Fire disturbances are increasing under global climate change and ecological transformations of forests are occurring. Specifically, shifts from productive closed-canopy feather moss forests to low-productivity open-canopy lichen (Cladonia spp.) woodlands have been observed in boreal forests of eastern Canada. It has been hypothesized that high severity of fires would be the cause of this change, but this is difficult to validate a posteriori on mature forest stands. Because charcoal properties are affected by fire severity, we have put forward the hypothesis that the amount and physicochemical properties of charcoal (C, N, H, O, ash, surface area) would be different and indicative of a greater fire severity for open-canopy forests compared to closed canopy ones. Our hypothesis was partly validated in that the amount of charcoal found on the ground of closed-canopy forests was greater than that of open-canopy forests. However, the physicochemical properties were not different, albeit a greater variability of charcoal properties for open canopy stands. These results do not allow us to fully validate or reject our hypothesis on the role of fire severity in the shift between open and closed canopy stands. However, they suggest that the variability in fire conditions as well as the amounts of charcoal produced are different between the two ecosystem types. Furthermore, considering the role that biochar may play in improving soil conditions and promoting vegetation restoration, our results suggest that charcoal may play a role in maintaining these two stable alternative ecosystem states.

In eastern Canada, boreal forests are locally experiencing a shift between two alternative stable states, productive closed-canopy feather moss (Pleurozium schreberi (Brid.) Mitt.) forests to low-productivity open lichen (Cladonia spp.) woodlands. While this shift has important consequences for ecosystem structure and productivity, little is known about the changes occurring in the diversity and composition of the soil microbial community which may be driven by this process. We evaluated the effects of 10-year moss transplantation on soil microbial communities in an open-lichen woodland. Treatments included: 1) removal of the lichen cover, 2) removal of the lichen cover followed by transplantation of a feather moss cover, 3) a control with the lichen cover kept in place (lichen control), and 4) a natural forest site with a feather moss cover (moss control). We found that changing the forest ground cover has a significant impact on the diversity, composition and function of soil microbial communities. Fungal alpha diversity was more sensitive to changes in lichen and moss cover, compared to bacterial diversity. Soil microbial community composition showed significant differences among all forest ground covers, but with greater similarities between the moss transplantation and control moss treatments. More importantly, changes of forest ground cover significantly affected the structure of microbial communities and fungal functional groups. Moss transplantation significantly increased the relative abundance of the organic nitrogen-scavenging fungal genus, Piloderma. Furthermore, moss transplantation significantly increased the overall relative abundance of ectomycorrhizal fungi and decreased the proportion of ericoid mycorrhizal fungi. Soil moisture and temperature were the main environmental variables associated to the shift in microbial community composition. Our study points out that moss transplantation in open-canopy lichen woodlands contributes to regulate and modify the composition, structure, and function of the soil microbial communities with potential implications for explaining the changes in ecosystem processes associated with these two forest ecosystems.

The boreal forest holds the world's largest soil carbon (C) reservoir. A large portion of it is contained in a thick organic layer originating from the slow decay of bryophytes. Because a thick organic layer slows down tree growth, reduces forest productivity, and thereby reduces the potential wood supply, silvicultural treatments that aim to maintain or restore forest productivity after harvesting often involve mechanical site preparation. However, while these treatments can increase growth and C storage in trees, they can also lead to accelerated decomposition of the soil organic matter, reducing C storage. In this study, we assessed the nine-years effect of two silvicultural treatments on soil C dynamics in forested peatlands of northwestern Quebec, compared to unharvested controls: (1) cut with protection of regeneration and soils (CPRS; low soil disturbance, also called careful logging around advanced growth (CLAAG)), (2) CPRS followed by mechanical site preparation (CPRS + MSP, plowing; severe soil disturbance). The mass loss rate of three bryophytes (Pleurozium schreberi, Sphagnum capillifolium, and Sphagnum fuscum) was measured over two growing seasons together with soil organic carbon (SOC) stocks. We also studied the different effects of temperature, water table level, depth, and type of soil layer on mosses decomposition.We observed a significant influence of silvicultural treatments, bryophyte species, and soil layer type (fibric, mesic, humic and mineral) on bryophyte mass loss, which was higher in the CPRS + MSP treatment (21.6 ± 0.13 % standard error) than in control sites (9.5 ± 0.21 %); CPRS alone resulted in an intermediate mass loss of 11.6 ± 0.23 %, for Sphagnum mosses. Bryophyte mass loss was significantly higher in fibric than humic layer. SOC stocks in the uppermost organic soil layer (fibric) were lower in the CPRS + MSP group than in the control group, while the CPRS group was intermediate; however, differences were not statistically significant for the other soil layer and for total SOC. We conclude that while CPRS + MSP accelerates Sphagnum moss decomposition in the topsoil layer, it has limited impact on total soil C stocks that are detectable with stock change methods.

Low productivity open lichen (Cladonia spp.) woodlands have been rapidly expanding in the closed-crown feather moss (Pleurozium schreberi (Brid.) Mitt.) boreal forest of eastern Canada. While open-woodland areas are progressing, there is little information on the recoverability of open lichen woodlands back to closed-canopy forests.

Top-down effects, like predation, are drivers of insect outbreaks, but bottom-up effects, like host nutritional quality, also influence outbreaks and could in turn be altered by insect-caused defoliation. We evaluated the prediction that herbivory leads to a positive feedback on outbreak severity as nutrient concentration in plant tissues increases through improved soil nutrient availability from frass and litter deposition. Over seven years of a spruce budworm outbreak, we quantified litter nutrient fluxes, soil nitrogen availability, and host tree foliar nutrient status along a forest susceptibility gradient. As the outbreak progressed, both soil nutrient fluxes and availability increased which, in turn, improved foliage quality in surviving host trees. This is consistent with boosted insect fitness and increased population density and defoliation as outbreaks grow. Our results suggest that a positive bottom-up feedback to forest ecosystems from defoliation may result in conditions favorable to self-amplifying population dynamics in insect herbivores that can contribute to driving broad-scale outbreaks.

 

Aim

The main objective of this study was to evaluate the influence of leachates from three typical boreal forest ground layers on young tree growth and to explore the linkages between the chemical composition of the leachates, tree growth, the allocation between belowground and aboveground parts, and ectomycorrhizal colonization.

Methods

An original 6-month greenhouse experiment was set up to investigate (i) the effects of lichen (Cladonia spp.) and feather moss (Pleurozium schreberii [Brid.] Mitt.) leachates on jack pine (Pinus banksiana Lamb.) growth and (ii) the effects of feather moss and Sphagnum spp. leachates on black spruce (Picea mariana [Mill.] B.S.P.) growth.

Results

Belowground growth and root allocation was reduced by lichen leachates in 2-year-old pine seedlings, while the impact was significant on both below- and aboveground growth in 6-month-old pine seedlings. A substance having the same migration time as usnic acid was detected in the lichen leachates by high-performance liquid chromatography. Sphagnum spp. and feather moss leachates were not found to have any effect on black spruce seedling growth, despite a higher supply of dissolved inorganic N in the feather moss leachates compared to the leachates of Sphagnum spp. and the control.

Conclusions

These results demonstrate that ground layer composition plays a crucial role in shaping the plant community in boreal ecosystems by influencing the chemical composition of the soil solution. They suggest that chemical interference may be another mechanism by which lichens promote the self-perpetuation of open woodlands in the closed-crown boreal forest.

Following a wildfire, organic carbon (C) accumulates in boreal-forest soils. The long-term patterns of accumulation as well as the mechanisms responsible for continuous soil C stabilization or sequestration are poorly known. We evaluated post-fire C stock changes in functional reservoirs (bioreactive and recalcitrant) using the proportion of C mineralized in CO2 by microbes in a long-term lab incubation, as well as the proportion of C resistant to acid hydrolysis. We found that all soil C pools increased linearly with the time since fire. The bioreactive and acid-insoluble soil C pools increased at a rate of 0.02 and 0.12?MgC?ha?1?yr?1, respectively, and their proportions relative to total soil C stock remained constant with the time since fire (8?% and 46?%, respectively). We quantified direct and indirect causal relationships among variables and C bioreactivity to disentangle the relative contribution of climate, moss dominance, soil particle size distribution and soil chemical properties (pH, exchangeable manganese and aluminum, and metal oxides) to the variation structure of in vitro soil C bioreactivity. Our analyses showed that the chemical properties of podzolic soils that characterize the study area were the best predictors of soil C bioreactivity. For the O layer, pH and exchangeable manganese were the most important (model-averaged estimator for both of 0.34) factors directly related to soil organic C bioreactivity, followed by the time since fire (0.24), moss dominance (0.08), and climate and texture (0 for both). For the mineral soil, exchangeable aluminum was the most important factor (model-averaged estimator of ?0.32), followed by metal oxide (?0.27), pH (?0.25), the time since fire (0.05), climate and texture (?0 for both). Of the four climate factors examined in this study (i.e., mean annual temperature, growing degree-days above 5??C, mean annual precipitation and water balance) only those related to water availability – and not to temperature – had an indirect effect (O layer) or a marginal indirect effect (mineral soil) on soil C bioreactivity. Given that predictions of the impact of climate change on soil C balance are strongly linked to the size and the bioreactivity of soil C pools, our study stresses the need to include the direct effects of soil chemistry and the indirect effects of climate and soil texture on soil organic matter decomposition in Earth system models to forecast the response of boreal soils to global warming.

The increase in open-crown forest stands in the closed-crown boreal forest of Quebec over the last 50 years prompts us to identify and understand the drivers responsible for stand opening. To do so, we studied 37 jack pine plots with varying degrees of canopy opening in the Eastern Canadian boreal forest to answer four questions: (1) Does stand opening result from a deficit in pine regeneration, from poor tree growth, or from both processes simultaneously? (2) In the event that pine stand opening results at least in part from poor tree growth, how early following stand initiation does the tree growth divergence occur between unproductive and productive plots? (3) Is poor tree growth in the unproductive plots related to water stress? Finally, (4) are there predisposing site factors and, if so, what are their contributions versus non-permanent factors such as disturbance history, vegetation, and soil dynamics? In the study area, jack pine stand openings resulted from both a poor regeneration density and weak tree growth. Tree growth divergence between productive and unproductive plots occurred very early during the post-disturbance forest succession and is not likely to result from water limitation during the early development of the trees as revealed by ?13C analysis of tree rings. Low-productivity plots were exclusively found on substrates with low base cation reserves. However, because plots of higher productivity were also found on these substrates, we conclude that stand susceptibility to regeneration failures may be greater on sites with such conditions. Variations in tree cover were mainly related to non-permanent environmental variables, suggesting that restoration of forest productivity is theoretically possible in the low-productivity sites investigated.

Identifying geochemical paleo?proxies of vegetation type in watersheds could become a powerful tool for paleoecological studies of ecosystem dynamics, particularly when commonly used proxies, such as pollen grains, are not suitable. In order to identify new paleological proxies to distinguish ecosystem types in lake records, we investigated the differences in the sediment geochemistry of lakes surrounded by two boreal forest ecosystems dominated by the same tree species: closed?canopy black spruce?moss forests (MF) and open?canopy black spruce?lichen woodlands (LW). This study was designed as a first calibration step between terrestrial modern soils and lacustrine sediments (0–1000 cal yr BP) on six lake watersheds. In a previous study, differences in the physical and geochemical properties of forest soils had been observed between these two modern ecosystems. Here we show that the geochemical properties of the sediments varied between the six lakes studied. While we did not identify geochemical indicators that could solely distinguish both ecosystem types in modern sediments, we observed intriguing differences in concentrations of C:N ratio, carbon isotopic ratio, and aluminum oxide species, and in the stabilization of their geochemical properties with depth. The C accumulation rates at millennial scale were significantly higher in MF watersheds than in LW watersheds. We suggest that these variations could result from organic matter inflows that fluctuate depending on forest density and ground vegetation cover. Further investigations on these highlighted geochemistry markers need to be performed to confirm whether they can be used to detect shifts in vegetation conditions that have occurred in the past.

In the managed portion of the boreal forest of eastern North America, logging has replaced fire as the most important disturbance agent. There, a large proportion of timber is harvested in forests susceptible of accumulating a thick Sphagnum layer that decreases forest productivity, a process called paludification. In such a context, understanding how disturbance type and severity of soil disturbances may affect post-disturbance microhabitat characteristics and understory community composition is critical for forest management. Different management techniques have been used such as careful logging and clearcutting, as well as winter and summer harvests, with various impacts on soils and forest regeneration. In the current study, we used 55 study sites representing a gradient of soil disturbance severity by harvesting (winter and summer) and fire (low and high severity) to compare their impacts on understory plant communities in the Clay Belt area of eastern Canada. At each site, understory community composition (vascular plants, bryophytes, and lichens) was assessed. We found a strong response of communities to overall severity as represented by disturbance type (careful logging, clearcutting, and fire), but little impact of fine scale disturbance severity (winter vs. summer, low vs. high severity disturbance) within each type of disturbance. Differences in community composition were reflected in the abundance of the various plant functional types, with invaders being more common in harvested sites, endurers being common in all disturbances except high severity fires, and avoiders being more common in older sites. Understory communities in harvested sites (<40?years old) were similar to communities typical of old sites originating from natural wildfire disturbances (75–100?years old low severity fires or 200?years old high severity fires) in terms of composition, but also Sphagnum spp. abundance. In order to maintain long-term forest productivity and manage forests in ways that more closely reproduce post-fire conditions, logging operations should aim at increasing soil disturbances, for example by using prescribed burns, in the Clay Belt area of eastern North America.

