Mixed plantations are garnering increased attention due to their potential to provide a broader array of benefits compared to monocultures. Although numerous studies have indicated promising complementarity between black spruce (Picea mariana) and tamarack (Larix laricina), few have delved into individual tree growth interactions to thoroughly assess early growth complementarity. We sampled 119 planted black spruce and their immediate environment to quantify and qualify any differences between two conditions in young plantations: mixed tamarack (?mixtures?) and black spruce monocultures (?monocultures?) within young plantations. We investigated the effect of neighbouring under four perspectives: tree competition, microenvironment, foliar nutrients, and soil nutrients. Our results showed increased values for black spruce foliar nitrogen total concentration, soil pH, and canopy closure in mixtures compared to monocultures. Furthermore, black spruce stem volume was increased by 38.1% in mixture compared to monoculture. Black spruce stem volume was negatively affected (86% decrease) by the combined effect of shrubs and non-crop trees under high competition pressure, despite the plantation being mechanically released in 2017. Collectively, our results suggest that black spruce growing in mixtures holds a greater growth potential than black spruce in monocultures.

Mechanical site preparation (MSP) is used prior to planting to control competing vegetation and enhance soil conditions, particularly in areas prone to paludification. Tree planting density can be adapted to the management context and objectives, as it influences yield and wood quality. However, the combined effects of MSP and planting density on understory vegetation composition, functional traits, and diversity remain uncertain. We thus conducted a study in the Clay Belt region of northwestern Quebec, Canada. After careful logging, the study area was divided into nine sites, each receiving one of three treatments: plowing, disc trenching, or no preparation. Sites were further divided into two, with black spruce (Picea mariana [Mill.] Britton, Sterns & Poggenb.) seedlings planted at either a low planting density of 1100 seedlings ha-1 or a high planting density of 2500 seedlings ha-1. After nine years, we assessed understory composition, diversity, key functional traits, sapling density and growth of planted trees. Careful logging alone led to a higher density of naturally established conifers compared to plowing or disc trenching. The interaction between planting density and MSP significantly influenced understory diversity and composition in plowed plots. Understory composition was affected by the soil C/N ratio, coniferous species, and deciduous species density. The growth of black spruce was notably enhanced with higher planting density in the plow treatment only. Neither planting density nor MSP alone affected tree height and diameter. Our results suggest that combining plowing with high-density planting can enhance stand growth and improve forest productivity. These findings guide future research on paludified forests.

<p>Vulnerability curves to cavitation (VC) and their derived parameters, such as <em>P₅₀</em>, are increasingly used and reported to assess forest vulnerability to drought and predict forest responses to climate change. Forest practitioners and policy-makers are encouraged to rely on these parameters to support species selection based on their sensitivity to drought. However, in the majority of studies, we consider that the variability of VC parameters is not clearly reported nor considered, which can lead to counterproductive decisions. In this opinion paper, we demonstrate the importance of precisely reporting the variability around VC parameters and the sources of this variability (plant materials, methods, etc.). We also identify the information that should be provided when reporting mean values of VC parameters. To support our argument, we built VCs for three <em>Picea</em> species and <em>Pinus strobus</em>, using different methods, and compared the value of <em>P₅₀</em> determined in our experiments with values from a literature review.</p>

The future climate of northern temperate forests is projected to be drier and warmer by the end of this century. As a result, more drought-induced forest dieback events are anticipated in northeastern North America, and assessing the vulnerability of dominant tree species to drought is critical for understanding the future composition of these forests. In a greenhouse experiment, we exposed two-year-old seedlings of Picea glauca (Moench) Voss, Picea mariana (Mill.) B.S.P. and Pinus strobus L. to three future climate treatments for southern Quebec, Canada, and evaluated their mortality, growth, and foliage water status responses to soil water availability and atmospheric drought. Using a unique approach, climate treatments emulated droughts of different frequencies, durations, and intensities. Treatments closely simulated one growing season, with changes in air temperature and relative humidity every six hours and daily adjustment in the amount of water delivered to the seedlings. The three species experienced high mortality (75%) in all water-limited treatments compared to a control treatment that provided non-limiting soil moisture (0% mortality). The biomass of the seedlings that survived was 40% lower than that of control seedlings. Our results confirmed that the hydraulic safety margins, defined as the difference between seasonal minimum water potential and xylem water potential leading to 12, 50 and 88% of hydraulic conductivity loss, were good predictors of probability of tree mortality. Therefore, hydraulic safety margins are useful functional traits that can be used to compare the vulnerability of various species to drought and then provide crucial information to practitioners and policymakers to adjust forest management to climate change. We showed that three dominant conifer species of northern temperate forests were highly vulnerable to drought in future climates. Because drought is projected to be a significant threat to forests, understanding potentially adaptive physiological responses to drought, such as hydraulic safety margins of tree seedlings, is important for predicting the response of forest regeneration and composition in warmer and drier climates.

