Effects of aspen canopy removal and root trenching on understory microenvironment and soil moisture

Effects of three aspen (Populus tremuloides Michx.) canopy removal treatments and root trenching on understory microenvironment and moisture were tested at Parkland and Boreal sites in Alberta, Canada. Aspen canopies moderated air temperature by reducing maximums and increasing minimums, and increased the frost-free period in the understory by reducing radiative frosts. When daily differences were found among canopy treatments, maximum absolute humidity was greater with complete canopy removal. Maximum daily relative humidity was greater in openings at night than with either full or partial canopy cover. Predictably, increasing aspen cover reduced PAR reaching the understory. Soil moisture response was highly variable, changing with site, aspen density and precipitation patterns, but there were only marginal differences due to root trenching. In the Parkland site, soil moisture conservation from aspen canopy and leaf litter effects were masked by tree uptake in most periods, but a net increase in soil water (+5.2%) was observed during drought. Soil and microclimatic conditions in thinned aspen stands suggest potentially favourable production benefits from developing and adopting agroforestry systems in these northern ecosystems.     

Long-term response of planted conifers, natural regeneration, and vegetation to harvesting, scalping, and weeding on a boreal mixedwood site

This study reports 14th-year response of a boreal mixedwood stand to different harvest intensities (uncut, 50% partial cut with and without removal of residuals after 3 years, and clearcut), spot site preparation treatments (none and scalped), and chemical weeding frequencies (none, single, and multiple) in northeastern Ontario. The response variables include the survival and growth of planted white spruce (Picea glauca [Moench] Voss) and jack pine (Pinus banksiana Lamb.), height and density of natural regeneration and shrubs, and cover of shrubs and non-woody vegetation. Harvesting and weeding generally improved survival and growth of planted trees, although white spruce survival did not significantly differ among the three weeding frequencies. Harvesting tended to increase heights of hardwood (mostly trembling aspen (Populus tremuloides Michx.)) and conifer (largely balsam fir (Abies balsamea (L.) Mill.).) natural regeneration, cover and density of shrubs, and cover of herbs, lichens, and ferns. Chemical weeding reduced height, density and cover of shrubs, height and density of hardwood regeneration, and fern cover, but increased moss and lichen cover. Spot scalping did not significantly affect planted seedling, natural regeneration, or the vegetation.Maximum survival and growth of planted white spruce and jack pine were achieved using a combination of clearcutting and multiple weeding. However, partial cutting followed by a single weeding produced acceptable survival and reasonable growth of planted trees, particularly for white spruce. Partial canopy removal alone substantially reduced the amount of hardwood regeneration, relative to clearcutting, but did not adequately suppress understory shrubs. Significant improvement in seedling growth following multiple weedings was evident primarily in the complete canopy removal treatments: 50% partial cut with removal of residuals after 3 years and clearcut. While the effects of harvesting and weeding on planted crop trees found in the 5th-year assessments generally persisted at year 14, survival decreased, likely due to light competition from developing hardwood and shrubs.     

Dependences of longleaf pine （Pinus palustris） natural reproduction on environments above ground

We studied relations between natural seedling reproduction and above ground environment in a longleaf pine ecosystem. Forty-eight 0.05 ha circular plots were sampled under single-tree selection, group-tree selection and control stands in three main longleaf pine areas in south Alabama, USA. We measured six above-ground environment factors, viz. canopy closure, stand density, basal area, average tree height, understory cover and PAR under canopy. We employed forward, back-ward and stepwise selection regression to produce one model. Three main variables：canopy closure, stand density and basal area, were left in the model; light, PAR and understory cover were not incorporated into the model at the 0.10 significance level. Basal area was a positive pa-rameter, while canopy closure and stand density were negative parame-ters. Canopy closure was the main parameter in the model. The model proved to be meaningful, and has potential to provide useful guidance for future work.     