In the boreal forest, open lichen woodlands have been described as an alternative stable state to closed-crown feather moss forest. In this study, we addressed the role of terricolous lichens in stabilizing open woodlands by hindering tree regeneration and/or growth. Based on field and greenhouse experiments, we compared germination and growth of jack pine (Pinus banksiana) on feather mosses (primarily Pleurozium schreberi) and lichens (primarily Cladonia stellaris), using bare mineral soil as a control. Drivers were investigated by (1) manipulating nutrient supply, (2) simulating shade of a closed canopy on the ground layer with the assumption this would mitigate lichen influence on pine growth, and (3) examining pine root ectomycorrhizal colonization and diversity as indicators of pine ability to take up nutrients. Total growth of 6-month-old greenhouse and 2–3-year-old field seedlings, as well as belowground growth of 2-year-old greenhouse seedlings, was significantly greater in moss than in lichen. Seed germination was not affected by ground cover type. Although field phosphorus and base cation availability was greater in mosses than in lichens, fertilization did not entirely compensate for the negative effects of lichens on pine growth in the greenhouse. Ground layer shading had no impact on pine growth. Lichens were associated with reduced abundance and modified composition of the root ectomycorrhizal community. By suggesting that terricolous lichens constitute a less favorable growth substrate than mosses for pine, our results support the hypothesis that lichens contribute to open woodland stability in the potentially closed-crown feather moss forest.

Wildfire is the primary abiotic disturbance in the boreal forest, and its long-term absence can lead to large changes in ecosystem properties, including the availability and cycling of nutrients. These effects are, however, often confounded with the effects of successional changes in vegetation toward nutrient-conservative species. We studied a system of boreal forested lake islands in eastern Canada, where time since last fire ranged from 50 to 1500 years, and where the relative abundance of the most nutrient-conservative tree species, black spruce, was largely independent of time since last fire. This allowed us to disentangle the effects of time since fire and the dominant vegetation on ecosystem properties, including nutrient stocks and concentrations. Effects of time since fire independent of vegetation composition mostly involved an increase in the thickness of the organic layer and in nitrogen concentration in both soil and leaves. Domination by black spruce had strong negative effects on nutrient concentrations and was associated with a shift toward more fungi and Gram-positive bacteria in the soil microbial community. Path modeling showed that phosphorus concentration was inversely related to organic layer thickness, which was in turn related to both time since fire and black spruce abundance, while nitrogen was more directly related to time since fire and the composition of the overstory. We conclude that discriminating between the effects of vegetation and time since fire is necessary for better understanding and predicting the long-term changes that occur in forest nutrient availability and ecosystem properties.

Forest management strongly influences the carbon (C) budget of boreal forests and their potential to mitigating greenhouse gas emissions. A better quantification of the net changes of carbon pools with time since harvesting is necessary to guide the development of climate-friendly forest management practices. The objective of this study was to assess the evolution of forest C pools, with a special focus on detrital biomass, in an 80-year post-harvesting chronosequence consisting of 36 very homogenous stem-only harvested plots from a humid boreal balsam fir forest of eastern Canada. Dead wood C stocks comprised of snags, stumps, downed woody debris and buried wood averaged 37?Mg?C?ha?1 and evolved according to an upward-facing «boomerang» shape pattern throughout the chronosequence (rapid decrease in the first years followed by a constant increase until the end of the time horizon). In contrast, soil C stocks (FH and mineral) averaged 156?Mg?C?ha?1 and remain constant through time. Stand C sequestration increased rapidly in the early stages up to age 50 when it reached about 250?Mg?C?ha?1, and then continued to accumulate at a slower rate. The temporal trends observed in C pools suggest that C originating from aboveground dead wood (snags, stumps, downed woody debris) is either leaving the system (respired or leached) or transferred into buried wood, and does not appear to influence the C stocks of the fine fraction of the organic and mineral soil horizons. However, the ultimate fate of dead wood C is still poorly understood and further research is needed in this field. We recommend fixing the length of harvest rotation at a minimum of 50?years for this ecosystem to allow the build-up of its dead wood capital, and to promote dead wood retention on site. We also recommend including buried wood in carbon inventories as this pool represents an important share of the detrital C stock in these humid boreal forests.

The accumulation of soil carbon (C) is regulated by a complex interplay between abiotic and biotic factors. Our study aimed to identify the main drivers of soil C accumulation in the boreal forest of eastern North America. Ecosystem C pools were measured in 72 sites of fire origin that burned 2–314 years ago over a vast region with a range of ? mean annual temperature of 3°C and one of ? 500 mm total precipitation. We used a set of multivariate a priori causal hypotheses to test the influence of time since fire (TSF), climate, soil physico?chemistry and bryophyte dominance on forest soil organic C accumulation. Integrating the direct and indirect effects among abiotic and biotic variables explained as much as 50% of the full model variability. The main direct drivers of soil C stocks were: TSF >bryophyte dominance of the FH layer and metal oxide content >pH of the mineral soil. Only climate parameters related to water availability contributed significantly to explaining soil C stock variation. Importantly, climate was found to affect FH layer and mineral soil C stocks indirectly through its effects on bryophyte dominance and organo?metal complexation, respectively. Soil texture had no influence on soil C stocks. Soil C stocks increased both in the FH layer and mineral soil with TSF and this effect was linked to a decrease in pH with TSF in mineral soil. TSF thus appears to be an important factor of soil development and of C sequestration in mineral soil through its influence on soil chemistry. Overall, this work highlights that integrating the complex interplay between the main drivers of soil C stocks into mechanistic models of C dynamics could improve our ability to assess C stocks and better anticipate the response of the boreal forest to global change.

Canada’s boreal forest represents an important contributor of the world’s wood supply industry. However, maintaining or increasing productivity of the boreal forest may be challenging in areas dominated by forested peatlands. Moreover, sustainable management of these forests must also consider other important aspects of the forest ecosystem such as biodiversity and carbon sequestration. To address these concerns, ecosystem-based management has been implemented in some Canadian jurisdictions, such as in regions where a large portion of the boreal forest is dominated by forested peatlands. The objectives of this paper are (1) to summarize our current understanding of how natural disturbances influence stand dynamics and biodiversity in forested peatlands, and (2) to review the main differences between natural and managed forest stands with respect to soil properties, stand productivity, understory plant communities. We also discuss how even-age management regime succeeds or fails to preserve old forests and how this loss affects both forest structure and habitat diversity at the landscape level. We conclude by showing how, in boreal forested peatlands, forest management could contribute to carbon sequestration and mitigate projected climate change.

The composition of the bryophyte layer influences boreal forest regeneration and growth through its effect on soil conditions. Canopy openings in boreal spruce stands can favor the expansion of Sphagnum spp. in the understory at the expense of feather mosses. We use an experimental approach in both paludified field and greenhouse fully-randomized conditions to examine the differential effects of these two ground cover types on black spruce germination and growth, specifically the role of nutrient limitation in generating these effects. We also tested the impact of ground cover shading, simulating the effect of a closed forest canopy on the ground layer, with the assumption that the stress induced to mosses, especially to Sphagnum spp., would have a beneficial impact on tree growth. The two moss types had no differential effects on spruce germination and 0–6-month-old seedling growth in the greenhouse. However, the growth of 2-year-old seedlings in the greenhouse was lower in Sphagnum spp. than in feather mosses. This negative effect was removed by fertilization, suggesting that soil nutrient availability could explain the seedling growth difference between moss types. Greenhouse 2-year-old seedlings also allocated a greater proportion of biomass to roots in Sphagnum spp. than in feather mosses. In the field, feather and Sphagnum spp. mosses had no differential effects on 3-year-old seedling growth, and ground cover shading did not have any short-term positive impact on spruce growth. Although they were not validated in the field, the results we obtained in the greenhouse suggest that the replacement of feather mosses by Sphagnum spp. mosses do not only affect spruce growth through the build-up of an organic layer often associated with low soil temperature and excess water, but also through more direct effects on nutrient availability. Therefore, silvicultural treatments that would favor Sphagnum spp. expansion at the expense of feather mosses, such as partial or total harvesting with protection of regeneration and soils, may result in subsequent tree growth problems even in sites with moderate organic layer accumulation.

Litter decomposition is a major driver of carbon (C) and nitrogen (N) cycles in forest ecosystems and has major implications for C sequestration and nutrient availability. However, empirical information regarding long-term decomposition rates of foliage and wood remains rare. In this study, we assessed long-term C and N dynamics (12–13 years) during decomposition of foliage and wood for three boreal tree species, under a range of harvesting intensities and slash treatments. We used model selection based on the second-order Akaike’s Information Criterion to determine which decomposition model had the most support. The double-exponential model provided a good fit to C mass loss for foliage of trembling aspen, white spruce, and balsam fir, as well as aspen wood. These litters underwent a rapid initial phase of leaching and mineralisation, followed by a slow decomposition. In contrast, for spruce and fir wood, the single-exponential model had the most support. The long-term average decay rate of wood was faster than that of foliage for aspen, but not of conifers. However, we found no evidence that fir and spruce wood decomposed at slower rates than the recalcitrant fraction of their foliage. The critical C:N ratios, at which net N mineralisation began, were higher for wood than for foliage. Long-term decay rates following clear-cutting were either similar or faster than those observed in control stands, depending on litter material, tree species, and slash treatment. The critical C:N ratios were reached later and decreased for all conifer litters following stem-only clear-cutting, indicating increased N retention in harvested sites with high slash loads. Partial harvesting had weak effects on C and N dynamics of decaying litters. A comprehensive understanding of the long-term patterns and controls of C and N dynamics following forest disturbance would improve our ability to forecast the implications of forest harvesting for C sequestration and nutrient availability. © 2017 Springer Science+Business Media New York

In the boreal forest, ground-layer composition may modulate the effects of precommercial thinning (PCT) on stand productivity by affecting tree growth conditions. Based on data from 15 years of PCT monitoring in black spruce (Picea mariana) and jack pine (Pinus banksiana) stands, the objectives of this study were to investigate the effects of PCT on ground-layer composition and the way ground-layer composition is related to tree growth, stand productivity, and the PCT impact on stand productivity. PCT favored lichen expansion in xeric sites. The positive impact of PCT on stand productivity after 15 years was lower in sites with high year-one lichen cover, suggesting that the aboveground positive effect of PCT on growth may have been mitigated by a belowground negative feedback resulting from lichen expansion in xeric sites. Although Sphagnum spp. cover was not affected by PCT, 15-year increase in stand productivity was lower in sites with high year-one Sphagnum spp. cover. These results suggest that xeric stands with high lichen cover should not be targeted for PCT because of either null or negative effects on stand productivity. Subhydric stands with high Sphagnum spp. cover should also be avoided because of lower potential stand productivity. © 2017 Society of American Foresters.

Digital soil mapping (DSM) involves the use of georeferenced information and statistical models to map predictions and uncertainties related to soil properties. Many remote regions of the globe, such as boreal forest ecosystems, are characterized by low sampling efforts and limited availability of field soil data. Although DSM is an expanding topic in soil science, little guidance currently exists to select the appropriate combination of statistical methods and model formulation in the context of limited data availability. Using the Canadian managed forest as a case study, the main objective of this study was to investigate to which extent the choice of statistical method and model specification could improve the spatial prediction of soil properties with limited data. More specifically, we compared the cross-product performance of eight statistical approaches (linear, additive and geostatistical models, and four machine-learning techniques) and three model formulations (“covariates only”: a suite of environmental covariates only; “spatial only”: a function of geographic coordinates only; and “covariates + spatial”: a combination of both covariates and spatial functions) to predict five key forest soil properties in the organic layer (thickness and C:N ratio) and in the top 15 cm of the mineral horizon (carbon concentration, percentage of sand, and bulk density). Our results show that 1) although strong differences in predictive performance occurred across all statistical approaches and model formulations, spatially explicit models consistently had higher R2 and lower RMSE values than non-spatial models for all soil properties, except for the C:N ratio; 2) Bayesian geostatistical models were among the best methods, followed by ordinary kriging and machine-learning methods; and 3) comparative analyses made it possible to identify the more performant models and statistical methods to predict specific soil properties. We make modeling tools and code available (e.g., Bayesian geostastical models) that increase DSM capabilities and support existing efforts toward the production of improved digital soil products with limited data.

At the northernmost extent of the managed forest in Quebec, Canada, the boreal forest is currently undergoing an ecological transition between two forest ecosystems. Open lichen woodlands (LW) are spreading southward at the expense of more productive closed-canopy black spruce–moss forests (MF). The objective of this study was to investigate whether soil properties could distinguish MF from LW in the transition zone where both ecosystem types coexist. This study brings out clear evidence that differences in vegetation cover can lead to significant variations in soil physical and geochemical properties. Here, we showed that soil carbon, exchangeable cations, and iron and aluminium crystallinity vary between boreal closed-canopy forests and open lichen woodlands, likely attributed to variations in soil microclimatic conditions. All the soils studied were typical podzolic soil profiles evolved from glacial till deposits that shared a similar texture of the C layer. However, soil humus and the B layer varied in thickness and chemistry between the two forest ecosystems at the pedon scale. Multivariate analyses of variance were used to evaluate how soil properties could help distinguish the two types at the site scale. MF humus (FH horizons horizons composing the O layer) showed significantly higher concentrations of organic carbon and nitrogen and of the main exchangeable base cations (Ca, Mg) than LW soils. The B horizon of LW sites held higher concentrations of total Al and Fe oxides and particularly greater concentrations of inorganic amorphous Fe oxides than MF mineral soils, while showing a thinner B layer. Overall, our results show that MF store three times more organic carbon in their soils (B+FH horizons, roots apart) than LW. We suggest that variations in soil properties between MF and LW are linked to a cascade of events involving the impacts of natural disturbances such as wildfires on forest regeneration that determines the vegetation structure (stand density) and composition (ground cover type) and their subsequent consequences on soil environmental parameters (moisture, radiation rate, redox conditions, etc.). Our data underline significant differences in soil biogeochemistry under different forest ecosystems and reveal the importance of interactions in the soil–vegetation–climate system for the determination of soil composition.