Little is known about the growth interactions of black spruce (Picea mariana) and tamarack (Larix laricina), two important commercial tree species of the Canadian boreal forest. We investigated growth relations between black spruce and tamarack in mature mixed stands. We sampled tree

Ecosystem-based forest management associated with partial harvesting (PH) is intended to balance ecological and economic values of sustainable forest management. The potential for delayed growth response and elevated mortality of advance regeneration following PH remains a critical concern, and may present a barrier to more widespread implementation of this approach. We used 835 permanent continuous forest inventory plots to examine the rate and time course of species-specific regeneration growth and mortality of eight tree species in the first fifteen years following operational partial harvests in the mixed-species forests of Maine, United States. We aimed to provide a quantitative understanding on how regeneration of different species responded to PH in terms of growth and mortality. In addition, we evaluated how the patterns and magnitudes of growth and mortality responses developed over time, if these responses occur gradually or suddenly, and if the patterns of the responses were persistent. We found that the response magnitude, temporal trajectories of responses, and the length of initial lag-period largely varied across species, PH treatments, and the variables examined. For sapling diameter growth, paper birch (Betula papyrifera Marshall) and red maple (Acer rubrum L.) showed immediate responses to high-intensity PH, while a five-year lag-period was observed in balsam fir (Abies balsamea (L.) Mill.), American beech (Fagus grandifolia Ehrh.), red spruce (Picea rubens Sarg.) and eastern hemlock (Tsuga canadensis (L.) Carrière) and a 10-year lag period in northern white-cedar (Thuja occidentalis L.). The initial increase in sapling mortality was observed in balsam fir, American beech, red maple and northern white-cedar, but not in other species. Sapling survival reached a stable state irrespective of species after the initial five-years following harvests. In partially harvested stands, identifying preharvest conditions related to postharvest density, growth, and mortality was complex and interacted with time since harvest. Our results suggest that broad application of PH only results in species-specific gains, losses, and delays in regeneration responses within mixed-species stands. Future research should consider PH in combination with other treatments to initiate immediate responses to a wider range of species.

In 2020, Quebec adopted a strategy to increase the quantity and quality of timber it produces. During a roundtable discussion held in the fall of 2021, experts in forestry and in related fields expressed their views on the new strategy and its implementation challenges. The main purpose of this article is to present the key observations from the roundtable. The observations addressed two themes: the general context in which the strategy was developed, and the context of its implementation on the ground. Although most of the panellists agreed on the relevance of such a strategy, particularly as regards to climate change mitigation and wealth creation, several questions remain. The challenge of harmonizing uses, regionalization, spatialization of management decisions, labour shortage, and uncertain ecosystem dynamics make it difficult to assess the strategy’s potential impact on the ground and its ability to achieve its targets.

En 2020, le Québec a adopté une stratégie nationale de production de bois (SNPB) afin d’augmenter la quantité et la qualité de la matière ligneuse produite. Au cours d’une table ronde tenue à l’automne 2021, des experts de la foresterie et de domaines connexes se sont prononcés sur cette nouvelle stratégie et sur les défis de mise en oeuvre qu’elle pose. L’objectif principal de cet article est de présenter les principaux constats émis au cours de cette table. Les constats ont été divisés en deux thématiques, soit le contexte général d’élaboration de cette stratégie et le contexte de sa mise en oeuvre en forêt. Bien que la plupart des panélistes s’entendent sur la pertinence de créer une telle stratégie, notamment en ce qui a trait à l’atténuation des changements climatiques et à la création de richesses, plusieurs interrogations persistent. Les défis d’harmonisation des usages, de régionalisation, de spatialisation des décisions d’aménagement, de manque de maind’oeuvre et de la dynamique incertaine des écosystèmes complexifient l’évaluation des retombées potentielles de la SNPB sur le terrain et sa capacité d’atteindre les cibles établies.

This data article describes datasets of plant community composition, dendrometric measurements, quantity and quality of snags of humid boreal stands (Quebec, Canada) from an experiment comparing silviculture scenarios of increasing intensity: (i) careful logging around advance growth (CLAAG); (ii) CLAAG followed by pre-commercial thinning; (iii) plantation followed by mechanical release; and (iv) plantation followed by chemical release and within five naturally disturbed sites. These data enable researchers to examine vegetation biodiversity recovery, ecosystem variables such as dead wood, and boreal stand productivity 20 years following the start of increasing-intensity silviculture scenarios. As a result, these data can be used to investigate the trade-off between keeping important ecosystem aspects of natural forests and maintaining and/or growing merchantable wood production at the stand level. This trade-off is the paradigm of forest ecosystem-based management, which aims to reduce the ecological distance between natural and managed forests in order to balance ecological challenges with the provision of socioeconomic services.

