control on the survival and growth of planted eastern white pine (Pinus strobus L.) seedlings through six growing seasons. Herbaceous vegetation control involved the suppression of grasses, forbs, ferns, and low-shrubs, and was maintained for 0, 2, or 4 years after white pine seedlings were planted. Woody control involved the removal of all tall-shrub and deciduous trees, and was conducted at the time of planting, at the end of the second or fifth growing seasons, or not at all. Seedling height and basal diameter responded positively and proportionally to duration of herbaceous vegetation control. Gains associated with woody control were generally not significant unless some degree of herbaceous vegetation control was also conducted. Only herbaceous control increased pine crown closure and rate of crown closure. Herbaceous control and the presence of 5000–15,000 stems per ha of young overtopping aspen were associated with reduced weevil (Pissodes strobi Peck.) injury and increased pine height growth. The study suggests that white pine restoration strategies on clearcut sites should focus on the proactive, early management of understory vegetation and the gradual reduction of overtopping cover from woody vegetation to create a seedling light environment that supports acceptable growth with minimal weevil damage.     

Logging residue removal after thinning in Nordic boreal forests: Long-term impact on tree growth

The aim of this study was to determine the effect of whole-tree harvesting (WTH) on the growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as compared to conventional stem harvesting (CH) over 10 and 20 years. Compensatory (WTH + CoF) and normal nitrogen-based (CH + F or WTH + F) fertilisation were also studied. A series of 22 field experiments were established during 1977-1987, representing a range of site types and climatic conditions in Finland, Norway and Sweden. The treatments were performed at the time of establishment and were repeated after 10-13 years at 11 experimental sites. Seven experiments were followed for 25 years.Volume increment was on average significantly lower after WTH than after CH in both 10-year periods in the spruce stands. In the pine stands thinned only once, the WTH induced growth reduction was significant during the second 10-year period, indicating a long-term response.Volume increment of pine stands was 4 and 8% and that of spruce stands 5 and 13% lower on the WTH plots than on CH during the first and the second 10-year period, respectively. For the second 10-year period the relative volume increment of the whole-tree harvested plots tended to be negatively correlated with the amount of logging residue. Accordingly, the relative volume increment decreased more, the more logging residue was harvested, stressing the importance of developing methods for leaving the nutrient-rich needles on site.If nutrient (N, P, K) losses with the removed logging residues were compensated with fertiliser (WTH + CoF), the volume increment was equal to that in the CH plots. Nitrogen (150-180 kg ha⁻¹) or N + P fertilisation increased tree growth in all experiments except in one very productive spruce stand. Pine stands fertilised only once had a normal positive growth response during the first 10-year period, on average 13 m³ ha⁻¹, followed by a negative response of 5 m³ ha⁻¹ during the second 10-year period. The fertilisation effect of WTH + F and WTH + CoF on basal area increment was both smaller and shorter than with CH + F.     

Diversity of northern plantations peaks at intermediate management intensity

Tree diversity is an important component of biodiversity. Management intensification is hypothesized to affect tree diversity. However, evidence to support the relationship between management intensity and tree diversity in northern forests is lacking. This study examined the effects of fertilization, site preparation, and brush control on tree species diversity, shade tolerance diversity and size diversity of jack pine (Pinus banksiana Lamb.), black spruce (Picea mariana [Mill.] B.S.P.), white pine (Pinus strobus L.) and white spruce (Picea glauca [Moench] Voss) plantations, 15 years after planting in Ontario, Canada. Species diversity and shade tolerance diversity were highly correlated, so were diameter size diversity and height size diversity. Fertilization did not affect the tree diversity indices of any plantations. Species diversity and shade tolerance diversity was interactively influenced by site preparation and brush control in the black spruce, white pine, and white spruce plantations, showing that the highest diversity occurred on sites with intensive site preparation without brush control, whereas on sites with brush control, diversity was higher with least intensity of site preparation. However, in the jack pine plantation, neither species diversity nor shade tolerance diversity differed with management intensification, and is attributed to the fast capture of site resources by the planted crop trees of jack pine which minimized establishment of non-crop species. Tree size diversity increased with site preparation intensity in the jack pine and black spruce plantations, while it decreased with brush control in the white pine and white spruce plantations. We concluded that (1) the effects of management intensification on diversity of northern plantations differ with growth habit of planted crop tree species and (2) species diversity and tree size diversity tend to be highest at intermediate levels of silvicultural intensification during the stand establishment phase, supporting the intermediate disturbance hypothesis.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Effects of soil temperature on biomass production and allocation in seedlings of four boreal tree species