Temporal changes of understory plant community in response to pre- and post-harvesting herbicide treatments and partial cutting in aspen-dominated boreal mixedwood stands

In response to concerns about the effects of traditional timber harvesting practices on biodiversity, we examined the effects of alternative silvicultural systems, including partial cutting and modified herbicide use on understory plant communities in an aspen-dominated mixedwood stand. These alternative silvicultural systems match disturbance rates that, based on the intermediate disturbance hypothesis, would support more diverse understory vegetation communities than uncut or clear-cut forests treated with a broadcast spray. Our results indicated that both understory vegetation cover and number of plant species increased at 5 and 10 years after timber harvesting in aspen-dominated boreal mixedwood stands. The highest amount of understory vegetation cover were found in the pre-harvesting herbicide spray treatment areas, likely because understory plants were not directly exposed to the herbicide, whereas the most species occurred in the partial cutting treatment, which represented the most diverse stand structure with both harvested and leave corridors. Understory composition by percent cover of individual species at 10 years post-harvesting was affected by all treatment attributes (i.e., level of harvesting removal, type and time of herbicide application, and mechanical site preparation); however, understory vegetation responded the most to harvesting level. Among treatments, the difference in understory composition was largely attributed to changes in understory species of different shade tolerance.     

Influences of gap position, vegetation management and herbivore control on survival and growth of white spruce (Picea glauca (Moench) Voss) seedlings

The boreal mixedwood forest type of the Canadian interior boreal is largely comprised of two dominant tree species: white spruce and trembling aspen (Picea glauca and Populus tremuloides). This forest type is expansive, providing important ecosystem services and economic production, yet such mixtures are difficult to establish after harvests. While aspen resprouts and grows vigorously following disturbance, spruce growth is relatively slow and is often limited by intense competition from associated vegetation, including aspen. To improve management, it is important to understand how environmental and vegetative conditions vary in relation to the competitive-facilitative relationship of spruce-aspen mixtures. In this study white spruce was planted across large canopy openings to determine whether survival and height growth is influenced by position within gap and by differing levels of competing vegetation control of aspen and understory plants. In addition, we addressed the issue of herbivory, which can pose a significant threat to planted spruce seedlings. Within each of four sites, linear gaps were created and five gap positions were recognized spanning the southern and northern forest understories, and southern, center and northern positions within each opening. Three different levels of vegetation management were implemented: a brush saw treatment in which all vegetation was cut to ground level, a mixing treatment in which all vegetation and rootstock was ground up, and a control with no vegetation management. The three herbivory treatments excluded large ungulates, small herbivores (rabbits, hares) or had no herbivore exclusion. Growth and survival of white spruce seedlings were measured for four years (1997-2000). Understory survival was significantly lower than within the gap, with the sheltered southern edge position providing the best initial environmental conditions and or ameliorative cover for spruce establishment. However, after four years the shelter effect starts to be inhibitive relative to center and northern gap positions, suggesting the removal of the canopy is necessary before spruce productivity declines. The optimal vegetation management treatment also changed over the study period. The most intensive treatment (mixing) initially showed a negative influence on survival and growth, but by year four, survival converged to approximately 75% for all treatments, and the mixing treatment produced the best height growth. The growth advantage became most evident in the center gap positions, which initially lagged the brushsaw and control treatments. Lastly, some growth losses from herbivory must be expected in boreal mixedwoods, although not enough to merit control. Results have implications for the timing and intensity of silvicultural treatments for harvesting and planting.     

Regeneration dynamics of planted seedling-origin aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis> Michx.)