Intraspecific trait variability (ITV) provides the material for species' adaptation to environmental changes. To advance our understanding of how ITV can contribute to species' adaptation to a wide range of environmental conditions, we studied five widespread understorey forest species exposed to both continental-scale climate gradients, and local soil and disturbance gradients. We investigated the environmental drivers of between-site leaf and root trait variation, and tested whether higher between-site ITV was associated with increased trait sensitivity to environmental variation (i.e. environmental fit). We measured morphological (specific leaf area: SLA, specific root length: SRL) and chemical traits (Leaf and Root N, P, K, Mg, Ca) of five forest understorey vascular plant species at 78 sites across Canada. A total of 261 species-by-site combinations spanning ~4300 km were sampled, capturing important abiotic and biotic environmental gradients (neighbourhood composition, canopy structure, soil conditions, climate). We used multivariate and univariate linear mixed models to identify drivers of ITV and test the association of between-site ITV with environmental fit. Between-site ITV of leaf traits was primarily driven by canopy structure and climate. Comparatively, environmental drivers explained only a small proportion of variability in root traits: these relationships were trait specific and included soil conditions (Root P), canopy structure (Root N) and neighbourhood composition (SRL, Root K). Between-site ITV was associated with increased environmental fit only for a minority of traits, primarily in response to climate (SLA, Leaf N, SRL). Synthesis. By studying how ITV is structured along environmental gradients among species adapted to a wide range of conditions, we can begin to understand how individual species might respond to environmental change. Our results show that generalisable trait–environment relationships occur primarily aboveground, and only accounted for a small proportion of variability. For our group of species with broad ecological niches, variability in traits was only rarely associated with higher environmental fit, and primarily along climatic gradients. These results point to promising research avenues on the various ways in which trait variation can affect species' performance along different environmental gradients.

Successive disturbances such as fire can affect post-disturbance regeneration density, with documented adverse effects on subsequent stand productivity. We conducted a simulation study to assess the potential of reactive (reforestation) and proactive (variable retention harvesting) post-fire regeneration failure mitigation strategies in a 1.37 Mha fire-prone boreal landscape dominated by black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.). We quantified their respective capacity to maintain landscape productivity and post-fire resilience, as well as their associated financial returns under current and projected (RCP 8.5) fire regimes. While post-fire reforestation with jack pine revealed to be the most effective strategy to maintain potential production, associated costs quickly became prohibitive when applied over extensive areas. Proactive strategies such as an extensive use of variable retention harvesting, combined with replanting of fire-adapted jack pine only in easily accessible areas, appeared as a more promising approach. Despite this, our results suggest an inevitable erosion of forest productivity due to post-fire regeneration failure events, highlighting the importance of integrating fire a priori in strategic forest management planning as well as its effects on long-term regeneration dynamics.

{Changing climates are altering the structural and functional components of forest ecosystems at an unprecedented rate. Simultaneously, we are seeing a diversification of public expectations on the broader sustainable use of forest resources beyond timber production. As a result, the science and art of silviculture needs to adapt to these changing realities. In this piece, we argue that silviculturists are gradually shifting from the application of empirically derived silvicultural scenarios to new sets of approaches, methods and practices, a process that calls for broadening our conception of silviculture as a scientific discipline. We propose a holistic view of silviculture revolving around three key themes: observe, anticipate and adapt. In observe, we present how recent advances in remote sensing now enable silviculturists to observe forest structural, compositional and functional attributes in near-real-time, which in turn facilitates the deployment of efficient, targeted silvicultural measures in practice that are adapted to rapidly changing constraints. In anticipate, we highlight the importance of developing state-of-the-art models designed to take into account the effects of changing environmental conditions on forest growth and dynamics. In adapt, we discuss the need to provide spatially explicit guidance for the implementation of adaptive silvicultural actions that are efficient, cost-effective and socially acceptable. We conclude by presenting key steps towards the development of new tools and practical knowledge that will ensure meeting societal demands in rapidly changing environmental conditions. We classify these actions into three main categories: re-examining existing silvicultural trials to identify key stand attributes associated with the resistance and resilience of forests to multiple stressors, developing technological workflows and infrastructures to allow for continuous forest inventory updating frameworks, and implementing bold, innovative silvicultural trials in consultation with the relevant communities where a range of adaptive silvicultural strategies are tested. In this holistic perspective, silviculture can be defined as the science of observing forest condition and anticipating its development to apply tending and regeneration treatments adapted to a multiplicity of desired outcomes in rapidly changing realities.}

Natural forest regeneration after natural or anthropogenic disturbance is difficult to predict given its high variability. The process is poorly documented for commercial northern hardwood species in the Acadian forest of eastern Canada. Our objective was to identify the silvicultural, environmental, and ecological factors that best explain the variability in sapling density and occurrence of two commercial tolerant hardwood species in New Brunswick: American beech (Fagus grandifolia Ehrh.) and sugar maple (Acer saccharum Marsh.). Forty-three permanent sample plots were established in 2002 and measured before harvesting in 2004. Sapling density and occurrence were measured 14 years after harvesting. The results showed that the interactions between the species and the residual merchantable basal area and between the species and the percent of hardwoods in the original stand best explained the sapling density and occurrence variation of tolerant hardwoods. The sapling density of sugar maple increased with increasing merchantable residual basal area. However, the effect of this variable was not significant for the density of American beech saplings. The density and occurrence of tolerant hardwood saplings both increased along with the percent of hardwoods in the original stand. These results provide an improved understanding about tolerant hardwood regeneration dynamics in New Brunswick forests.