Strategies for increasing the mobilization of forest biomass supply chains for bioenergy production require continuous assessments of the spatial and temporal availability of biomass feedstock. Using remote sensing products at a 250-m pixel resolution, estimates of theoretical biomass availability from harvest residues and fire-killed trees were computed by combining Canada-wide maps of forest attributes (2001) and of yearly (2002–2011) fires and harvests. At the national scale, biomass availability was estimated at 47 ± 18 M ODT year?1 from fire-killed trees and at 14 ± 2 M ODT year?1 from harvest residues. Mean biomass densities in burned and harvested pixels were estimated at 34 ± 3.0 ODT ha?1 and at 24 ± 1.2 ODT ha?1, respectively. Mean biomass densities also varied dramatically among ecozones, from 14 ODT ha?1 to 206 ODT ha?1 and from 6 ODT ha?1 to 63 ODT ha?1 for burned and harvested pixels, respectively. Spatial averaging with a 100-km radius window shows distinct hotspots of biomass availability across Canada. The largest hotspots from fire-killed trees reached 3.6 M ODT year?1 in the Boreal Shield and the Boreal Plains ecozones of northern Alberta and Saskatchewan, where fires are large and frequent. The largest hotspots from harvest residues reached 1.2 M ODT year?1 in the Montane Cordillera ecozone of British Columbia. The use of spatially explicit remote sensing products yields estimates of theoretical biomass availability that are methodologically consistent across Canada. Future development should include validations with on-the-ground forest inventories as well as the factoring in of environmental, technical and economic considerations to implement operational biomass supply chains.

In the boreal forest, long-lasting canopy gaps are associated with lichens on dry sites and with Sphagnum spp. on wet sites. We hypothesize that ground layer composition plays a role in maintaining gaps through its effects on fine root biomass (Ø?2mm) and soil nutrient availability. Along gradients of canopy openness in both jack pine-lichen and black spruce-moss forests, the relationships between canopy closure, ground layer composition, tree fine root biomass and soil nutrients were analyzed and decomposed using path analysis. The effects of lichen and Sphagnum spp. removal on tree fine root biomass and soil nutrients were tested in situ. Although variations in pine fine root biomass were mainly explained by stand aboveground biomass, lichen removal locally increased fine root biomass by more than 50%, resin extractable soil potassium by 580% and base cations by 180%. While Sphagnum cover was identified as a key driver of stand aboveground biomass reduction in paludified forest sites, its removal had no short-term effects on spruce fine root biomass and soil nutrients. Our results suggest that lichens, unlike Sphagnum spp., affect tree growth via direct effects on soil nutrients. These two different patterns call for different silvicultural solutions to maintain productive stands.

We studied three hybrid poplar plantations in Quebec (Canada) established on sites with varying soil and environmental characteristics to investigate the effects of logging residues on the water potential, carbon isotope ratio and foliar nutrients of planted trees. On each site, four treatments representing different residue loads, as well as treatments aimed at manipulating specific factors of the environment (Herbicide, Geotextile) were applied to test their effects on seedling water potential, carbon isotope ratio and foliar nutrients. Along with analyses of variance, we used structural equation modelling to infer causal relationships of logging residues on height, basal diameter and foliar nutrition of trees through their effects on soil temperature, soil water content and competing vegetation cover. Logging residues decreased soil temperature at all sites and woody plants cover at one site out of three. Height, basal diameter and unit leaf mass were strongly related to each other. Foliar ?13C, N concentration and unit leaf mass increased with decreasing cover of woody plants suggesting an important role of competition for resources. Overall, logging residues had no direct influence on hybrid poplar dimensions after two growing seasons: their effects on the microenvironment of this resource demanding tree species were either cancelling out each other, or were not significant enough to have a significant impact on the growth drivers measured. For example, presence of logging residues might reduce soil temperature, impeding overall seedling performance. Our study highlights the fact that any given silvicultural method aimed at manipulating logging residues has a complex influence involving the interaction of multiple environmental drivers and that the net effect on tree productivity will depend on species and site specific conditions.

1.A broad and diversified group of compounds, secondary metabolites, are known to govern species interactions in ecosystems. Recent studies have shown that secondary metabolites can also play a major role in ecosystem processes, such as plant succession or in the process of litter decomposition, by governing the interplay between plant matter and soil organisms. 2.We reviewed the ecological role of the three main classes of secondary metabolites and the methodological challenges and novel avenues for their study. We highlight emerging general patterns of the impacts of secondary metabolites on decomposer communities and litter decomposition and argue for the consideration of secondary compounds as key drivers of soil functioning and ecosystem functioning. 3.Synthesis. Gaining a greater understanding of plant-soil organisms relationships and underlying mechanisms, including the role of secondary metabolites, could improve our ability to understand ecosystem processes. We outline some promising directions for future research that would stimulate studies aiming to understand the interactions of secondary metabolites across a range of spatio-temporal scales. Detailed mechanistic knowledge could help us to develop models for the process of litter decomposition and nutrient cycling in ecosystems and help us to predict future impacts of global changes on ecosystem functioning.

Fire is considered the major disturbance in boreal forests. Nonetheless, in several areas logging has become the primary driver of forest dynamics. In many areas of the boreal forest, stands may undergo paludification (i.e. the accumulation of thick, poorly decomposed organic layers over the mineral soil) in the prolonged absence of fire, which reduces forest productivity. Whereas high-severity fires (HSF) may restore forest productivity by burning the soil organic layer (SOL), low-severity fires (LSF) mainly burn the soil surface and do not significantly reduce SOL thickness. In the Clay Belt region of eastern Canada, an area prone to paludification, forest stands have historically been harvested by clearcutting (CC), but concerns about the protection of soils and tree regeneration lead to the replacement of CC by careful logging (CL). Whereas CC disturbs the SOL and is thought to favor tree growth, CL has little impact on the SOL. Furthermore, it has been suggested that prescribed burning after clearcut (CCPB) could also be used to control paludification. Using a retrospective approach, this study sought to understand how CC, CL, and CCPB compare to LSF and HSF with respect to soil properties, SOL thickness, vegetation ground cover, tree nutrition, and stand height in paludified black spruce stands of the Clay Belt region. HSF led to significantly taller trees than CL and LSF, but did not differ from CC and CCPB. Foliar N was significantly higher in HSF and CCPB sites relative to CL and LSF, with an intermediate value in CC sites. Ground cover of Rhododendron groenlandicum was significantly lower in HSF and CC sites relative to LSF, with intermediate values in CL and CCPB sites. Sphagnum spp. ground cover was significantly lower in HSF and CCPB sites relative to CL, with intermediate values in CC and LSF sites. High-severity fire sites had a significantly thinner SOL than the four other disturbances. Finally, regression tree analysis showed that SOL thickness represented the best predictor of tree height, whereas segmented regression showed that tree height was negatively correlated to SOL thickness and revealed a cut-off point circa 23 cm, which suggests that tree growth is impeded beyond this threshold. These results support the idea that management strategies intending to regenerate paludified forests should primarily aim at reducing organic layer thickness, either through mechanical disturbance or combustion.

In existing carbon budget models, carbon stocks are not explicitly related to forest successional dynamics and environmental factors. Yet time-since-last-fire (TSLF) is an important variable for explaining successional changes and subsequent carbon storage. The objective of this study was to predict the spatial variability of aboveground biomass carbon (ABC) as a function of TSLF and other environmental factors across the landscape at regional scales. ABC was predicted using random forest models, both at the sample-plot level and at the scale of 2-km2 cells. This cell size was chosen to match the observed minimum fire size of the Canadian large fire database. The percentage variance explained by the empirical sample-plot level model of ABC was 50%. At that scale, TSLF was not significantly related to ABC. At the 2-km2 scale, ABC was influenced mainly by the proportions of cover density classes, which explained 83% of the variance. Changes in cover density were related to TSLF at the same 2-km2 scale, indicating that the increase in cover density following fire disturbance is a dominant mechanism through which TSLF acts upon ABC at the scale of landscapes.

Over long time periods, paludification reduces aboveground productivity resulting in forest retrogression. Paludified forests are typified by intense accumulation of the soil organic layer and a reduction in soil temperatures and nutrient availability. En route to paludification, early successional forests experience large inputs of deadwood biomass during the senescence of the post-fire cohort, much of which may be entombed in this rapidly growing soil organic layer. Here we examined the effects of paludification across a .2000-year chronosequence of black spruce forests on wood decomposition using three complementary approaches.We (1) repeatedly measuredwood density of logs through time, (2) utilize a time-series of logs that varied in time since death, and (3) estimate woody biomass at the stand level as it progresses from live trees to snags, logs and ultimately to buried or decomposed deadwood. Together these approaches demonstrated a 6– 7-year delay before the onset of rapid decomposition.We also found strong evidence that paludification results in a large proportion of logs becoming buried in the soil organic layer. Stand level modeling indicates that the rates of accumulation of buried deadwoodwere greatest following the senescence of the post-fire cohort when both soil organic layer build-up and creation of deadwood peaked. Following this period of high deadwood creation, stands enter a retrogressive state whereby productivity continues to decline albeit more slowly. Continued losses inwoody carbon biomass fromtrees during this retrogressive state are offset by lower wood decomposition rates and a high biomass of accumulated buried deadwood, essentially stabilizing the wood based carbon budget in these ecosystems.We recommend that partial cutting be conducted prior to or near the senescence of the post-fire cohort to improve emulation of natural forest succession in terms of both live tree and deadwood biomass. Furthermore, deadwood during this period has an extremely short residence time and the dynamics of deadwood should recover much quicker than if harvesting is conducted later in succession when there is less live tree biomass and deadwood has longer residence times.

Plant species mixtures are often seen as being able to achieve higher productivity and carbon (C) sequestration than their single-species counterparts, but it is unclear whether this is true in natural forests. Here, we investigated whether naturally-regenerated mixtures of common North American boreal tree species were more productive and stored more C than single-species stands. We also examined how closely the different C pools and fluxes were interrelated and whether these relationships varied with species composition. Single- and mixed-species stands of trembling aspen, black spruce and jack pine on mesic sites were selected in two regions of the Canadian boreal forest to assess aboveground and belowground productivity and C storage. Although previous studies conducted in these stands found synergistic effects of tree species mixtures on specific C pools and fluxes, such as higher organic layer C stocks and higher fine root productivity in some mixtures, no effects were detected on combined C pools or fluxes at the ecosystem level in the current study. Aspen abundance was linked with higher aboveground tree productivity, higher aboveground living biomass and higher soil heterotrophic respiration, indicating that aspen acts as a key driver of ecosystem C storage and fluxes in these natural forest ecosystems, more so than species richness. However, our results do not rule out the possibility of increased productivity and C storage in mixed stands under environmental conditions or stand developmental stages that are different from the ones studied here. Furthermore, when the entire forest ecosystem is considered (not only tree parts), synergistic effects of tree species mixtures may be more difficult to observe because the beneficial effect of species mixing on one specific C pool may be counterbalanced by a negative effect on another pool.

Mixedwood forests occupy a large extent of boreal regions and have the potential for sequestering large amounts of carbon. In the context of forest ecosystem management, partial cutting prescriptions are increasingly being applied to boreal mixedwood stands. Partial harvesting is expected to maintain carbon pools and dynamics within the limits of those of natural stands. Changes in live tree, deadwood (standing snags, downed logs), forest floor and mineral soil carbon pools were assessed over a 9-year period in a replicated large-scale experiment, which included unharvested controls, two variants of partial harvesting and clear-cuts. We also measured leaf litter and deadwood inputs and decay rates. Carbon flux through leaf litterfall recovered rapidly following partial harvesting. Carbon flux from live trees to deadwood pools was a dominant process in partially harvested stands where snags and downed log carbon pools remained similar to those of natural stands. Hence, the nature of litter inputs diverged strongly among clear-cut and partially harvested treatments. Leaf and wood decay rates were higher in the partial cuts and controls than in clear-cuts. No significant differences in forest floor and mineral soil carbon were observed 9 years after harvesting. Carbon sequestration in live tree biomass was the carbon pool that most strongly differentiated the treatments allowing partial harvesting to maintain forest stands as net carbon sinks.

Climate change is altering insect disturbance regimes via temperature-mediated phenological changes and trophic interactions among host trees, herbivorous insects and their natural enemies in boreal forests. Range expansion and increase in outbreak severity of forest insects are occurring in Europe and North America. The degree to which northern forest ecosystems are resilient to novel disturbance regimes will have direct consequences on the provisioning of goods and services from these forests and on long-term forest management planning. Among major ecological disturbance agents in the boreal forests of North America is a tortricid moth, the eastern spruce budworm, which defoliates fir (Abies spp.) and spruce (Picea spp.). Northern expansion of this defoliator in eastern North America and climate-induced narrowing of the phenological mismatch between the insect and its secondary host, black spruce (Picea mariana), may permit greater defoliation and mortality in extensive northern black spruce forests. While spruce budworm outbreak centres have appeared in the boreal black spruce zone historically, defoliation and mortality were minor. Potential increases in outbreak severity and tree mortality raise concerns about the future state of this northern ecosystem. Severe spruce budworm outbreaks could decrease stand productivity compared with their occurrence in more diverse, southern balsam fir forest landscapes that have coevolved with outbreaks. Furthermore, depending on the proportion of balsam fir and deciduous species present and fire recurrence, changes in regeneration patterns and in nutrient cycling could alter ecosystem dynamics and replace black spruce by more productive mixed-wood forest, or by less productive ericaceous shrublands. Long-term monitoring, manipulative experiments and process modeling of climate-induced phenological changes on herbivorous insect pests, their host tree species and natural enemies in northern forests are therefore crucial to predicting species range shifts and assessing ecological and economic impacts.