Aspen (Populus tremuloides Michx.) is a foundational tree species of the North American boreal forest. After disturbance, clonal aspen stands quickly achieve canopy closure by sending up numerous clonal shoots (root suckers) from their lateral root system. Controlled aboveground disturbance will commonly induce prolific root suckering and thereby increase stem density in clonal aspen stands, but it is unclear if increases in stem density will be observed in planted seedling-origin aspen stands. The objectives of this study were to determine (1) overall root suckering response of planted aspen to aboveground disturbance; (2) if different cut heights of the stem or infliction of root damage impact the number of root suckers produced. We found that planted aspen regenerated readily after disturbance, averaging five root suckers per cut tree. However, individual response was highly variable, ranging from zero to 29 root suckers per root system. Of the cut trees, 75% produced at least one root sucker and 60% produced at least one stump sprout. Cutting trees close to the soil surface produced more root suckers than leaving a 25 cm stump. While root system size (mass and length) was well correlated with aboveground measures of planted aspen, root suckering was not related to root system size. As a result of increased forest reclamation efforts in the boreal forest region the planting of aspen has become a more common practice, necessitating a better understanding of the regeneration dynamics and root suckering potential of these planted and seedling-origin aspen forests.     

Will changes in root-zone temperature in boreal spring affect recovery of photosynthesis in Picea mariana and Populus tremuloides in a future climate?

Future climate will alter the soil cover of mosses and snow depths in the boreal forests of eastern Canada. In field manipulation experiments, we assessed the effects of varying moss and snow depths on the physiology of black spruce (Picea -mariana (Mill.) B.S.P.) and trembling aspen (Populus tremuloides Michx.) in the boreal black spruce forest of western Québec. For 1 year, naturally regenerated 10-year-old spruce and aspen were grown with one of the following treatments: additional N fertilization, addition of sphagnum moss cover, removal of mosses, delayed soil thawing through snow and hay addition, or accelerated soil thawing through springtime snow removal. Treatments that involved the addition of insulating moss or snow in the spring caused lower soil temperature, while removing moss and snow in the spring caused elevated soil temperature and thus had a warming effect. Soil warming treatments were associated with greater temperature variability. Additional soil cover, whether moss or snow, increased the rate of photosynthetic recovery in the spring. Moss and snow removal, on the other hand, had the opposite effect and lowered photosynthetic activity, especially in spruce. Maximal electron transport rate (ETR(max)) was, for spruce, 39.5% lower after moss removal than with moss addition, and 16.3% lower with accelerated thawing than with delayed thawing. Impaired photosynthetic recovery in the absence of insulating moss or snow covers was associated with lower foliar N concentrations. Both species were affected in that way, but trembling aspen generally reacted less strongly to all treatments. Our results indicate that a clear negative response of black spruce to changes in root-zone temperature should be anticipated in a future climate. Reduced moss cover and snow depth could adversely affect the photosynthetic capacities of black spruce, while having only minor effects on trembling aspen.     

The influence of canopy patch mosaics on understory plant community composition in boreal mixedwood forest

We assessed the composition of understory vascular plant communities in relation to the mosaic of canopy patch types, and their associated structure and environment, within unmanaged, mature boreal mixedwood forests in western Canada. Within a 30 km² area, we sampled patches of four different canopy types: conifer-dominated, broadleaf-dominated, mixed conifer-broadleaf, and canopy gaps (total n = 98). There were significant differences in understory composition among the four patch types (based on multi-response permutation procedure (MRPP)) and these were mainly due to differences in relative abundances of understory species. The understory communities of conifer patches were characterized by low abundances of shade intolerant species while shade-tolerant and evergreen species were indicators (based on an indicator species analysis (ISA)). Understory communities under gap and broadleaf patches were characterized by higher abundances of grasses and shade intolerant species. Gap, broadleaf, and mixed patches had higher abundances of certain shrub species than did conifer patches. The patch types also differed in terms of their environmental conditions. Conifer patches had drier, cooler soils and the lowest understory light. Broadleaf patches had the warmest soils while understory light during the leaf-off period was similar to that of canopy gaps. Gap patches had the lowest litter cover and PO₄⁻ availability and the highest light. Seven environmental variables (soil moisture, soil temperature, total light during the leaf-off period, cover of coarse and fine downed woody material, and availability of NH₄⁺ and Ca²⁺) were significantly related to understory species composition (in a constrained ordination by means of a distance-based redundancy analysis (db-RDA); 16.5% of variation in understory community data explained). Even within a single patch type, there was substantial environmental variation that was related to understory species composition. Our study suggests that the mosaic of canopy patches within mixedwood forests supports coexistence of both early and late successional understory plant species in mixedwood stands. Maintaining the mixture of canopy patch types within mixedwood stands will be important for conserving the natural patterns of understory plant composition in boreal mixedwood forests.     