Understory vegetation plays an important role in maintaining biodiversity, soil nutrient cycling and carbon stocks, yet the ability of understory functional traits to predict forest productivity after harvesting disturbance is unknown. Our objective was to investigate the utility of individual traits (the community-weighted trait mean) and combined information from multiple traits (functional diversity indices) of understory plants in predicting tree diameter growth of black spruce (Picea mariana (Mill.) B.S.P.) after two harvesting methods (clearcutting and careful logging). We used vegetation and tree growth data from a network of field plots (171 plots on 43 sites) established in black spruce dominated forests across Ontario and Quebec, in northeastern Canada. Functional traits (11 traits) of dominant understory species that reflect plant morphology, regeneration strategy, and resource utilization were collected. We then used generalized linear mixed models (GLMMs) to model the response of tree diameter increment to functional diversity indices, community-weighted trait means and their interactions with harvesting method. Our results showed that: 1) at the functional diversity level, functional richness had a positive relationship with tree growth in clear-cut forests, while functional evenness showed a negative and functional divergence a positive relationship with tree growth in careful logging forests; 2) at the individual trait level, understory species with relatively large morphological trait values often promoted tree growth in both forest types, with stronger relationships in careful logging than in clear-cut forests. For regeneration traits, heavier seed weight or permanent seed bank persistence of understory plants had negative relationships with tree growth, and those relationships were either only found in careful logging forests or did not depend on harvesting type. A positive relationship was found between vegetative propagation species and tree growth, which was stronger in careful logging than in clear-cut forests. Species preferring either humid or xeric habitat had positive relationships with tree growth; both shade-tolerant and shade-intolerant species also positively correlated to tree growth. In conclusion, our study shows the important roles of understory traits in predicting tree growth based on both niche complementarity and mass-ratio hypotheses. Moreover, disturbance severity influenced the balance or degree of the positive and negative interactions that occurred between understory plants and black spruce tree growth. We suggest that identifying the threshold of harvesting disturbance severity that can activate facilitation interactions between understory plants and crop trees would be an important topic for future study.

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

Sustainable forest management implies successful regeneration after disturbances. Low N availability and competition can, however, limit tree establishment in boreal ecosystems. To develop silviculture strategies that maintain productivity in such context, we established a field trial in northern Québec, Canada. We evaluated if a companion N2-fixing species (Alnus alnobetula) promotes or hinders Picea mariana and Pinus banksiana establishment over six growing seasons. We tested if Alnus has a facilitation effect through nutritional processes and a competition effect through light interception. Foliar stable nitrogen isotope ratio (?15N?=?15N/14N, ‰) results confirmed that Alnus obtains a substantial part of its N through biological fixation and represents an N source in this system. Although we did not observe increased foliar N concentrations in either conifer species in the presence of Alnus, Pinus growth was nonetheless higher in presence of Alnus, whereas no difference was observed for Picea. In the plots where Alnus cohabited with the conifers, the former had a negative impact on seedling growth, suggesting a significant competition for light. Overall, the net effect of Alnus was positive for Pinus and neutral for Picea. Our results have significant implications for silviculture in N-limited systems, especially in the context of climate change that imposes increased levels of stress on regeneration.

In eastern Canada, spruces (Picea spp.) and pines (Pinus spp.) are among the main commercial species being logged for their lumber or wood fiber. Annually, about 175 million seedlings are planted in areas totaling ~100,000 ha. Appropriate microsite selection is essential during reforestation operations, given that it can improve the chances of survival and initial growth of the seedlings. In fir (Abies spp.) and spruce forests of eastern Canada, the optimal characteristics of establishment microsites have yet to be identified; these would be determined by different physical and climatic variables operating at several scales. Our study determined the influence of climatic (regional-scale), edaphic (stand-scale), local (microsite-scale) and planting conditions on the establishment substrate and initial growth of black spruce (Picea mariana Britton, Sterns and Poggenb.) and jack pine (Pinus banksiana Lamb.). Substrate characterization and growth monitoring (three growing seasons) for the two species were conducted on 29 planted cutblocks that were distributed over an east–west climatic gradient (precipitation and temperature) in the balsam fir and black spruce–feather moss forests of Quebec (Canada). Linear mixed models and multivariate analyses (PCAs) determined the effects of climatic, edaphic and micro-environmental variables and their interactions on the establishment substrate and seedling initial growth. The predictive models explained, respectively, 61% and 75% of the growth variability of black spruce and jack pine. Successful establishment of black spruce and jack pine depended upon regional conditions of precipitations and temperature, as well as on their interactions with stand-scale edaphic variables (surface deposit, drainage and slope) and local variables (micro-environmental) at the microsite-scale (establishment substrate types and substrate temperature). Mineral, organo-mineral and organic establishment substrates exerted mixed effects on seedling growth according to regional precipitation and temperature conditions, as well as their interactions with edaphic and local variables at the stand and microsite-scales, respectively. View Full-Text