We quantified the effects of different loads of forest logging residues on the microenvironment (soil temperature, soil volumetric water content, competing vegetation cover, and available nutrients) of planted hybrid poplars one year after planting, and assessed the corresponding seedling growth until the third growing season. In four experimental plantations across Quebec (Canada), we used a factorial design of four residue loads that were applied at the tree-level over three planted species: hybrid poplars ( Populus spp.), black spruce ( Picea mariana (Mill.) BSP), and either jack pine ( Pinus banksiana Lamb.) or white spruce ( Picea glauca (Moench) Voss), depending upon the site. Logging residues linearly decreased competing vegetation cover on two of four sites and reduced fluctuations in soil temperature on all sites. Logging residues also decreased summer soil temperatures at all sites through a negative quadratic effect. On one site, the frequency of freeze-thaw cycles increased under logging residues, while logging residues increased soil water content on another site, for certain measurement events. Logging residues did not affect available nutrients. Seedlings showed no consistent growth response to logging residues for three years after planting, except for a beneficial effect of logging residues on hybrid poplar growth on one site. Because logging residues affected seedling microclimate and competing vegetation, their maintenance and on-site spatial arrangement on site could be used to manipulate the growing conditions for planted trees.

Experimental research on beetle responses to removal of logging residues following clearcut harvesting in the boreal balsam fir forest of Quebec revealed several abundant rove beetle (Staphylinidae) species potentially important for long-term monitoring. To understand the trophic affiliations of these species in forest ecosystems, it was necessary to analyze their gut contents. We used microscopic and molecular (DNA) methods to identify the gut contents of the following rove beetles: Atheta capsularis Klimaszewski, Atheta klagesi Bernhauer, Oxypoda grandipennis (Casey), Bryophacis smetanai Campbell, Ischnosoma longicorne (Mäklin), Mycetoporus montanus Luze, Tachinus frigidus Erichson, Tachinus fumipennis (Say), Tachinus quebecensis Robert, and Pseudopsis subulata Herman. We found no apparent arthropod fragments within the guts; however, a number of fungi were identified by DNA sequences, including filamentous fungi and budding yeasts [Ascomycota: Candida derodonti Suh & Blackwell (accession number FJ623605), Candida mesenterica (Geiger) Diddens & Lodder (accession number FM178362), Candida railenensis Ramirez and Gonzáles (accession number JX455763), Candida sophie-reginae Ramirez & González (accession number HQ652073), Candida sp. (accession number AY498864), Pichia delftensis Beech (accession number AY923246), Pichia membranifaciens Hansen (accession number JQ26345), Pichia misumaiensis Y. Sasaki and Tak. Yoshida ex Kurtzman 2000 (accession number U73581), Pichia sp. (accession number AM261630), Cladosporium sp. (accession number KF367501), Acremoniumpsammosporum W. Gams (accession number GU566287), Alternaria sp. (accession number GU584946), Aspergillus versicolor Bubak (accession number AJ937750), and Aspergillusamstelodami (L. Mangin) Thom and Church (accession number HQ728257)]. In addition, two species of bacteria [Bradyrhizobium japonicum (Kirchner) Jordan (accession number BA000040) and Serratia marcescens Bizio accession number CP003942] were found in the guts. These results not only provide evidence of the consumer-resource relations of these beetles but also clarify the relationship between rove beetles, woody debris and fungi. Predominance of yeast-feeding by abundant rove beetles suggests that it may play an important role in their dietary requirements.

Increased interest in biomass harvesting for bioenergetic applications has raised questions regarding the potential ecological consequences on forest biodiversity. Here we evaluate the initial changes in the abundance, species richness and community composition of rove (Staphylinidae) and ground beetles (Carabidae), immediately following 1) stem-only harvesting (SOH), in which logging debris (i.e., tree tops and branches) are retained on site, and 2) whole-tree harvesting (WTH), in which stems, tops and branches are removed in mature balsam fir stands in Quebec, Canada. Beetles were collected throughout the summer of 2011, one year following harvesting, using pitfall traps. Overall catch rates were greater in uncut forest (Control) than either stem-only or whole-tree harvested sites. Catch rates in WTH were greater than SOH sites. Uncut stands were characterized primarily by five species: Atheta capsularis, Atheta klagesi, Atheta strigosula, Tachinus fumipennis/frigidus complex (Staphylinidae) and to a lesser extent to Pterostichus punctatissimus(Carabidae). Increased catch rates in WTH sites, where post-harvest biomass was less, were attributable to increased catches of rove beetles Pseudopsis subulata, Quedius labradorensis and to a lesser extent Gabrius brevipennis. We were able to characterize differences in beetle assemblages between harvested and non-harvested plots as well as differences between whole tree (WTH) and stem only (SOH) harvested sites where logging residues had been removed or left following harvest. However, the overall assemblage response was largely a recapitulation of the responses of several abundant species.

Sustainable management of boreal ecosystems involves the establishment of vigorous tree regeneration after harvest. However, two groups of understory plants influence regeneration success in eastern boreal Canada. Ericaceous shrubs are recognized to rapidly dominate susceptible boreal sites after harvest. Such dominance reduces recruitment and causes stagnant conifer growth, lasting decades on some sites. Additionally, peat accumulation due to Sphagnum growth after harvest forces the roots of regenerating conifers out of the relatively nutrient rich and warm mineral soil into the relatively nutrient poor and cool organic layer, with drastic effects on growth. Shifts from once productive black spruce forests to ericaceous heaths or paludified forests affect forest productivity and biodiversity. Under natural disturbance dynamics, fires severe enough to substantially reduce the organic layer thickness and affect ground cover species are required to establish a productive regeneration layer on such sites. We succinctly review how understory vegetation influences black spruce ecosystem dynamics in eastern boreal Canada, and present a multi-scale research model to understand, limit the loss and restore productive and diverse ecosystems in this region. Our model integrates knowledge of plant-level mechanisms in the development of silvicultural tools to sustain productivity. Fundamental knowledge is integrated at stand, landscape, regional and provincial levels to understand the distribution and dynamics of ericaceous shrubs and paludification processes and to support tactical and strategic forest management. The model can be adapted and applied to other natural resource management problems, in other biomes.

Changes in forest composition as a result of forest management, natural disturbances, and climate change may affect the accumulation of soil organic carbon (SOC). We examined the influence of common boreal tree species (trembling aspen, black spruce, and jack pine), either in pure stands or in conifer-broadleaf mixtures, on the amount, distribution, and quality of SOC in two regions of the Canadian boreal biome. Long-term laboratory incubations were used to assess SOC quality by quantifying proportions of fast carbon (C) (that is, proportion of total C released during the first 100 days of incubation) and active C (that is, modeled proportion of total C that can be potentially released). Total amounts of SOC did not differ between stand types, but the effects of stand type on SOC stocks and quality differed with soil depth. Among stand types, aspen stands had the greatest relative proportion of total SOC in deeper mineral layers and the lowest amount of active C in the organic layer. For these reasons, the SOC stock that developed under aspen was more stable than in the other stand types. Although black spruce stands allowed a greater accumulation of SOC in surface layers, these stocks, however, might become more vulnerable to extra losses if environmental conditions are to become more favorable to decomposition in the future. Our work highlights that boreal forest composition influences the stability of SOC stocks and how climate change could alter this large C pool.

• Although fine roots (< 2 mm in diameter) account for a major share of the production of terrestrial ecosystems, diversity effects on fine root productivity and their mechanisms remain unclear.
• We hypothesized that: (i) fine root productivity increases with tree species diversity, (ii) higher fine root productivity is a result of greater soil volume filling due to species-specific patterns of root placement and proliferation, and (iii) differences in fine root productivity and soil volume filling associated with tree species diversity are more pronounced in summer when plants are physiologically active and demand for water and nutrients is at its greatest.
• We investigated the effects of tree species diversity on fine root productivity and soil volume filling of boreal forest stands that have grown naturally for 85 years on similar sites.
• Annual fine root production was 19–83% higher in evenly mixed- than single-species-dominated stands, and increased with tree species evenness, but not tree species richness. Fine root biomass was higher in evenly mixed- than single-species-dominated stands in summer months, but not in spring or fall. Higher fine root productivity in evenly mixed- than single-species-dominated stands was realized by filling more soil volume horizontally and vertically in the forest floor in the mixtures of deep- and shallow-rooted species vs. the deeper mineral soil in the mixtures of deep-rooted species.
• Synthesis. Our results provide some of the first direct evidence for below-ground species complementarity in heterogeneous natural forests, by demonstrating that tree species evenness increases fine root productivity by filling/exploiting the soil environment more completely in space and time, driven by differences in the inherent rooting traits of the component species and variations of root growth within species.

Question: The effect of overstorey composition on above?ground dynamics of understorey vegetation is poorly understood. This study examines the understorey biomass, production and turnover rates of vascular and non?vascular plants along a conifer–broadleaf gradient of resource availability and heterogeneity.

Location: Canadian boreal forests of northwest Quebec and Ontario.

Methods: We sampled mature stands containing various proportions of black spruce (Picea mariana (Mill.) BSP), trembling aspen (Populus tremuloides Michx.) and jack pine (Pinus banksiana Lamb.). Above?ground biomass of the understorey vegetation was assessed through harvesting; annual growth rates were calculated as the differences between biomass in 2007 and 2008, as estimated by allometric relationships, and turnover rates were estimated as net primary production divided by the biomass in 2007.

Results: Higher aspen presence, linked to greater nutrient availability in the forest floor, was generally associated with higher vascular biomass and production in the understorey. This effect was less pronounced in sites of high intrinsic fertility. In contrast, bryophyte biomass was positively associated with conifer abundance, particularly in wet sites of the Quebec study area. Non?linear responses resulted in total understorey biomass being lower under mixed canopies than under pure aspen or pure conifer canopies. Turnover rates did not differ with overstorey composition.

Conclusions: While resource availability is a main driver of understorey productivity, resources as drivers appear to differ with differences in understorey strata components, i.e. vascular versus non?vascular plants. Resource heterogeneity induced by a mixed canopy had overall negative effects on understorey above?ground productivity, as this productivity seemed to rely on species adapted to the specific conditions induced by a pure canopy.

Processes governing tree interspecific interactions, such as facilitation and competition, may vary in strength over time. This study tried to unveil them by performing dendrometrical analyses on black spruce Picea mariana, trembling aspen Populus tremuloides and jack pine Pinus banksiana trees from pure and mixed mature boreal forest stands in the Clay Belt of northwestern Quebec and on the tills of northwestern Ontario. We cored 1430 trees and cut 120 for stem analysis across all stand composition types, tree species and study regions. Aspen annual growth rate was initially higher when mixed with conifers, but then progressively decreased over time compared to pure aspen stands, while jack pine growth rate did not differ with black spruce presence throughout all stages of stand development. When mixed with aspen, black spruce showed a contrary response to aspen, i.e. an initial loss in growth but a positive gain later. On the richer clay soil of the Quebec Clay Belt region, however, both aspen and spruce responses in mixed stands reversed between 37 and 54 years. Overall, our results demonstrate that interspecific interactions were present and tended to change with stand development and among species. Our results also suggest that the nature of interspecific interactions may differ with soil nutrient availability. © 2011 The Authors.

In boreal forested peatlands, we disturbed Sphagnum spp. and Pleurozium schreberi carpets to see how disturbance influenced substrate physico-chemistry, and growth and foliar nutrition of planted Picea mariana seedlings. Carpets were hand disturbed using gardening tools to a depth of approximately 25 cm. Carpet disturbance was aimed at disrupting only the organic layer and did not result in the mixing of organic matter with mineral soil. Disturbed carpets, whether P. schreberi or Sphagnum spp., were warmer than undisturbed carpets and had a lower cover of ericaceous shrubs. Pleurozium schreberi carpets had a higher decomposition index than Sphagnum spp. carpets, whereas disturbance had no effect on this variable. Pleurozium schreberi had higher N_tot and dissolved organic N concentrations (DON) than Sphagnum spp., whereas disturbance increased NH ₄ ⁺ availability in both substrates. Moss disruption increased seedling growth rates as well as their foliar N and P concentrations in both substrates and these variables remained higher in P. schreberi than in Sphagnum spp. within a given treatment. Seedling growth was positively correlated to substrate N_tot, NH ₄ ⁺ and DON concentrations, and to foliar N and P concentrations, and negatively to substrate C/N and ericaceous shrub cover. Disruption of the moss carpets without mineral soil mixing improved black spruce seedling growth and nutrition in both moss types but the superiority of P. schreberi compared to Sphagnum spp. as a growing substrate remained present.

Context

Following forest harvest, mechanical site preparation (MSP) is commonly used to regenerate harvested sites. In boreal forested peatlands, however, the effectiveness of MSP to regenerate harvested sites is likely to be hampered by thick organic layers.

Aim

We sought to determine the capability of different MSP techniques to improve growth conditions of planted black spruce seedlings in boreal forested peatlands where closed-crown productive forests could revert to unproductive forested peatlands in the absence of severe soil disturbance.

Methods

The effects of disc scarification, mounding and patch scarification on soil chemistry and seedling growth were contrasted.

Results

Seedlings of site-prepared plots were 15% taller than those of untreated ones, irrespective of the MSP technique used, likely owing to the greater abundance of exposed mineral soil and mesic substrates created. Mounding and patch scarification were able to expose mineral soil over a greater proportion (>25% vs. <10%) of the treated area compared with disc scarification and control, whereas the combined surface area of exposed mineral soil and mesic substrates was higher in every MSP treatments relative to the control (>57% vs. 41%, respectively). Individual seedling growth was influenced by substrate type and drainage. Seedlings planted in moderately and well-drained mesic substrates and mineral soil were 25% taller than those planted in poorly drained fibric substrates.