Variability of herbaceous productivity along Nothofagus pumilio forest-open grassland boundaries in northern Chilean Patagonia

In order to develop a general model of aboveground net primary production (ANPP) of herbaceous communities in grazing systems that combine forested and open grasslands in temperate areas, biomass production and a set of biotic and abiotic variables were measured at four adjacent forest and grassland sites in Chile’s northern Patagonia for two consecutive growing seasons. At each site, one transect of 80 m long (40 m in open grassland and 40 m in forest) × 10 m wide was established. ANPP was significantly higher in open grasslands but no gradual change in biomass production was observed from inside the forest towards the open grassland. In open grasslands ANPP was spatially uniform but highly variable between years of contrasting weather conditions, whereas in forests it was more spatially heterogeneous and less variable over time. ANPP was highly correlated with cattle consumption. Structural equation models developed for the whole system confirm that ANPP was driven mainly by photosynthetically active radiation (PAR) and available nitrogen. However, we found important differences between forests and the adjacent open grasslands. In forests ANPP was enhanced by positive feedbacks between the amount of transmitted PAR through the canopy and soil nutrient input via cattle dung deposition. In open grasslands nitrogen availability appeared to be the main limiting factor but also influenced by weather conditions (dryer or wetter years). The coexistence of forests and grasslands patches, with different susceptibility of ANPP to meteorological and soil nutrient availability, highlights the importance of implementing an integrated silvopastoral system with lenga (Nothofagus pumilio [Poepp. & Endl.] Krasser) in northern Patagonia.     

Understory plant communities of boreal mixedwood forests in western Canada: Natural patterns and response to variable-retention harvesting

We examined patterns of variation in richness, diversity, and composition of understory vascular plant communities in mixedwood boreal forests of varying composition (broadleaf, mixedwood, conifer) in Alberta, Canada, before and for 2 years following variable-retention harvesting (clearcut, 20 and 75% dispersed green tree retention, control). Broadleaf-dominated forests differed from mixedwood or conifer-dominated forests in that they had greater canopy cover, litter depth, soil nitrogen, warmer soils, as well as greater shrub cover, herb and shrub richness and diversity (plot scale). In contrast, conifer, and to a lesser extent mixedwood, forest had greater β diversity than broadleaf forest. Overall, mixedwood and conifer forests were similar to one another, both differed from broadleaf forest. Several species were found to be significant indicators of broadleaf forest but most of these also occurred in the other forest types. Understory composition was related to canopy composition and edaphic conditions. Variable-retention harvesting had little effect on understory cover, richness, or diversity but resulted in reduced richness and β diversity at a larger scale. The clearcut and 20% treatments affected composition in all forest types. Early successional species and those common in disturbed sites were indicators of harvesting while evergreen, shade-tolerant understory herbs were indicators of the control forest and 75% retention harvest. We conclude that it is important to maintain a range of variation in canopy composition of mixedwood forests in order to conserve the associated understory communities. The presence of conifers in these forests has a particularly important influence on understory communities. The threshold for a lifeboat effect of variable-retention harvesting is between 20 and 75% retention. Examination of richness and β diversity at a variety of scales can provide interesting information on effects of harvesting on spatial reorganization and homogenization of understory plant communities.     