In the boreal forest of eastern Canada, a large proportion of black spruce (Picea mariana [Mill.] Britton, Sterns & Poggenb.) stands are affected by paludification. Edaphic conditions that are created by paludification processes, including an abundance of microsites with high moisture and low nutrient contents, hinder forest regeneration. Disturbance of paludified sites by mechanical soil preparation (MSP) reduces organic layer thickness, while generating a range of substrates for regeneration establishment. Yet, little information is available regarding the effects of these substrates on tree growth. Our objective was to determine the effect of organic, mineral and organo-mineral substrates that are created following MSP of a paludified site on the growth and root development of black spruce seedlings in a semi-controlled environment. We demonstrated that substrate exerted a significant effect on seedling growth and foliar concentrations of N, P and K. Increase in height and diameter were respectively greatest on clay (mineral) and mesic substrates. Substrate effects did not affect total biomass increases or final root biomass. Foliar nutrients (N, P, K) were relatively high in seedlings that were established on mesic substrates and relatively low for those established on clay substrates. To ensure successful seedling establishment, we recommend the application of MSP techniques that expose organic-mesic substrates on sites that are susceptible to paludification.

Climate change is projected to increase fire severity and frequency in the boreal forest, but it could also directly affect post-fire recruitment processes by impacting seed production, germination, and seedling growth and survival. We reviewed current knowledge regarding the effects of high temperatures and water deficits on post-fire recruitment processes of four major tree species (Picea mariana, Pinus banksiana, Populus tremuloides and Betula papyrifera) in order to anticipate the effects of climate change on forest recovery following fire in the boreal biome. We also produced maps of future vulnerability of post-fire recruitment by combining tree distributions in Canada with projections of temperature, moisture index and fire regime for the 2041–2070 and 2071–2100 periods. Although our review reveals that information is lacking for some regeneration stages, it highlights the response variability to climate conditions between species. The recruitment process of black spruce is likely to be the most affected by rising temperatures and water deficits, but more tolerant species are also at risk of being impacted by projected climate conditions. Our maps suggest that in eastern Canada, tree species will be vulnerable mainly to projected increases in temperature, while forests will be affected mostly by droughts in western Canada. Conifer-dominated forests are at risk of becoming less productive than they currently are, and eventually, timber supplies from deciduous species-dominated forests could also decrease. Our vulnerability maps are useful for prioritizing areas where regeneration monitoring efforts and adaptive measures could be developed.

Low productivity caused by paludification in some parts of the closed black spruce (Picea mariana (Mill.) B.S.P) dominated boreal forest threatens the provision of ecosystem services, including wood fiber production. The accumulation, over time, of organic matter in paludified soils leads to an anaerobic environment that reduces microbial activity, decelerates decomposition of organic matter, and generates nutrient-poor microsites for regeneration. Consequently, it results in significant impacts on site productivity. Considering its ability to disturb the soil, mechanical site preparation (MSP) is viewed as a potential treatment that can help restore productivity of paludified sites following harvesting. We conducted a field experiment to verify if (1) the availability of microsites conducive to reforestation varies with MSP, microtopography (slope and aspect) and initial OLT conditions; (2) the growth of planted seedlings depends on the intensity of mechanical disturbance of the organic layer, type of microsite, planting density, presence of Ericaceae, and the planting position and depth; (3) there are direct and indirect causal relationships between microsites availability after MSP, OLT, microtopography, planting quality and seedlings growth; and (4) if mechanical site preparation and microsite type exposed affect the Ericaceae cover after planting. Our results confirmed that MSP is effective in establishing conditions that permit a productive regeneration cohort on these paludified sites. To ensure successful establishment of plantations on these sites, it is necessary, however, to distinguish between those that are slightly or moderately paludified from those that are highly paludified, as treatment effectiveness of different MSP types depends on organic layer thickness. Our results also show that preference should be given to some microsite types as clay and mixed-substrate microsites for planting to ensure sufficient availability of water and nutrients for seedlings.