Conclusion

All three MSP techniques were effective because they succeeded in creating high-quality microsites despite thick organic layers.

As part of a multidisciplinary project on carbon (C) dynamics of the ecosystems characterizing the Eastmain Region Watershed (James Bay, Quebec), the objective of this study is to compare the soil C stocks and soil organic matter quality among the main upland vegetation types in a boreal region subjected to a high fire frequency. On average, the organic layer contained twice the amount of C than the mineral soil. Closed canopy vegetation types had greater C stocks both in the mineral and in the organic layers than the other more open canopy vegetation types. Landscape features such as drainage and surficial deposit could not discriminate between vegetation types although closed vegetation types were on average found on wetter site conditions. Average soil C contents varied more than 2-fold across vegetation types. On the opposite, except for the organic layer C:N ratio, which was smaller in closed vegetation types, other measured soil organic matter properties (namely specific rate of evolved C after a long-term incubation, hydrolysis acid-resistant C as well as the rate of changes in soil heterotrophic respiration with increasing temperature (Q10)) remained within a narrow range between vegetation types. Therefore, total soil C stocks were a major determinant of both labile C and estimated summer soil heterotrophic respiration rate. The homogeneity of soil organic matter quality across the vegetation types could be attributable to the positive relationship between soil C storage and soil C fluxes observed in this landscape experiencing a high fire frequency. The low variability in soil C quality could help simplify the modelling of soil C fluxes in this environment.

It has been suggested that without sufficient soil disturbance, harvest in boreal forested peatlands may accelerate paludification and reduce forest productivity. The objectives of this study were to compare the effects of harvest methods (clearcutting vs. careful logging) and season (summer vs. winter harvest) on black spruce regeneration and growth in boreal forested peatlands of eastern Canada, and to identify the soil variables that favour tree growth following harvest. Moreover, we sought to determine how stand growth following harvest compared with that observed following fire. The average tree height of summer clearcuts was greater than that of summer carefully logged stands and that of all winter harvested sites. Summer clearcutting also resulted in a higher density of trees > 3 m and > 4 m tall and in a 50% reduction in Rhododendron groenlandicum cover, a species associated with reduced black spruce growth. Height growth of sample trees was related to foliar N and P concentrations, and to soil total N, pH and available Ca and Mg but not to harvest method or season. Our results also indicate that summer clearcutting could produce stand productivity levels comparable to those observed after high-severity soil burns. These results suggest that summer clearcutting could be used to restore forest productivity following harvest in forested peatlands, and offer further support to the idea that sufficient levels of soil disturbance may be required to restore productivity in ecosystems undergoing paludification.

The slow decomposition rate of boreal forest floor bryophytes contributes both to maintaining high soil C reserves as well as affecting conditions for tree growth by maintaining excessively high soil water content, cooling the soil and slowing nutrient cycles. In this study, mass loss of three bryophyte species (Pleurozium schreberi, Sphagnum capillifolium, S. fuscum) was measured in unharvested, partial cut and low-retention cut forest blocks. Mesh decomposition bags containing the three species and wood sticks were placed at two depths in colonies of either P. schreberi or S. capillifolium (environment) in the three harvest treatments and retrieved after two growing seasons. Mass loss was primarily related to substrate type (P. schreberi > S. capillifolium > wood sticks > S. fuscum) and secondarily to depth. Harvest treatment and environment (P. schreberi or S. capillifolium) only weakly affected sphagna mass loss. The weak effect of harvest treatment suggests that conditions created by low retention cuts do not to stimulate decomposition in this system and are not important enough to stimulate carbon loss, or to counteract paludification. On the other hand, the strong effect of bryophyte type indicates that conditions affecting bryophyte colonization and succession are of great importance in driving carbon and nutrient cycles.

Summary

1. The diversity–productivity debate has so far been focused above-ground, despite that below-ground production can account for approximately half of total annual net primary production, mostly from fine roots.

2. Here, we investigate the fine root productivity of mature, fire-origin stands of Populus tremuloides–Picea spp.–Abies balsamea (mixed-species stands) and relatively pure P. tremuloides (single-species stands) in two regions of North American boreal forest to better understand the link between plant diversity and below-ground productivity in forest ecosystems. We hypothesized that: (i) mixed-species stands have higher fine root productivity compared with single-species stands and (ii) this difference may be the result of greater soil space filling by the fine roots due to the contrasting rooting traits of the component species in the mixed-species stands.

3. We found that fine root productivity, measured by annual production and total biomass, was higher in mixed- than single-species stands. We also found that mixed-species stands had lower and higher horizontal and vertical fine root biomass heterogeneity, respectively, indicating that soil space is more fully occupied by fine roots in the mixed- than single-species stands.

4. In all, our study supports that below-ground niche differentiation may be a key driver of higher fine root productivity in mixed stands of species with contrasting rooting traits than single-species stands by facilitating greater soil space filling of fine roots and soil resource exploitation.

Abstract: Machinery traffic restrictions during forest harvest have been adopted to minimize soil damage and protect tree regeneration. However, this practice is questioned for paludifying black spruce (Picea mariana (Mill.) BSP) stands in which severe soil disturbance by wildfire restores forest productivity. The objective of this study was to determine, 8 years after harvest, how soil disturbance created by clearcutting and careful logging affected black spruce natural regeneration and growth and how this effect varied by soil type. While regeneration density was higher following careful logging, stocking was not influenced by harvest method. Regenerating stands were taller following clearcutting despite potentially greater damages to preestablished regeneration. Compared with careful logging, clearcutting also resulted in reduced cover of Sphagnum spp. and ericaceous shrubs. Spruce stem density and stocking were both higher on organic and subhydric soils and lower on mesic soils. No significant interactions were observed between harvest method and soil type, indicating that the observation of taller black spruce stands and adequate stocking with clearcutting may be applicable to all soil types considered in this study. These results suggest that an adequate level of soil disturbance is an important part of forest regeneration, particularly in ecosystems where an autogenic reduction in productivity occurs.

Résumé : Afin de préserver les sols et la régénération durant la récolte, la circulation de la machinerie forestière est fréquemment restreinte à des sentiers définis. Toutefois, cette pratique est remise en question dans les peuplements d’épinette noire (Picea mariana (Mill.) BSP) susceptibles à la paludification où de sévères perturbations des sols causées par le feu sont aptes à améliorer la productivité des peuplements. L’objectif de cette étude était de déterminer comment la perturbation des sols causée lors de la coupe totale et la coupe de protection affectent la régénération de l’épinette noire 8 ans après la récolte, et comment cet effet est conditionné par le type de sol. Alors que la densité de régénération de l’épinette noire était supérieure après la coupe de protection, le coefficient de distribution n’était pas influencé par la méthode de récolte. Les peuplements d’épinette régénérés après coupe totale étaient plus hauts en dépit d’un potentiel de dommage plus grand à la régénération préétablie. Comparée à la coupe de protection, la coupe totale a aussi résulté en une réduction de la couverture au sol de la sphaigne et des éricacées. La densité et le coefficient de distribution de l’épinette noire étaient supérieurs sur les sols organiques et subhydriques et inférieurs sur les sols mésiques. Aucune interaction significative n’a été observée entre la méthode de récolte et le type de sol, indiquant qu’une plus grande hauteur des peuplements et un coefficient de distribution d’épinette adéquat après coupe totale pourrait être applicable à toute la gamme de types de sol considérée dans cette étude. Ces résultats suggèrent qu’un niveau de perturbation des sols suffisant lors des opérations forestières est nécessaire, particulièrement dans les écosystèmes où des processus autogéniques réduisant la productivité s’opèrent.

Abstract: Plant species distribution and plant community composition vary along environmental gradients. At the continental scale, climate plays a major role in determining plant distribution, while at the local and regional scales vegetation patterns are more strongly related to edaphic and topographic factors. The projected global warming and alteration of the precipitation regime will influence tree physiology and phenology, and is likely to promote northward migration of tree species. However the influence of soil characteristics on tree species migration is not as well understood. Considering the broad tolerance of most tree species to variations in soil factors, soils should not represent a major constraint for the northward shift of tree species. However, locally or regionally, soil properties may constrain species migration. Thus, while climate change has the potential to induce a northward migration of tree species, local or regional soil properties may hinder their migratory response. These antagonistic forces are likely to slow down potential tree migration in response to climate change. Because tree species respond individualistically to climate variables and soil properties, new tree communities are likely to emerge from climate change.

Résumé : La distribution des espèces et la composition des communautés végétales varient le long de gradients environnementaux. À l’échelle continentale, le climat joue un rôle primordial dans la distribution des espèces végétales, alors qu’aux échelles locale et régionale la distribution des espèces est surtout influencée par les variables édaphiques et la topographie. Le réchauffement climatique projeté et l’altération du régime des précipitations sont susceptibles d’influencer la physiologie et la phénologie des arbres, et d’entraîner leur migration vers le nord. Toutefois, l’influence des caractéristiques des sols sur le potentiel migratoire des arbres est peu connue. Puisque la plupart des espèces d’arbres possèdent une grande tolérance aux variations édaphiques, les sols ne devraient pas constituer un obstacle majeur à leur migration vers le nord. Toutefois, localement ou régionalement, les sols pourraient posséder des propriétés qui pourraient restreindre la migration des espèces. Alors que les changements climatiques ont le potentiel de provoquer la migration des espèces vers le nord, localement ou régionalement les propriétés des sols pourraient limiter leur capacité migratoire. Ces forces en opposition pourraient restreindre le potentiel migratoire des arbres en réponse aux changements climatiques. Parce que les espèces d’arbres répondent de manière individuelle aux variables climatiques et aux propriétés des sols, de nouvelles communautés d’arbres pourraient résulter des changements climatiques.

Abstract:

A renewed interest in the intensive harvesting of forest biomass as a source of bioenergy in North America raises concerns about the impacts that this practice may have on the maintenance of forest soil productivity. In Canada, such concerns were first voiced in the 1970s, and studies were launched to investigate and predict the impact of intensive forest biomass removal on site productivity. Most of these studies focused on static nutrient budgets. In Canada and around the world, more detailed process models were also developed to study carbon, nitrogen and base cation cycles under different forest harvesting intensities. However, the validity of modelling results is still constrained by our lack of knowledge on the capacity of ecosystems to supply nutrients. A few sets of field trials have been established in Canada to gather empirical data on the impact of biomass removal on soil nutrient reserves as well as on tree nutrition and growth. Although still fairly recent, these field trials, along with the older ones established in other countries with similar site conditions and climates, provide opportunities to refine our understanding of the resilience of ecosystem processes and of the impacts of intensive biomass removal on ecosystem functions. Although numerous knowledge gaps and questions remain, some jurisdictions around the world have nevertheless issued policy directives and developed guidelines for biomass harvesting. As described by the concept of adaptive forest management, ecological monitoring of harvesting operations, scientific field testing and modelling can all interact to produce better knowledge that could then help improve policy directives.

Résumé:

L’intérêt renouvelé pour la récolte intensive de biomasse forestière comme source de bioénergie en Amérique du Nord soulève des inquiétudes quant aux impacts de cette pratique sur le maintien de la productivité des sols. Au Canada, ces inquiétudes ont été émises pour la première fois dans les années 1970 et des travaux de recherche ont été entrepris pour étudier et prédire les effets de la récolte de biomasse sur la productivité des sites. La plupart de ces études étaient basées sur des budgets nutritionnels statiques. Au Canada et ailleurs dans le monde, des modèles plus détaillés incluant la description des processus écologiques ont aussi été développés pour étudier les cycles du carbone, de l’azote et des cations basiques sous différentes intensités de récolte. Cependant, la validité des résultats de modélisation est toujours contrainte par notre connaissance limitée de la capacité des écosystèmes à fournir des éléments nutritifs. Quelques dispositifs expérimentaux ont été établis au Canada pour recueillir des données empiriques sur l’impact de la récolte de biomasse sur les réserves nutritionnelles du sol ainsi que sur la croissance et la nutrition des arbres. Bien que relativement récents, ces dispositifs, ainsi que des dispositifs plus anciens installés dans d’autres pays aux conditions de site et de climat semblables à celles du Canada, nous offrent l’opportunité de raffiner notre compréhension de la résilience des processus écologiques et des impacts de la récolte de biomasse sur le fonctionnement des écosystèmes. Bien que les incertitudes et les questions restent nombreuses, certaines juridictions dans le monde ont tout de même fourni un cadre politique et développé des lignes directrices pour la récolte de biomasse. Tel que décrit par le concept d’aménagement forestier adaptatif, le suivi écologique des opérations de récolte, les études sur le terrain et la modélisation peuvent interagir afin d’améliorer les connaissances scientifiques qui pourront par la suite être utilisées de façon à ajuster le cadre politique.

Abstract: Litter quality is often considered the main driver of decomposition rate. The objective of this study was to investigate the relative contribution of two other tree-driven mechanisms, litter mixing and forest floor conditions, to foliar litter decomposition and nutrient dynamics for trembling aspen (Populus tremuloides Michx.) and black spruce (Picea mariana (Mill.) BSP) using a microcosm approach. Results based on mixed linear models show that the greater influence over these processes was obtained through litter quality followed by forest floor conditions and litter mixing. Specifically, the results indicate that significantly more C and nutrients were mineralized (i) from aspen than from spruce litter, (ii) from spruce litter in mixture with aspen litter than from spruce litter applied singly, and (iii) from litter incubated on forest floor from the aspen stand rather than from the spruce stand, except for nutrients in the spruce litter. Collectively, our results show that the litter and forest floor material from aspen both favour decomposition and nutrient mineralization processes. Hence, we provide evidence that the effect of tree species on litter decomposition may not only be caused by the properties of its litter but also, indirectly, by the specific conditions and the decomposer community that tree species develop in their forest floor.