Belowground competition from overstory trees influences Douglas-fir sapling morphology in thinned stands

We evaluated effects of belowground competition on morphology of naturally established coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) saplings in 60- to 80-year-old thinned Douglas-fir stands in southwestern Washington. We separately quantified belowground competition from overstory and understory sources using trenching and understory removal. In this light-limited environment of 26 ± 16% (std. dev.) full sunlight, 2-year exclusion of tree root competition by trenching increased sapling stem biomass by 18%, total aboveground biomass by 21%, number of interwhorl buds by 68%, total foliar biomass by 33%, and foliar biomass on branch components over 4 years old by 143%. Belowground competition did not influence shoot:root ratio or foliar efficiency (i.e., stem growth per unit foliage biomass). Sapling needle size, specific leaf area, and internodal distance also were not affected by belowground competition; these variables were apparently a function of the low-light environment. The principal source of belowground competition was roots of overstory trees; effects of belowground competition from understory vegetation were minor. Thus, under a partial overstory, morphology of Douglas-fir regeneration was influenced by both belowground and aboveground competition from overstory trees. In this environment, understory vegetation control would not likely influence belowground competition to an extent that would affect sapling morphology.     

Interacting effects of canopy gap, understory vegetation and leaf litter on tree seedling recruitment and composition in tropical secondary forests

We experimentally investigated interacting effects of canopy gaps, understory vegetation and leaf litter on recruitment and mortality of tree seedlings at the community level in a 20-year-old lowland forest in Costa Rica, and tested several predictions based on results of previous studies. We predicted that experimental canopy gaps would greatly enhance tree seedling recruitment, and that leaf litter removal would further enhance recruitment of small-seeded, shade-intolerant seedlings in gaps. We created a large (320–540 m²) gap in the center of 5 out of 10 40 m × 40 m experimental plots, and applied the following treatments bimonthly over a 14-month-period in a factorial, split–split plot design: clipping of understory vegetation (cut, uncut), and leaf litter manipulations (removal, addition, control). As expected, experimental gaps dramatically increased tree seedling recruitment, but gap effects varied among litter treatments. Litter addition reduced recruitment in gaps, but enhanced recruitment under intact canopy. Species composition of recruits also differed markedly between gap treatments: several small-seeded pioneer and long-lived pioneer species recruited almost exclusively in gaps. In contrast, a few medium-to-large-seeded shade-tolerant species recruited predominantly under intact canopy. Leaf litter represents a major barrier for seedling emergence and establishment of small-seeded, shade-intolerant species, but enhances emergence and establishment of large-seeded, shade-tolerant species, possibly through increased humidity and reduced detection by predators. Periodic clipping of the understory vegetation marginally reduced tree seedling mortality, but only in experimental gaps, where understory vegetation cover was greatly enhanced compared to intact canopy conditions. Successful regeneration of commercially valuable long-lived pioneer trees that dominate the forest canopy may require clear-cutting, as well as weeding and site preparation (litter removal) treatments in felling clearings. Management systems that mimic natural canopy gaps (reduced-impact selective logging) could favor the regeneration of shade-tolerant tree species, potentially accelerating convergence to old-growth forest composition. In contrast, systems that produce large canopy openings (clear-cutting) may re-initiate succession, potentially leading to less diverse but perhaps more easily managed “natural plantations” of long-lived pioneer tree species.     

Canopy cover effects on nitrogen and phosphorus status in understory vegetation in oak and pine stands

The rate of change of leaf mass, N and P levels in understory vegetation at various levels of canopy cover were measured for 2 years following canopy cover manipulations in northern red oak (Quercus rubra L.) and red pine (Pinus resinosa Ait.) stands in northern Lower Michigan, USA. Canopy cover treatments consisted of clearcut, 25% (50% during first sampling year), 75%, and uncut. Leaf mass, and N and P contents were significantly higher in the clearcut treatment than in other canopy cover treatments, except for the 25% treatment in red pine stands. Leaf N concentrations in understory vegetation were also significantly higher in the clearcut (1991, 20.8mgg^–1; 1992, 22.4mgg^–1) than in the uncut treatment (1991, 16.5mgg^–1; 1992, 16.9mgg^–1). Canopy type (northern red oak and red pine) had little influence on understory nutrient status and leaf mass. In addition, fronds of bracken ferns in all canopy cover treatments in both northern red oak and pine stands were a major sink of nutrients in the understory. The results of this study showed that partial canopy removal generally had only a minor impact on understory leaf production and nutrient status compared with clearcuts during the 2-year period following canopy removal.     