One-year old seedlings of trembling aspen (Populus tremuloides Michx.), black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss), and jack pine (Pinus banksiana Lamb.) were subject to seven soil temperatures (5, 10, 15, 20, 25, 30 and 35 °C) for 4 months. All aspen seedlings, about 40% of jack pine, 20% of white spruce and black spruce survived the 35 °C treatment. The seedlings were harvested at the end of the fourth month to determine biomass and biomass allocation. It was found that soil temperature, species and interactions between soil temperature and species significantly affected root biomass, foliage biomass, stem biomass and total mass of the seedling. The relationship between biomass and soil temperature was modeled using third-order polynomials. The model showed that the optimum soil temperature for total biomass was 22.4, 19.4, 16.0 and 13.7 °C, respectively, for jack pine, aspen, black spruce and white spruce. The optimum soil temperature was higher for leaf than for root in jack pine, aspen and black spruce, but the trend was the opposite for white spruce. Among the species, aspen was the most sensitive to soil temperature: the maximum total biomass for aspen was about 7 times of the minimum value while the corresponding values were only 2.2, 2.4 and 2.3 times, respectively, for black spruce, jack pine and white spruce. Soil temperature did not significantly affect the shoot/root (S/R) ratio, root mass ratio (RMR), leaf mass ratio (LMR), or stem mass ratio (SMR) (P>0.05) with the exception of black spruce which had much higher S/R ratios at low (5 °C) and high (30 °C) soil temperatures. There were significant differences between species in all the above ratios (P<0.05). Aspen and white spruce had the smallest S/R ratio but highest RMR while black spruce had the highest S/R but lowest RMR. Jack pine had the highest LMR but lowest SMR while aspen had the smallest LMR but highest SMR. Both LMR and SMR were significantly higher for black spruce than for white spruce.     

A comparison of lodgepole pine responses to varying levels of trembling aspen removal in two dry south-central British Columbia ecosystems

We used manual cutting to manipulate trembling aspen (Populus tremuloides Michx.) density and spatial arrangement in relation to crop lodgepole pine (Pinus contorta Dougl. Ex Loud. var. latifolia Engelm.) on two sites in contrasting dry, cool to cold ecosystems of south-central British Columbia. In the dry, cool interior Douglas-fir ecosystem (IDFdk3), we reduced the density of tall aspen (aspen at least as tall as target pine) to 0 (broadcast removal), 1000, 2500, or 4000 stems/ha when the planted lodgepole pine was 6 years old. Eight years later, pine height/diameter ratio (HDR) was significantly lower in the broadcast removal and 1000 stem/ha treatments than in the control. There were no other significant growth responses and pine survival and vigour were good regardless of treatment. In contrast, in a dry, cold sub-boreal pine spruce ecosystem (SBPSxc) where treatments were applied at a stand age of 11 years, naturally regenerated lodgepole pine stem diameter increased significantly in the broadcast removal treatment relative to the untreated control within 2 years. After 4 years, HDR had declined significantly relative to the control where tall aspen density was ≤1000 stems/ha. There were no significant pine responses where 2500 tall aspen stems/ha were retained or where tall aspen were removed only within a 1-m radius around pine. The greater difference in height (height differential) between aspen and pine at the SBPSxc than the IDFdk3 site may partly explain the differing response of lodgepole pine to treatment. Trends of decreasing sucker density with increasing aspen retention were evident at both sites, but differences were significant (p ≤ 0.05) only at the SBPSxc site.     

Persistent effects of fire severity on early successional forests in interior Alaska

There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.