Résumé : La qualité de la litière est souvent considérée comme le facteur déterminant de la décomposition. L'objectif de cette étude était d'étudier la contribution relative de deux autres mécanismes contrôlés par l'arbre, soit le mélange de litière et les propriétés de la couverture morte, à la décomposition et à la dynamique des nutriments de la litière foliaire du peuplier faux-tremble (Populus tremuloides Michx.) et de l'épinette noire (Picea mariana (Mill.) BSP) en utilisant une approche en microcosme. Les résultats basés sur des modèles mixtes linéaires montrent que l'influence la plus grande sur ces processus a été obtenue via la qualité de la litière, suivie des propriétés de la couverture morte et de mélange de litière. Spécifiquement, les résultats indiquent que significativement plus de C et de nutriments ont été minéralisés (i) dans la litière de peuplier que dans la litière d'épinette, (ii) dans la litière d'épinette en mélange avec la litière de peuplier que dans la litière d'épinette seule et (iii) dans les litières incubées sur la couverture morte provenant du peuplement de peuplier que sur celle provenant du peuplement d'épinette, excepté pour les nutriments de la litière d'épinette. Collectivement, nos résultats montrent que la litière et la couverture morte de peuplier favorisent les processus de décomposition et de minéralisation des nutriments. Ainsi, cette étude prouve que l'influence des essences forestières sur la décomposition de la litière est non seulement reliée aux propriétés de leur litière, mais aussi, indirectement, aux conditions spécifiques et à la communauté de décomposeurs que les essences forestières développent dans leur couverture morte.

Abstract: This study investigates the potential of mixed forest stands as better aboveground carbon sinks than pure stands. According to the facilitation and niche complementarity hypotheses, we predict higher carbon sequestration in mature boreal mixedwoods. Aboveground carbon contents of black spruce (Picea mariana (Mill.) Britton, Sterns, Poggenb.) and trembling aspen (Populus tremuloides Michx.) mixtures were investigated in the eastern boreal forest, whereas jack pine (Pinus banksiana Lamb.) and trembling aspen were used in the central boreal forest. No carbon gain was found in species mixtures; nearly pure trembling aspen stands contained the greatest amount of aboveground carbon, black spruce stands had the least, and mixtures were intermediate with amounts that could generally be predicted by linear interpolation with stem proportions. These results suggest that for aspen, the potentially detrimental effect of spruce on soils observed in other studies may be offset by greater light availability in mixtures. On the other hand, for black spruce, the potentially beneficial effects of aspen on soils could be offset by greater competition by aspen for nutrients and light. The mixture of jack pine and trembling aspen did not benefit any of these species while inducing a loss in trembling aspen carbon at the stand level.

Résumé : Cette étude vise à déterminer si les peuplements forestiers mixtes peuvent être des puits de carbone aériens plus efficaces que les peuplements purs. En accord avec les hypothèses de facilitation et de séparation des niches écologiques, nous prédisons une plus grande séquestration du carbone dans les peuplements mixtes. Nous avons donc déterminé les stocks de carbone aériens dans des mélanges d’épinettes noires (Picea mariana (Mill.) Britton, Sterns, Poggenb.) et de peupliers faux-tremble (Populus tremuloides Michx.) situés dans la forêt boréale de l’est, tandis que des mélanges de pin gris (Pinus banksiana Lamb.) et de peupliers faux-tremble furent plutôt utilisés dans la forêt boréale centrale. Les mélanges d’espèces ne présentaient aucun gain en carbone. Les peuplements dominés par le tremble contenaient la plus grande quantité de carbone aérien, les pessières la plus petite quantité, et les mélanges en contenaient des quantités intermédiaires qui pouvaient généralement être prédites par interpolation linéaire en fonction de la proportion de tiges de chaque espèce. Ces résultats suggèrent que dans le cas du tremble, l’effet potentiellement néfaste de l’épinette sur les sols peut être compensé par une meilleure disponibilité de la lumière dans les peuplements mixtes. Parallèlement, dans le cas de l’épinette noire, l’effet potentiellement bénéfique du peuplier faux-tremble sur les sols pourrait être contrebalancé par une compétition accrue pour la lumière et les nutriments en sa présence. Le mélange de pin gris et de peuplier faux-tremble n’a apporté aucun bénéfice pour aucune des deux espèces tout en induisant une perte en carbone chez le tremble à l’échelle du peuplement.

Deforestation usually results in significant losses of soil organic carbon (SOC). The rate and factors determining the recovery of this C pool with afforestation are still poorly understood. This paper provides a review of the influence of afforestation on SOC stocks based on a meta-analysis of 33 recent publications (totaling 120 sites and 189 observations), with the aim of determining the factors responsible for the restoration of SOC following afforestation. Based on a mixed linear model, the meta-analysis indicates that the main factors that contribute to restoring SOC stocks after afforestation are: previous land use, tree species planted, soil clay content, preplanting disturbance and, to a lesser extent, climatic zone. Specifically, this meta-analysis (1) indicates that the positive impact of afforestation on SOC stocks is more pronounced in cropland soils than in pastures or natural grasslands; (2) suggests that broadleaf tree species have a greater capacity to accumulate SOC than coniferous species; (3) underscores that afforestation using pine species does not result in a net loss of the whole soil-profile carbon stocks compared with initial values (agricultural soil) when the surface organic layer is included in the accounting; (4) demonstrates that clay-rich soils (> 33%) have a greater capacity to accumulate SOC than soils with a lower clay content (< 33%); (5) indicates that minimizing preplanting disturbances may increase the rate at which SOC stocks are replenished; and (6) suggests that afforestation carried out in the boreal climate zone results in small SOC losses compared with other climate zones, probably because trees grow more slowly under these conditions, although this does not rule out gains over time after the conversion. This study also highlights the importance of the methodological approach used when developing the sampling design, especially the inclusion of the organic layer in the accounting.

The occurrence of aspen (Populus tremuloides Michx.) patches within stands dominated by black spruce (Picea mariana Mill. BSP) has been shown to increase litter decomposition and nutrient cycling rates by improving soil physical and chemical properties. It is well known, however, that these processes are also influenced by the structure of the soil biota, but this factor has received less attention. In this study, relationships between forest floor properties and soil invertebrates were studied along black spruce-trembling aspen gradients in three stands of the eastern boreal forest of Canada. The forest floor layer of 36 plots differing in aspen basal area was sampled and analyzed to determine physical and chemical properties, the rates of decomposition of standard substrates, net N mineralization, as well as microbial basal respiration and metabolic quotient. Soil invertebrates were also collected using funnel-extraction and pitfall trapping methods. Based on redundancy analyses, we found that forest floor properties, the abundance and composition of soil invertebrates, and the rates of belowground processes changed along the spruce-aspen gradient. The increase in aspen basal area was associated with a reduction in forest floor thickness, moisture content and microbial biomass, and with an increase in the concentration of nutrients. It was also accompanied by changes in soil faunal communities, as soil invertebrates were associated with specific soil properties. In general, macroinvertebrates (i.e., Lumbricidae, Formicidae, Carabidae, Staphylinidae and Gastropoda) were related to the nutrient-rich forest floor associated with aspen, whereas microarthropods and Enchytraeidae tended to be negatively related to aspen basal area. According to mixed linear models, decomposition rates of standard substrates and net ammonification significantly increased along the spruce-aspen gradient. Given the functional significance of macroinvertebrates in soils, these results suggest that aspen favours the elaboration of a macrofaunal community, which in turn accelerates the rate of soil processes by having either direct or indirect influence on microbial activity. Moreover, this study shows that the changes in soil processes and in the biodiversity of soil organisms related to the presence of mixed stands can operate only in the immediate surroundings of a given tree species. Therefore, coarse-scale tree species mixing in a forest stand may have a different effect on soil biodiversity and soil processes than fine-scale mixing.

Abstract: In many northern forest ecosystems, soil organic matter accumulation can lead to paludification and forest productivity losses. Paludification rate is primarily influenced by topography and time elapsed since fire, two factors whose influence is often confounded and whose discrimination would help forest management. This study, which was conducted in the black spruce (Picea mariana (Mill.) BSP) boreal forest of northwestern Quebec (Canada), aimed to (1) quantify the effect of slope and time since fire on paludification rates, (2) determine whether soil organic layer depth could be estimated by surface variables that can potentially be remotely sensed, and (3) relate the degree of paludification to tree productivity. In this study, soil organic layer depth was used as an estimator of the degree of paludification. Slope and postfire age strongly affected paludification dynamics. Young stands growing on steep slopes had thinner organic layers and lower organic matter accumulation rates compared with young stands growing on flat sites. Black spruce basal area and Sphagnum cover were strong predictors of organic layer depth, potentially allowing mapping of paludification degree across the landscape. Tree productivity was negatively related to organic layer depth (R² = 0.57). The equations developed here can be used to quantify forest productivity decline in stands that are undergoing paludification, as well as potential productivity recovery given appropriate site preparation techniques.

Résumé : Dans plusieurs écosystèmes forestiers nordiques, l'accumulation de matière organique peut mener à la paludification des sols et entraîner des pertes de productivité forestière. Le taux de paludification est principalement influencé par la topographie et le temps écoulé depuis le dernier feu, deux facteurs dont l'influence est souvent confondue et dont la séparation aiderait à l'aménagement forestier. Cette étude réalisée dans la pessière noire (Picea mariana (Mill.) BSP) du nord-ouest du Québec (Canada) avait comme objectifs : (1) de quantifier l'effet de la pente et du temps depuis le dernier feu sur le taux de paludification, (2) de déterminer si l'épaisseur de la couche organique pouvait être estimée à partir de deux variables de surface quantifiables au moyen de la télédétection, et (3) d'établir la relation entre le degré de paludification et la productivité des arbres. Dans cette étude, l'épaisseur de la couche organique du sol a été utilisée pour estimer le degré de paludification. La pente et l'âge des peuplements après feu influencent fortement la dynamique de paludification. La couche organique était plus mince et avait un taux d'accumulation plus faible dans les jeunes peuplements établis sur des pentes fortes que dans les peuplements d'âge similaire établis sur terrain plat. La surface terrière en épinette noire et le recouvrement de sphaignes avaient un fort pouvoir de prédiction de l'épaisseur de la couche organique, ce qui pourrait permettre de cartographier le degré de paludification à l'échelle du paysage. Enfin, la productivité des arbres était négativement reliée à l'épaisseur de la couche organique (R² = 0,57). Les équations développées dans ce travail peuvent être utilisées pour quantifier le déclin de productivité forestière dans les peuplements sujets à la paludification, ainsi que le potentiel de récupération de productivité à la suite d'une préparation de terrain appropriée.

Long-term forest productivity decline in boreal forests has been extensively studied in the last decades, yet its causes are still unclear. Soil conditions associated with soil organic matter accumulation are thought to be responsible for site productivity decline. The objectives of this study were to determine if paludification of boreal soils resulted in reduced forest productivity, and to identify changes in the physical and chemical properties of soils associated with reduction in productivity. We used a chronosequence of 23 black spruce stands ranging in postfire age from 50 to 2350 years and calculated three different stand productivity indices, including site index. We assessed changes in forest productivity with time using two complementary approaches: (1) by comparing productivity among the chronosequence stands and (2) by comparing the productivity of successive cohorts of trees within the same stands to determine the influence of time independently of other site factors.

Charcoal stratigraphy indicates that the forest stands differ in their fire history and originated either from high- or low-severity soil burns. Both chronosequence and cohort approaches demonstrate declines in black spruce productivity of 50–80% with increased paludification, particularly during the first centuries after fire. Paludification alters bryophyte abundance and succession, increases soil moisture, reduces soil temperature and nutrient availability, and alters the vertical distribution of roots. Low-severity soil burns significantly accelerate rates of paludification and productivity decline compared with high-severity fires and ultimately reduce nutrient content in black spruce needles. The two combined approaches indicate that paludification can be driven by forest succession only, independently of site factors such as position on slope. This successional paludification contrasts with edaphic paludification, where topography and drainage primarily control the extent and rate of paludification. At the landscape scale, the fire regime (frequency and severity) controls paludification and forest productivity through its effect on soil organic layers. Implications for global carbon budgets and sustainable forestry are discussed. ©2007 ESA

The influence of forest stand composition on soil was investigated by comparing the forest floor (FH) and upper mineral soil (0–20 cm) nutritional properties of jack pine and aspen stands on two soil types of contrasting fertility, a coarse-textured and a fine-textured deposit, in a replicated design. The studied tree species are pioneers that are found after major disturbances in the southern boreal forest of western Quebec and that differ in their nutrient requirements but not in their growth rate. Soil organic matter as well as total and available N, P, K, Ca, Mg contents were determined and the relationships with nutrient accumulation in tree biomass were studied. On both soil types a greater total and available nutrient accumulation in the forest floor layer was observed in aspen than in jack pine whereas such differences between stand types could not be detected in the mineral soil. Differences in FH nutrient content between stand types were larger on coarse deposits than on fine-textured soils. These results support the hypothesis that tree species with greater nutrient requirements cause an enrichment of the surface soil at least in the short term. The modulation of tree species effect by soil type was contrary to the pattern observed in other studies since a greater expression of this effect was observed on poorer soils. Differences in soil nutrient content were related to levels of organic matter accumulation.© 2007 Ecosystems.