Variation in canopy structure, light and soil nutrition across elevation of a Sri Lankan tropical rain forest

Mixed dipterocarp forests are perhaps the single most important rain forest type in the wet tropics. Only a few studies have purposefully examined differences in resource availability across mixed dipterocarp forest landscapes by simply measuring the abiotic variables of light, soil nutrition and soil water availability in relation to forest structure. We sought to directly measure the environment of canopy gaps across elevation and geology—from lowland mixed dipterocarp forest (100 m amsl) to lower montane dipterocarp forest (1200 m amsl) in southwest Sri Lanka. Middle elevation gap sites (300–900 m amsl) were subdivided into valley, mid-slope and ridge topographic positions. Eighteen natural disturbances all of which were canopy openings caused by tree fall, were randomly selected within primary rain forest that ranged across 100–1200 m elevation. Plots were placed in gap centers and in adjacent understories and measurements taken of forest structure (basal area, canopy height, canopy cover index, CCI), shade (light sensors—photosynthetically active radiation [PAR], canopy hemispherical photographs—global site factor [GSF]) and soil nutrition (pH, exchangeable Al, K, Mg and Ca; Total N; and plant available P). Soil moisture was measured at bi-weekly intervals for five years across middle elevation sites only (300–900 m amsl). Stand basal area, mean canopy height, and canopy cover index all declined with increase in elevation. Understory PAR and GSF decreased with increases in canopy height, basal area and CCI. Size of canopy opening decreased with increase in elevation, but PAR and GSF increased. Valley sites had significantly greater levels of mean percent soil water content as compared to mid-slope and ridge sites of middle elevation sites. However, at the onset of the southwest monsoons in May all sites were similar. Differences were most pronounced during the dry season (December–April). No differences in soil moisture content could be found between gap and understory microsites. K and Ca in gap centers and adjacent forest understories increased with increase in elevation and change in associated geology. pH increased and Al decreased with elevation and associated geology but only for forest understory conditions. Results demonstrate strong differentiation in soil and light resources with elevation that appears related to size of tree-fall disturbance, stature of the forest, topographic position and underlying geology and soil-weathering environment. This suggests that forest management and conservation practices need to develop and tailor techniques and treatments (silviculture) to the forest that emulate and/or account for change in elevation, geology and topographic position. Further studies are needed to identify which are the primary underlying mechanisms (e.g. temperature, wind, soil nutrients, soil moisture availability) defining change in forest structure across elevation.     

Trends in land cover change and isolation of protected areas at the interface of the southern boreal mixedwood and aspen parkland in Alberta, Canada

The Beaver Hills region of central Alberta is located at the interface of the southern boreal mixedwood forest and the aspen parkland, an area now dominated by agriculture, urban and industrial development. Increasing anthropogenic land cover will eventually isolate remaining natural habitats currently protected in parks and reserves. This paper analyzes land cover and land cover change (LCC) in the Beaver Hills moraine and surrounding areas using a structured hierarchical satellite imagery classification applied to Landsat Multi Spectral Scanner and Thematic Mapper from 1977, 1987, and 1998. Our goal was to quantify deforestation and habitat fragmentation trends and then discuss how this information could be used to develop a conservation approach that will protect current areas against further habitat loss. We found that the rate of deforestation in the lands surrounding the moraine was similar to the broad trend at the southern periphery of the Canadian boreal forest region: annual rate of change in forest cover was −0.82%/year. However, in the Beaver Hills there was a net gain of +0.61%/year, due to regeneration of low quality agricultural lands. All fragmentation indices used indicated an increase in forest fragmentation. We designed a network of protected areas and remaining large forest patches, based on the UNESCO-MAB biosphere model. Our results underline concerns regarding the increasing isolation of national parks and biological reserves in Canada.     