1. Considering intraspecific trait variability (ITV) in ecological studies has improved our understanding of species persistence and coexistence. These advances are based on the growing number of leaf ITV studies over local gradients, but logistical constraints have prevented a solid examination of ITV in root traits or at scales reflecting species’ geographic ranges.
2. We compared the magnitude of ITV in above? and below?ground plant organs across three spatial scales (biophysical region, locality and plot). We focused on six understorey species (four herbs and two shrubs) that occur both in disturbed and undisturbed habitats across boreal and temperate Canadian forests. We aimed to document ITV structure over broad ecological and geographical scales by asking: (a) What is the breadth of ITV across species range?scale? (b) What proportion of ITV is captured at different spatial scales, particularly when local scale disturbances are considered? and (c) Is the variance structure consistent between analogous leaf and root traits, and between morphological and chemical traits?
3. Following standardized methods, we sampled 818 populations across 79 forest plots simultaneously, including disturbed and undisturbed stands, spanning four biophysical regions (~5,200 km). Traits measured included specific leaf area (SLA), specific root length (SRL) and leaf and root nutrient concentrations (N, P, K, Mg, Ca). We used variance decomposition techniques to characterize ITV structure across scales.
4. Our results show that an important proportion of ITV occurred at the local scale when sampling included contrasting environmental conditions resulting from local disturbance. A certain proportion of the variability in both leaf and root traits remained unaccounted for by the three sampling scales included in the design (36% on average), with the largest amount for SRL (54%). Substantial differences in magnitude of ITV were found among the six species, and between analogous traits, suggesting that trait distribution was influenced by species strategy and reflects the extent of understorey environment heterogeneity.
5. Even for species with broad geographical distributions, a large proportion of within?species trait variability can be captured by sampling locally across ecological gradients. This has practical implications for sampling design and trait selection for both local studies and continental?scale modelling.

Roots directly affect planted seedling adaptation to new growing conditions at reforestation sites. To test the influence of root characteristics on the short-term response of seedlings to limiting resources (water, nutrient, or oxygen), we conducted two experiments. We compared (1) the growth and physiology of three types of four-year-old black spruce (Picea mariana (Mill.) BSP) seedlings (Containerized, highly developed initial roots restricted to a plug; bareroot, less developed but unrestricted initial roots; deeply-planted containerized, restricted initial and adventitious roots) to different combinations of irrigation and fertilization. We also investigated (2) the cellular plasticity of adventitious and initial roots to three irrigation regimes including flooding. Bareroot seedlings had better relative growth rates in height than containerized seedlings, probably due to their larger initial size. On the other hand, containerized seedlings took better advantage of fertilization, as shown by a higher relative growth rate in diameter compared to bareroot seedlings and were less affected by water limitation, possibly due to the root plug acting as an additional water reserve capacity. For containerized seedlings, the presence of adventitious roots was beneficial to height growth and physiological performances compared to seedlings with initial roots only. Adventitious roots showed great cell plasticity, particularly under flooding conditions.

Black spruce (Picea mariana [Mill.] BSP) is a boreal tree species characterized by the formation of an adventitious root system. Unlike initial roots from seed germination, adventitious roots gradually appear above the root collar, until they constitute most of mature black spruce root system. Little is known about the physiological role they play and their influence on tree growth relative to initial roots. We hypothesized that adventitious roots present an advantage over initial roots in acquiring water and nutrients. To test this hypothesis, the absorptive capacities of the two root systems were explored in a controlled environment during one growing season. Black spruce seedlings were placed in a double?pot system allowing irrigation (25 and 100% water container capacity) and fertilization (with or without fertilizer) inputs independent to initial and adventitious roots. After 14?weeks, growth parameters (height, diameter, biomass), physiology (net photosynthetic rate, stomatal conductance, shoot water potential) and nutrient content (N, P, K, Ca and Mg foliar content) were compared. Most measured parameters showed no difference for the same treatment on adventitious or initial roots, except for root biomass. Indeed, fertilized black spruce seedlings invested heavily in adventitious root production, twice as much as initial roots. This was also the case when adventitious roots alone were irrigated, while seedlings with adventitious roots subjected to low irrigation produced initial root biomass equivalent to that of adventitious roots. We conclude that black spruce seedlings perform equally well through adventitious and initial roots, but if resources are abundant, they strongly promote development of adventitious roots.

Planting shock can last for several years during which seedling growth may significantly slow down. This duration depends upon the time that is required for the root system to acclimatize to its new environment. We hypothesized that characteristics of the root system are important factors for black spruce (Picea mariana [Mill.] BSP) seedling establishment. To test this hypothesis, we compared growth and physiology of three stock types: containerized seedlings, with a dense initial root system restricted to the volume of a root plug; deeply planted containerized seedlings, with the stem buried during production to develop dense adventitious and initial root systems; and bareroot seedlings, with a less developed but unrestricted initial root system. The three stock types were planted on three microsites that are typical of reforestation sites found in eastern Canadian boreal forest: (1) bare mineral soil; (2) in presence of ericaceous shrubs; or (3) in presence of Sphagnum spp. and ericaceous species. Labelled nitrogen fertilizer was added to half of the seedlings to compare nitrogen uptake. We measured seedling growth (height, diameter, above- and belowground biomass), nutrition (including 15N uptake) and 13C discrimination (as an indicator of water stress) at the end of the second growing season. Except at the root level, there were few interactions between stock types and microsites. For fertilized microsites, we speculate that ericaceous species captured most added nutrients since seedlings that were planted directly on mineral soil had greater height and diameter increments than seedlings with competing vegetation. In unfertilized microsites, Sphagnum spp. and ericaceous species had significant negative effects on seedling growth. For stock type, the presence of a root plug allowed containerized and deeply planted seedlings to have a lower ?13C, suggesting less water stress compared to bareroot seedlings. Strong initial root system development of containerized seedlings resulted in greater total biomass. We hypothesized that the presence of adventitious roots in deeply planted seedlings positively influenced nutrient uptake, particularly N. As nutrient availability is one of the most limiting factors in the boreal forest, we suggest using deeply planted containerized seedlings to reduce planting stress.