Abstract

The surface of the soil in recently harvested or burned lowland black spruce (Picea mariana (Mill.) BSP) sites is composed of a fine mosaic of different bryophytes (mostly Sphagnum spp. and feathermosses), disturbed organic material originating mostly from mosses at different stages of decay, and exposed mineral soil. Growth substrates were compared in lowland black spruce stands regenerating after either careful logging or wildfire. The 3-year annual increment for black spruce seedlings was greatest with substrates of feathermosses, mainly Pleurozium schreberi (Brid.) Mitt., fibric material of P. schreberi origin, and a mixture of fibric P. schreberi and humic materials; it was least with fibric Sphagnum spp., mineral soil, and decaying wood substrates. The most favourable substrates for growth were characterized by better black spruce N and P foliar status. Our results also suggest that categories of growth substrates in the rooting zone reflect nutritional quality better than categories of growth substrates on the soil surface. To maintain or increase black spruce growth following careful logging of sites prone to paludification, we recommend fill-planting of seedlings in substrates originating from P. schreberi; management techniques that favour P. schreberi over Sphagnum mosses should also be developed.

Résumé

La surface du sol de sites récemment récoltés ou brûlés dans des peuplements d'épinettes noires (Picea mariana (Mill.) BSP) de basses terres est composée d'une fine mosaïque de différents types de bryophytes, principalement les sphaignes et les mousses hypnacées, de matière organique perturbée composée de différents types de mousses à différents degrés de décomposition et de sol minéral exposé. Les substrats de croissance qu'on retrouve dans des peuplements d'épinettes noires de basses terres provenant de sites régénérés suite à la coupe avec protection de la régénération et des sols ou suite à un feu ont été comparés. Les résultats suggèrent que la croissance en hauteur des semis d'épinette noire sur 3 ans est plus élevée avec les substrats de Pleurozium schreberi (Brid.) Mitt., de matériel fibrique composé de P. schreberi et d'un mélange de matériels fibrique (composé de P. schreberi) et humique qu'avec la sphaigne fibrique, le sol minéral et le bois mort. Les substrats de croissance les plus favorables à la croissance sont caractérisés par une meilleure nutrition en azote et phosphore. Nos résultats suggèrent également que la classification des substrats de croissance au niveau des racines est plus indicative de leur valeur nutritive que la classification des substrats de croissance localisés en surface. En se basant sur ces résultats, pour maintenir ou augmenter la croissance en hauteur de l'épinette noire après coupe sur des sites susceptibles à la paludification, nous recommandons que la plantation dans des substrats formés de P. schreberi et qu'un aménagement qui favorise la présence de P. schreberi au détriment des sphaignes soient favorisés. ©2006 NRC Canada

Black spruce (Picea mariana (Mill.) BSP) is the most important commercial tree species in the eastern boreal forest of Canada. Only limited work has been conducted to assess the quality of the various substrates that are found on postdisturbed sites prone to paludification having an effect on black spruce seedling growth. The objectives of this study were (1) to use a pot test to assess black spruce seedling performance on substrates found at the soil surface, in the rooting zone of undisturbed soil, at depths that become available to spruce roots after soil disturbance by wildfire or through management; and (2) to determine the nutritional quality of these substrates when constraints of poor drainage are artificially removed. Black spruce growth was greatest with fibric Pleurozium in the rooting zone, and with burned fibric Pleurozium and living Sphagnum at the soil surface. Good seedling growth on different substrates found in the rooting zone was associated with higher N and P foliar concentrations. Based on these results, we recommend targeted planting of black spruce seedlings in substrates of Pleurozium origin and the development of management techniques that promote Pleurozium schreberi.

Abstract

Questions: 1. How does the spatial structure of the organic layer affect tree sapling physiology? 2. Are the organic layer and Picea mariana height growth spatially structured at different scales? 3. Does microtopography influence the accumulation of organic matter and does organic layer thickness affect height growth?

Locations: Picea mariana forests, northwestern Quebec, Canada.

Methods: We assessed the spatial pattern of each variable in one wildfire site and one harvest site using semivariograms and correlograms. We measured the cross-correlation between relative elevation and organic layer thickness, and between organic layer thickness and growth using cross-correlograms.

Results: Picea mariana height growth was autocorrelated to a greater extent in the wildfire site (103 m) than in the harvest site (43 m). The spatial structure of organic layer thickness was similar in both sites. Deeper depressions in the harvest site, as illustrated by spatial variance in relative elevation at short distances (ca. 50 m), and by high autocorrelation values, increased the accumulation of organic matter within 20 m.

Conclusions: The interaction between microtopography and organic matter accumulation led to paludification and poor growth of Picea mariana at the harvest site. Paludification at the wildfire site was independent of microtopography and was probably a result of stand development.

The effects of harvesting and slash treatments on soil nutrient dynamics were assessed in boreal aspen stands growing on mesic clayey sites. Stem-only harvested stands were compared with unharvested controls according to a complete block design with three replications. Within harvested areas, four slash treatments (stem-only harvesting [SOH], whole-tree harvesting [WHT], wood chip application, and slash burns) were compared. Treatments created a gradient of slash that ranged from 52.3 Mg ha–1 in stem-only to 13.8 Mg ha–1 in control stands. The amount of slash had no effect on wood decomposition rates but was strongly associated with higher forest floor organic C, Kjeldahl N and base cation concentrations (Cae and Mge), base saturation, pH, and effective cation exchange capacity (CEC) and lower microbial C/N when control stands were compared with stem-only harvested stands. Slash burn severity was too low to significantly reduce slash loads and induce base cation release but severe enough to reduce forest floor microbial C and N concentrations (–48 and –55%, respectively) for at least one complete growing season. Slash burn also induced increases in forest floor available P (54%) concentration compared with other slash treatments. Chipping reduced forest floor microbial N concentration by 25% and increased microbial C/N by 28% but had no impact on nutrient availability. Differences between WTH and SOH were linked to the abundance of slash. Finally, the results illustrate that whatever the treatment, the amount of slash left on the ground is the main factor found to affect soil microbial community characteristics and soil nutrient availability.

Abbreviations : CEC, effective cation exchange capacity • CHT, chipping harvesting treatment • PCA, principal component analysis • SB, slash-burn treatment • SOH, stem-only harvesting treatment • WTH, whole-tree harvesting treatment

Abstract

To assess nutrient dynamics in decomposing logs of trembling aspen (Populus tremuloides Michx.), white birch (Betula papyrifera Marsh.), white spruce (Picea glauca (Moench) Voss), and jack pine (Pinus banksiana Lamb.), we monitored mass losses and changes in N and P contents in dead boles from a chronosequence of sites following stand-replacing disturbances. To assess the importance of wood decomposition to nutrient cycling, we compared net estimates of nutrient release from logs with net nutrient immobilization in live-tree biomass of stands as a function of time since disturbance. Mineralization rates were 0.060, 0.053, 0.038, and 0.020 center dot year(-1) for trembling aspen, white birch, white spruce, and jack pine logs, respectively. Trembling aspen boles released large quantities of N and P during the first year of decomposition (51 kg center dot ha(-1) of N and 7 kg center dot ha(-1) of P, assuming a bole volume of 150 m(3)center ! dot ha(-1)). White birch boles acted initially as a nutrient sink and delayed the release of immobilized nutrients until a period when the stand ' s net nutrient immobilization rates were highest. Jack pine boles appeared to be intermediate in terms of their contribution as a sink or a source of nutrients but, in mature stands, provided up to 40% of N and 26% of P immobilized annually in tree biomass. As pure stands of white spruce are rare in boreal Quebec, information on nutrient accumulation in white spruce stands was not available.

Résumé

Après avoir échantillonné les débris ligneux (DL) et la matière organique du sol (MO) présents dans la couverture morte, les billes au sol et le sol minéral de surface (0–30 cm) dans 14 peuplements entre les stades intermédiaire et final de succession, établis sur une variété de types de sol et soumis à différents régimes climatiques dans le nord-ouest des États-Unis, nous avons trouvé que 44 % à 84 % du carbone (C) se retrouvait dans les DL et la MO en surface tandis que plus de 80 % de l'azote (N) se retrouvait dans le sol minéral. Dans plusieurs forêts du nord-ouest, la lutte contre les incendies de forêt et les changements naturels dans la composition des peuplements ont eu pour effet d'augmenter la quantité de DL et de MO qui peut être perdue à cause des incendies de forêt. Les peuplements qui ont des concentrations élevées de MO à la surface du sol ont plus de chances de perdre de grandes quantités de C et de N après un feu de surface de forte sévérité. Selon le guide de la qualité des sols du Service forestier américain, nous estimons que 6 % à 80 % du réservoir de C jusqu'à 30 cm dans le sol minéral pourrait être perdu lors d'un incendie considéré comme néfaste pour la productivité du sol. Ces estimations devraient varier selon le régime climatique local, la sévérité du feu dans la zone brûlée, la dimension et la classe de décomposition des DL et la répartition de la MO dans les portions organique et minérale du sol de surface. Les pertes estimées de N dues au feu étaient beaucoup plus faibles (< 1 % à 19 %). D'autres études sur la quantité et la répartition de la MO dans ces peuplements sont nécessaires pour évaluer le risque d'incendie de forêt, déterminer les impacts de différentes sévérités du feu sur les DL et les réservoirs de MO du sol ainsi que pour établir un lien entre les pertes de C et de N et la productivité des peuplements. ©2006 NRC Canada

In the absence of fire in black spruce-feathermoss stands, a thick forest floor layer dominated by bryophytes and sphagnum accumulates. This layer is associated with wet, cool and nutrient-poor soil conditions conducive to the paludification process and pushing the ecosystem towards an unproductive open black spruce forest. The presence of Populus tremuloides in theses stands may halt this process because this species has a high nutrient cycling rate and a litter that represses moss cover. The main hypothesis of this study is that, despite similar abiotic conditions (slope and drainage), the presence of Populus tremuloides in a stand dominated by Picea mariana affects surface soil nutrient availability, total N, pH as well as the decomposition process. The abundance of Populus tremuloides trees was associated with higher exchangeable cations, cationic exchangeable capacity and pH of the forest floor layer on all sites. A decrease in organic matter thickness with increasing aspen presence was also found on all sites, suggesting that this species affects the decomposition process by the quality of its litter as well as by a general improvement of soil physical and chemical properties. The decomposition rate of a standard substrate as well as in vitro potential net nitrogen mineralization were positively related to Populus tremuloides on only one of the three sites, and non-significant on the other sites. Strong immobilization of added nitrogen during incubation was observed on all sites and was not related to aspen, which suggested that in these stands, the soil microbial community is uniformly and strongly nitrogen limited. The zone of influence of Populus tremuloides was evaluated in areas around the soil sampling plot ranging from 3 to 7 m. The results revealed that this zone varies with soil properties. The results suggest that the presence of Populus tremuloides accelerate nutrient cycling, which could affect stand productivity to some extent.

Northern peatlands occupy approximately 4% of the global land surface and store about 30% of the global soil carbon (C). A compilation of C accumulation rates in northern peatlands indicated a long-term average rate of C accumulation of 24.1 g m–2 year–1. However, several studies have indicated that on a short-time scale and given the proper conditions, these ecosystems can exhibit very high rates of C accumulation (up to 425 g m–2 year–1). Peatland development is related to precipitation and temperature, and climate change is expected to have an important impact on the C balance of this ecosystem. Given the expected climate change, we suggest that most of the northern forested peatlands located in areas where precipitation is expected to increase (eastern Canada, Alaska, FSU, and Fennoscandia) will continue to act as a C sink in the future. In contrast, forested peatlands of western and central Canada, where precipitation is predicted to decrease, should have a reduction in their C sequestration rates and (or) could become a C source. These trends could be affected by forest management in forested peatlands and by changes in fire cycles. Careful logging, as opposed to wildfire, will facilitate C sequestration in forested peatlands and boreal forest stands prone to paludification while silvicultural treatments (e.g., drainage, site preparation) recommended to increase site productivity will enhance C losses from the soil, but this loss could be compensated by an increase in C storage in tree biomass.

Les tourbières nordiques occupent approximativement 4 % de la surface terrestre et emmagasinent environ 30 % du carbone (C) qu'on retrouve dans le sol. Une revue sur les taux d'accumulation de C, dans les tourbières nordiques, indique que l'accumulation moyenne de C à long terme est de 24,1 g m–2 a–1. Cependant, plusieurs études ont démontré que sur une courte période de temps, et dans des conditions appropriées, ces écosystèmes peuvent montrer de hauts taux d'accumulation de C (jusqu'à 425 g m–2 a–1). Le développement des tourbières dépend des régimes de précipitation et de température, et les changements climatiques sont susceptibles d'avoir un effet important sur la balance du C dans cet écosystème. Selon les changements climatiques prévus, nous suggérerons que la plupart des tourbières boisées en milieu nordique, où des augmentations de précipitations sont prévues (est du Canada, Alaska, ex-URSS et Fenno Scandinavie), vont continuer à être des puits de C dans le futur. À l'opposé, dans les tourbières boisées de l'ouest et du centre du Canada où les précipitations doivent diminuer, il devrait y avoir une réduction du taux de séquestration de C et/ou ces tourbières deviendront une source de C. Ces tendances pourraient être affectées par l'aménagement forestier dans les tourbières boisées, ainsi que par les changements dans les cycles des feux. Contrairement au feu, la coupe facilite la séquestration du C dans les tourbières boisées et dans les sites sujets à la paludification, alors que les traitements sylvicoles (drainage, préparation de terrain), recommandés pour augmenter la productivité des sites augmentent les pertes de C par décomposition, mais cette perte pourrait être compensée par une augmentation de C dans la biomasse végétale.