Understory vegetation response after 30 years of interval prescribed burning in two ponderosa pine sites in northern Arizona,USA

Southwestern USA ponderosa pine (Pinus ponderosa C. Lawson var. scopulorum Engelm.) forests evolved with frequent surface fires and have changed dramatically over the last century. Overstory tree density has sharply increased while abundance of understory vegetation has declined primarily due to the near cessation of fires. We examined effects of varying prescribed fire-return intervals (1, 2, 4, 6, 8, and 10 years, plus unburned) on the abundance and composition of understory vegetation in 2007 and 2008 after 30+ years of fall prescribed burning at two ponderosa pine sites. We found that after 30 years, overstory canopy cover remained high, while understory plant canopy cover was low, averaging <12% on all burn intervals. We attributed the weak understory response to a few factors – the most important of which was the high overstory cover at both sites. Graminoid cover and cover of the major grass species, Elymus elymoides (squirreltail), increased on shorter fire-return intervals compared to unburned plots, but only at one site. Community composition differed significantly between shorter fire-return intervals and unburned plots at one site, but not the other. For several response variables, precipitation levels appeared to have a stronger effect than treatments. Our findings suggest that low-severity burn treatments in southwestern ponderosa pine forests, especially those that do not decrease overstory cover, are minimally effective in increasing understory plant cover. Thinning of these dense forests along with prescribed burning is necessary to increase cover of understory vegetation.     

Leaf area dynamics of a boreal black spruce fire chronosequence

Specific leaf area (SLA) and leaf area index (LAI) were estimated using site-specific allometric equations for a boreal black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Stands ranged from 3 to 131 years in age and had soils that were categorized as well or poorly drained. The goals of the study were to: (i) measure SLA for the dominant tree and understory species of boreal black spruce-dominated stands, and examine the effect of various biophysical conditions on SLA; and (ii) examine leaf area dynamics of both understory and overstory for well- and poorly drained stands in the chronosequence. Overall, average SLA values for black spruce (n = 215), jack pine (Pinus banksiana Lamb., n = 72) and trembling aspen (Populus tremuloides Michx., n = 27) were 5.82 +/- 1.91, 5.76 +/- 1.91 and 17.42 +/- 2.21 m2 x kg-1, respectively. Foliage age, stand age, vertical position in the canopy and soil drainage had significant effects on SLA. Black spruce dominated overstory LAI in the older stands. Well-drained stands had significantly higher overstory LAI (P < 0.001), but lower understory LAI (P = 0.022), than poorly drained stands. Overstory LAI was negligible in the recent (3-12 years old) burn sites and highest in the 70-year-old burn site (6.8 and 3.0 in the well- and poorly drained stands, respectively), declining significantly (by 30-50%) from this peak in the oldest stands. Understory leaf area represented a significant portion (> 40%) of total leaf area in all stands except the oldest.     

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.     

Spatial and seasonal variability of photosynthetic parameters and their relationship to leaf nitrogen in a deciduous forest

We used gas exchange techniques to estimate maximum rate of carboxylation (V(cmax)), a measure of photosynthetic capacity, in the understory and upper crown of a closed deciduous forest over two seasons. There was extensive variability in photosynthetic capacity as a result of vertical canopy position, species type, leaf age and drought. Photosynthetic capacity was greater in oaks than in maples and greater in the overstory than in the understory. Parameter V(cmax) was maximal early in the season but declined slowly throughout most of the summer, and then more rapidly during senescence. There was also an apparent decline during drought in some trees. Variability in V(cmax) as a result of species or vertical canopy gradients was described well by changes in leaf nitrogen per unit area (N(a)). However, temporal changes in V(cmax) were often poorly correlated with leaf nitrogen, especially in spring and summer and during drought. This poor correlation may be the result of a seasonally dependent fractional allocation of leaf nitrogen to Rubisco; however, we could not discount Rubisco inactivation, patchy stomatal closure or changes in mesophyll resistance. Consequently, when a single annual regression equation of V(cmax) versus N(a) was used for this site, there were substantial errors in the temporal patterns in V(cmax) that will inevitably result in modeling errors.