Forest regeneration is a key element in achieving sustainable forest management. Partial harvest methods have been used extensively in temperate broadleaf and mixedwood ecosystems to promote regeneration on poorly stocked sites and to maintain forest composition and productivity. However, their effectiveness in promoting conifer establishment has yet to be demonstrated in unmanaged boreal forests, especially those dominated by black spruce (Picea mariana (Mill.) BSP) where constraints for regeneration differ from those found in more meridional regions. We aimed to evaluate conifer seedling density and dimensions, 10 years after the onset of a gradient of silvicultural treatments varying in harvesting intensities, and to identify the critical factors driving the regeneration process. Study blocks of even-aged black spruce stands in the eastern Canadian boreal forest were submitted to three variants of shelterwood harvesting: a seed-tree harvest, a clear-cut and an untreated control. Shelterwood and seed-tree harvesting were combined with spot scarification to promote regeneration. Shelterwood and seed-tree harvesting produced a density of conifer regeneration sufficient to maintain forest productivity, but they did not promote seedling growth. Black spruce was the predominant species in terms of regeneration density, with proportions 3–5× higher than that for balsam fir (Abies balsamea (L.) Mill.). Ten years after treatment, seed-origin black spruce seedlings were abundant in skidding trails, while layers dominated the residual strips. Balsam fir density was not influenced by treatment nor by tree position relative to skidding trails. Balsam fir and black spruce had different responses to treatment in terms of height and diameter, the former exhibiting a better growth performance and larger diameter in the residual strips. Spot scarification created micro-sites that had a significant impact on the regeneration process. Overall, our results support that shelterwood and seed-tree harvesting combined with scarification enable adequate regeneration in black spruce stands, confirming these treatments as viable silvicultural alternatives to clear-cutting when required by sustainable forest management objectives. © 2018 Montoro Girona, Lussier, Morin and Thiffault.

La maîtrise de la végétation concurrente est essentielle afin que les plantations présentent les rendements escomptés, mais ses effets dépendent de son moment d’application. Ainsi, notre objectif était d’évaluer l’effet, après 15–20 ans, d’un délai d’application du dégagement mécanique par rapport au moment requis selon les procédures opérationnelles au Québec (Canada) pour des plantations de Picea glauca et Picea mariana. Nous avons utilisé trois dispositifs expérimentaux de reboisement hâtif comprenant les traitements suivants : i) témoin, non dégagé; ii) dégagement mécanique l’année requise (Requis); iii) dégagement mécanique avec un délai d’une année par rapport à Requis (Requis+1); et, iv) dégagement mécanique avec un délai de deux années (Requis+2). Nos résultats montrent que dans un scénario de reboisement hâtif comprenant un traitement de nettoiement au stade gaulis, il n’y a pas d’impact significatif sur la croissance à retarder l’application d’un dégagement mécanique jusqu’à deux années après le moment requis. Cette marge opérationnelle devrait être utilisée afin d’assurer la réalisation des dégagements; l’omission du dégagement a un effet négatif marqué sur la production aux échelles de l’arbre et du peuplement. Le dégagement a en effet augmenté la hauteur et le diamètre des arbres plantés ainsi que la surface terrière totale des peuplements.

Early successional competition among boreal forest tree and shrub species and its effects on growth of commercial tree species have been a major source of uncertainty in establishing efficient precommercial thinning and brushing prescriptions. We examined the effect of prethinning competitor density, postthinning competitor regrowth density, prethinning stem diameter, and the timing of thinning operations on the growth response of black spruce (Picea mariana [Mill.] BSP) and jack pine (Pinus banksiana Lamb.). In addition, we examined the mortality rate of hardwoods after thinning and the number of new shoots produced per surviving thinned stem. For jack pine, growth response was greatest when thinning occurred between 4 and 9 years after establishment, whereas for black spruce we observed no significant relationship between growth response and the timing of treatment. For jack pine, growth response was significantly affected by pretreatment competitor density, posttreatment competitor regrowth density, and pretreatment stem diameter. For black spruce, no significant relationship was observed between growth response and any variables. Mortality rates and production of new shoots in hardwoods varied significantly between species. Considering the high regrowth potential of willow (Salix spp.) and alder (Alnus spp.), we recommend that stands exhibiting low densities of these species should be left unthinned. Our results help foresters identify stands that require precommercial thinning and call for modification of currently used thinning strategies.