In the western boreal forest of Quebec, black spruce stand productivity is approximately 1 m3/ha/year. The low productivity of these stands is often attributed to the paludification process, which is sustained by low quality black spruce litter and the influence of black spruce on soil moisture. In contrast, aspen hasten nutrient cycling, suggesting that the presence of aspen in black spruce-dominated stands could offset the effect of black spruce on soil processes and positively affect stand productivity. We hypothesised that aspen in black spruce-dominated stands could: (1) increase black spruce DBH, height and volume per stem, (2) increase black spruce productivity without affecting black spruce volume in the stand, and (3) increase total stand volume. In 2001, twelve 14 m diameter plots were sampled for DBH and height of every stem on three black spruce-dominated sites containing various proportions of aspen. Using stem analysis, the time to grow from a height of 5 m to a height of 10 m was determined on three dominant black spruces in each plot. Statistical analyses revealed that DBH, height and volume per black spruce stem were not affected by aspen. However, total black spruce volume decreased with increasing aspen basal area in sites 2 and 3, suggesting that the presence of aspen reduced black spruce density. In site 1, black spruce volume was not affected by aspen, indicating, for total stand productivity, a net gain in aspen fibre. Along a gradient of increasing aspen basal area, the time to grow 5 m decreased in sites 1 and 3. These results suggest that the presence of aspen influences black spruce productivity, although this influence is site-specific and could be dependent on the proportion of aspen, its hierarchical position in the canopy, and the nutrient status of the site. To some extent, this could explain the absence of a general trend concerning mixed stand productivity. © 2005 Elsevier B.V. All rights reserved.

In the southeastern boreal forest of Canada, the presence of mixed stands of black spruce (Picea mariana (Mill.) BSP) and trembling aspen (Populus tremuloides Michx.) growing in similar abiotic conditions offers the opportunity to study the influence of aspen on stand volume and spruce growth. A regression analysis performed on field data from the ministère des Ressources naturelles du Québec showed a significant relationship between the relative basal area of aspen (aspen relative basal area was determined by the ratio of aspen basal area to total basal area of the stand) and the total stand merchantable volume after accounting for stand density. However, the relationship between total black spruce volume and relative basal area of aspen was not significant, implying that the volume gain was, in fact, aspen fibre. The positive effects of aspen on black spruce DBH and height were only present when the proportion of aspen in the stand ranged between 0% and 41% of the total stand basal area. These results suggest that aspen uses a different niche than black spruce. Furthermore, the significant increase in black spruce dominant height along the aspen gradient suggests that aspen enhances soil fertility by its influence on nutrient availability. The management of mixed stands, which make up an important proportion on the landscape, offers an example as to how commercial management of the forest can be in agreement with ecosystem management.

Dans la forêt boréale du Sud-Est du Canada, la présence de peuplements mixtes d'épinette noire (Picea mariana (Mill.) BSP) et de peuplier faux-tremble (Populus tremuloides Michx.) croissant dans des conditions abiotiques similaires a offert l'opportunité d'étudier l'influence du peuplier faux-tremble sur le volume du peuplement et la croissance de l'épinette noire. Une analyse de régression effectuée sur des données d'inventaire du ministère des Ressources naturelles du Québec révèle qu'il y a une relation significative entre la surface terrière relative en peuplier et le volume marchand total du peuplement. Cependant, la relation entre la surface terrière relative en peuplier faux-tremble et le volume marchand en épinette noire n'est pas significative, suggérant un gain net en fibre de peuplier. L'effet positif du peuplier faux-tremble sur le DHP et la hauteur de l'épinette noire est présent seulement lorsque le peuplier faux-tremble couvre de 0 à 41 % de la surface terrière du peuplement. Ces résultats suggèrent que les deux espèces utilisent une niche écologique différente. De plus, l'augmentation significative de la hauteur de l'épinette noire le long du gradient de peuplier faux-tremble suggère que la présence du peuplier augmente la fertilité du sol par son influence sur la disponibilité des nutriments. L'aménagement de peuplements mixtes, qui sont abondamment représentés dans le paysage forestier, pourrait offrir une situation où la mission commerciale de la forêt pourrait être en accord avec l'aménagement écosystémique.©2004 NRC Canada

The relationships between site quality indices (SQI) (estimated height at age 50) of black spruce and Jack pine and site characteristics were investigated over a large territory in the boreal forest of Quebec. The relationships with degree-days (DD) and parent material, which are considered as permanent site factors, were significant for both species. Linear regressions with these two parameters as independent variables explained about 40% of the variability in site quality indices for black spruce and Jack pine. The addition of soil chemistry and biological data to this model indicated a significant contribution of exchangeable Mg concentrations or Al:CEC molar ratios in the forest floor and lichen cover to the prediction of site quality indices. Inclusion of these variables increased the model R² up to 60% for both species. Whether these variables reflect permanent site conditions or conditions related to the history of the sites is unknown. © 2004 Elsevier B.V. All rights reserved.

The SAFE (sylviculture et aménagement forestiers écosystémique) project was set up in 1998 in the Lake Duparquet Research and Teaching Forest to test stand-level silvicultural treatments designed to reflect different aspects of natural forest dynamics. In the winter of 1998–1999, four levels of forest harvesting, including a no-harvest and a clearcut treatment, were applied to even-aged trembling aspen (Populus tremuloides Michx.) stands according to a complete block design with three replications. Two partial cut treatments removed 33% and 61% of the stand basal area. During the first growing season, harvesting induced a large increase in indigenous understorey biomass that paralleled changes in the canopy opening. Aspen sucker density increased from 4916 stems/ha in the control to 28 751 and 63 333 stems/ha in the one-third and two-thirds harvesting treatments and 102 916 stems/ha in the clearcut. Most changes in nutrient cycling occurred in the second year and included an increase in forest floor organic C, total N, and base cation availability and a decrease in microbial C/N ratio. These changes may have occurred in response to reduced vegetation uptake and woody debris abundance.

Le projet SAFE (sylviculture et aménagement forestiers écosystémique) a débuté en 1998 dans la Forêt d'enseignement et de recherche du lac Duparquet afin de tester différents traitements sylvicoles inspirés de la dynamique naturelle des peuplements. Quatre intensités de récolte, incluant la coupe totale, la récolte de 33 et 61% de la surface terrière totale et un traitement témoin non coupé, ont été appliquées à des peuplements équiennes de tremble (Populus tremuloides Michx.) au cours de l'hiver 1998–1999, selon un plan en block complet avec trois répétitions. La végétation de sous bois a répondu par une augmentation de sa biomasse proportionnelle à l'ouverture du couvert. Au cours de la première année, la densité des drageons de tremble était de 4916 tiges/ha dans le traitement témoin, 28 751 et 63 333 tiges/ha dans les coupes partielles et 102 916 tiges/ha dans la coupe totale. La plupart des changements observés dans le cycle des nutriments sont survenus dans la couverture morte pendant la deuxième année suivant la coupe. Parmi ceux-ci figurent une augmentation du C organique, du N total et des cations basiques échangeables et une diminution du ratio C/N de la biomasse microbienne. La diminution du prélèvement par les plantes et les changements dans l'abondance des débris ligneux expliqueraient ces changements.©2004 NRC Canada

Ectomycorrhizal (ECM) diversity was measured in 12 mixed-wood stands in the Abitibi region of north-western Quebec. Stands were of similar age and were situated on similar mineral soil deposits, but supported varying proportions of ECM host trees. Host roots were sampled in a manner that enabled their separation into species on the basis of wood anatomy. Shannon diversity indices for the ECM colonizing each host species were determined on the basis of ECM anatomy. The diversity of overstory trees, understory plants and host roots, as well as overstory tree composition, root density and pertinent abiotic factors were measured and used as independent variables in multiple regressions against ECM diversity. We found a positive relationship between overstory tree diversity and ECM diversity, which appears related to fungal host specificity. Although no direct relationship was seen between ECM diversity and soil factors, levels of exchangeable base cations were related to ECM fungal species composition which correlated with ECM diversity at the scale sampled.

Woodland caribou (Rangifer tarandus caribou) require a diversity of forested habitats over large areas and may thus be particularly affected by the large-scale changes in the composition and age-class distribution of forest landscapes induced by the northern expansion of forest management. In this study we examine habitat characteristics associated to the use of calving areas by woodland caribou females and calves at different spatial scales. Thirty females were captured and collared with Argos satellite transmitters that allowed to locate 14 calving areas. Field surveys were conducted at each of these areas to measure the landscape composition of forest cover types and local vegetation characteristics that are used for both forage conditions and protection cover. At the scale of the calving area, univariate comparisons of the amount of forest cover types between sites with and without calves showed that the presence of calves was associated to mature black spruce forest with a high percent cover of terrestrial lichens. Within calving grounds, univariate comparisons showed that vegetation features like ericaceans and terrestrial lichens, that are important food resources for lactating females, were more abundant in calving areas where females were seen with a calf in mid-July than in areas where females were seen alone. The protection of the vegetation cover against predators was however similar between calving areas with or with-out a calf. Logistic regression results also indicated that vegetation characteristics associated to forage conditions were pos-itively associated to calf presence on calving grounds. Our results suggest that foraging conditions should be given more attention in analyses on habitat requirements of woodland caribou.

A sustainable forest management system requires that a balance must be reached between ecosystem nutrient losses and gains in the course of a rotation. In order to determine the influence of stand characteristics (species composition, density, site potential productivity), method of forest harvesting (stem-only versus whole-tree) as well as rotation length on nutrient losses induced by biomass harvesting, a geochemical balance (nutrient inputs minus outputs) was computed from published information and forest inventory databases for the southern portion of the boreal forest of Quebec. Losses were compared with potential nutrient gains that varied according to soil types. We provided a tool for assessing the risk of having a negative nutrient budget (outputs>inputs ) that forest managers can use with information that is already available to them. This exercise was conducted for five commercial tree species, namely paper birch (Betula papyrifera Marsh.), aspen (Populus tremuloides Michx.), balsam fir (Abies balsamea (L.) Mill.), jack pine (Pinus banksiana Lamb.), and black spruce (Picea mariana (Mill.) B.S.P.), grouped into four site indexes and three stand density classes. Strong differences in nutrient exportation in biomass appeared between stands of different compositions as well as between stands of the same species but of different classes of productivity or density. The most nutrient-demanding tree species were trembling aspen and balsam fir. As expected, whole-tree harvesting caused a greater drain on nutrient reserves than stem-only harvesting, but this effect varied strongly with tree species and was greatest for balsam fir and lowest for jack pine. Harvesting the forest before or after the age of financial maturity, which might be desirable under some circumstances, generally created a lesser nutrient drain but this was at the expense of biomass production. Aspen was an exception to this rule showing a greater nutrient drain for stands harvested prior to the age of financial maturity. Implications for the development of indicators of sustainable forestry and for future research are discussed.

Forest overstory composition influences both light and nutrient availability in the mixed boreal forest. The influence of stand composition on understory cover and biomass was investigated on two soil types (clay and till deposits). Four forest composition types were considered in this study: aspen (Populus tremuloides Michx.), paper birch (Betula papyrifera Marsh.), jack pine (Pinus banksiana Lamb.) and a mixture of balsam-fir (Abies balsamea (L.) Mill.) and white spruce (Picea glauca (Moench) Voss). The cover of all understory species was recorded while the biomass of two important and ubiquitous species was measured: mountain maple (Acer spicatum Lam.) of the shrub layer and large-leaved aster (Aster macrophyllus L.) of the herb layer. Soil analyses were conducted to evaluate the influence of overstory composition on understory biomass through its influences on soil characteristics. Analyses of variance showed a significant effect of forest canopy type on mountain maple biomass, understory cover and shrub cover but not on herb cover and large-leaved aster biomass. Path analysis was performed to explore the relationships between canopy type, nutrient availability and understory biomass. Contrary to what was expected, the variation in plant biomass associated with forest composition was weakly related to soil nutrient availability and more strongly related to stand structural attributes.

Concerns over decreases in soil nitrogen reserves and productivity following the removal of logging residues (windrowing, shearblading and piling.) have been raised by numerous researchers. Medium-term impacts of this practice on soil N reserves and availability and on indices of organic matter quality were assessed for balsam fire (Abies balsamea (L.) Mill.), white birch (Betula paperyfera Marsh.) and white spruce (Picea glauca (Moench) Voss) stands growing on dry to fresh clayey sites in northwestern Quebec, Canada. Unharvested control stands, whole-tree harvested cutovers. and windrowed sites were compared. Fifteen years following harvesting and windrowing, forest floor Kjeldahl N concentrations and content and forest floor in situ net N mineralization rates (undisturbed closed top cores incubation) were affected by harvesting but not by windrowing. No differences in mineralization constant, potentially mineralizable N and cumulative mineralized N (526 day incubation period) were found between treatments, suggesting that treatment differences in field net N mineralization rates were the result of interactions between residual ecosystem structures such as forest floor. coarse woody debris and vegetation and meteorological conditions. If these trends persist over time, it could signal that, while whole-tree harvesting does not have a direct effect on soil organic matter quality, long-term impacts on N dynamics could result from changes in ecosystem structures. Slash removal following whole-tree harvesting did not have any additional negative impact. (C) 2002 Elsevier Science B.V. All rights reserved. © 2002 Elsevier B.V. All rights reserved.

Variation in canopy composition can influence ecosystem processes, such as nutrient cycling and light transmittance, even when environmental soil conditions are similar. To determine whether forest cover type influences species composition of the understory vegetation (herbs and shrubs), the composition of this layer was studied on two different surface deposits, clay and till, and under four different forest cover types dominated, respectively, by Populus tremuloides Michx. (aspen), Betula papyrifera Marsh. (white birch), Pinus banksiana Lamb. (jack pine), and Picea glauca (Moench) Voss - Abies balsamea (L.) Mill. (spruce-fir) over similar environmental conditions. Detrended correspondence analysis and analysis of variance performed on the ordination scores revealed that understory plant composition was highly affected by surface deposit and forest cover. The gradient observed in the correspondence analysis proceeds from aspen, white birch, spruce-fir, to jack pine. Indicator species were identified for each surface deposit and cover type, and most of them were associated with either jack pine or aspen. The richness, evenness, and diversity of the understory vegetation did not vary between cover types, but were affected by surface deposit. By controlling ecosystem processes such as light transmittance and nutrient cycling, forest cover influences understory composition.