Ecosystem-based management, now a dominant forestry paradigm, implies reducing the gap between variability of natural and managed forests (i.e. ecological distance) to reconcile ecological issues with production of socioeconomic services. Here, we tested whether a trade-off exists between conserving key ecosystem attributes of natural forests and maintaining and/or increasing merchantable wood production at the stand scale in humid boreal stands. Using 20-y data from an experimental design comparing silviculture scenarios of increasing intensity, (i) careful logging around advance growth (CLAAG); (ii) CLAAG followed by pre-commercial thinning; (iii) plantation followed by mechanical release; and (iv) plantation followed by chemical release, we examined plant community composition, stand structure and the quantity and the quality of snags. We also assessed timber productivity by comparing scenarios in terms of conifer and merchantable (diameter at breast height > 9 cm) tree dimensions. We used data from stands originating from a spruce budworm outbreak as a baseline to understand scenario impacts on variability of key attributes and productivity. Our results showed increasing differences in these attributes between natural and managed stands with increasing silviculture intensity: the diameter structure became more homogenized, light demanding species richness and abundance increased and the quantity and the quality of snags decreased. Therefore, our results showed that the ecological distance from naturally disturbed stands was lower after CLAAG than after the other silviculture scenarios. However, CLAAG favored an increase in the density of deciduous trees and a decrease of conifer snag density that have the potential to affect resilience of mature stands. Pre-commercial thinning resulted in crop trees reaching larger diameter than following CLAAG only and in the decrease of birch tree density, with no effect on deciduous regeneration density ? 60 cm in height. We measured higher basal area of merchantable trees in plantations than in stands originating from natural regeneration scenarios, with mechanical and chemical release scenarios resulting in similar crop tree productivity. Globally, our study confirmed a general antagonism between the impacts of silviculture on key ecosystem attributes and forest productivity, posing a challenge for reconciling ecological issues with the production of socioeconomic services. At the stand level, results support that retention forestry could emulate natural disturbances by conserving biological legacies during harvest in humid boreal forests. Further research is needed to determine retention parameters to achieve expected wood production while maintaining variability of key attributes in humid boreal forests. © 2017 Elsevier B.V.

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.

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.

Some regenerating stands of the boreal forest exhibit low juvenile growth after major disturbances, which compromise sustainable forest management objectives. In black spruce-feather moss stands of eastern Canada subject to paludification, careful logging methods could decrease stand productivity with time, by preventing a beneficial reduction in organic soil thickness. This project aimed at confirming decreases in juvenile growth between stands originating from careful logging and the former stands originating from old fires on the same sites. Stem analyses showed that stands originating from CPRS had significant better juvenile height growth than the former stands,but significant lower growth than post-recent fire stands, in the study region. If organic matter thickness apparently played a role in the growth differences observed between fire and harvesting, it was not the only factor determining stand productivity. The cohort status, the climatic regime and the quality of the residual organic matter are other factors that seem to drive productivity according to our results. Our results show that post-harvest management approaches (e.g., site preparation) should be used to increase yields after harvest for the sites to express their full growth potential.

La paludification est l'accumulation de la couche organique partiellement décomposée sur le sol minéral saturé en eau. La paludification réduit la régénération et la croissance des arbres, principalement en raison des faibles températures et la teneur élevée en eau dans la zone racinaire, réduisant ainsi la décomposition de la couche organique, et par conséquent, la disponibilité des éléments nutritifs. Dans la Ceinture d'Argile de l'ouest du Québec et l'est de l'Ontario, les forêts tendent vers une paludification naturellement, mais ce processus pourrait être contré par des méthodes d'exploitation forestière. Notre objectif était d'identifier laquelle des deux techniques de préparation mécanique du terrain (PMT) couramment utilisées est la mieux adaptée pour réduire l’épaisseur de la couche organique (ECO) et générer des microsites de plantation favorables après récolte dans les sites paludifiés. Neuf blocs expérimentaux (entre 20 ha–61 ha chacun) ont été délimités dans un secteur forestier de 35 km2 avec des niveaux variables de paludification. Le secteur a été récolté en utilisant la coupe avec protection de la régénération et des sols (CPRS). Les neuf blocs expérimentaux ont ensuite été traités avec une herse forestière, un scarificateur à disques (T26) ou laissés comme témoins non traités (récolte seulement), avec trois blocs répétés par traitement. Nous avons mesuré l'ECO avant et après PMT et déterminé la qualité des microsites à l'intérieur de chaque bloc. Les résultats ont révélé des différences significatives d'ECO entre les traitements de PMT et la récolte seulement. Globalement, la herse forestière était la meilleure technique de PMT, car cette dernière a réduit davantage l'ECO que le scarificateur T26 et a généré le pourcentage le plus élevé de bons microsites, sauf dans le seuil d'ECO initiale 44 cm–56 cm. Nos résultats favorisent l'utilisation réussie de la PMT dans les forêts paludifiées.

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.

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.