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We investigated how the surrounding environment influences the growth of dominant trees and their responses to temperature and insect epidemics in boreal forests of eastern Canada. We focused on 82 black spruce and jack pine focal trees in stands spanning a double gradient of species diversity and soil texture within a 36 km2 area of western Quebec. For these trees, we compared their diameter at breast height, growth rates, temperature–growth relations, and growth during insect defoliator epidemics. We used linear models to study how surrounding tree attributes and soil properties affected the growth of focal trees. Models showed that tree growth responses and responses to temperature and insect epidemics were generally negative with higher intraspecific competition and positive with greater tree species diversity. Growth of both species benefitted from lower soil sand content. Our research offers novel insights on the potential role of the surrounding environment, notably tree competition and species diversity, in mitigating the vulnerability of eastern Canada's boreal trees to anthropogenic climate change and insect epidemics.

Clear-cutting has been the dominant harvesting method used in boreal forest silviculture. Reducing the potential negative effects of intensive forestry activities on ecosystems, e.g., the simplification and homogenization of stand structure, requires diversifying silvicultural practices to promote forest resilience in the face of climate change. Priority therefore lies in developing, evaluating, and adapting partial cutting as a potential silvicultural option for ensuring the sustainable management of boreal forests. In this chapter, we summarize the findings of two large-scale experiments conducted in Canadian boreal forests that tested new silvicultural approaches and explore their implications for forest management. We discuss the effects of these treatments on tree growth, tree mortality, regeneration, and biodiversity, and we examine the challenges of existing silvicultural approaches in the context of climate change.

Long-term disturbance histories, reconstructed using diverse paleoecological tools, provide high-quality information about pre-observational periods. These data offer a portrait of past environmental variability for understanding the long-term patterns in climate and disturbance regimes and the forest ecosystem response to these changes. Paleoenvironmental records also provide a longer-term context against which current anthropogenic-related environmental changes can be evaluated. Records of the long-term interactions between disturbances, vegetation, and climate help guide forest management practices that aim to mirror “natural” disturbance regimes. In this chapter, we outline how paleoecologists obtain these long-term data sets and extract paleoenvironmental information from a range of sources. We demonstrate how the reconstruction of key disturbances in the boreal forest, such as fire and insect outbreaks, provides critical long-term views of disturbance-climate-vegetation interactions. Recent developments of novel proxies are highlighted to illustrate advances in reconstructing millennial-scale disturbance-related dynamics and how this new information benefits the sustainable management of boreal forests in a rapidly changing climate.

Phyllosphere microbial communities have received considerable attention given their important influence on their plant hosts and on ecosystem functioning. In a context where climate change threatens the sustainability of ecosystems, it is important to understand how phyllosphere microbes will respond to changes in their environment. We used 16S rRNA gene amplicon sequencing to quantify phyllosphere bacterial communities of black spruce (Picea mariana (Mill.) B.S.P.) exposed to nitrogen canopy enrichment and soil warming in the boreal forest of Quebec, Canada. The treatments were applied from April to September 2015 and the sampling was done in September. Neither treatment influenced the overall community structure and diversity of black spruce phyllosphere bacterial communities. However, some bacterial taxa and inferred microbial functions did differ among treatments, revealing in particular a stronger response of some bacteria to soil warming rather than nitrogen enrichment. Our results suggest that soil warming could potentially induce more changes in phyllosphere bacterial taxa abundances and functions than could nitrogen addition, with potential consequences for microbial diversity and boreal forest ecosystem function under likely climate change scenarios. Our study suggests avenues for further research to integrate a more mechanistic understanding of the importance of phyllosphere microbes for black spruce and boreal forest ecosystems.

Despite growing interest in predicting plant phenological shifts, advanced spring phenology by global climate change remains debated. Evidence documenting either small or large advancement of spring phenology to rising temperature over the spatio-temporal scales implies a potential existence of a thermal threshold in the responses of forests to global warming. We collected a unique data set of xylem cell-wall-thickening onset dates in 20 coniferous species covering a broad mean annual temperature (MAT) gradient (−3.05 to 22.9°C) across the Northern Hemisphere (latitudes 23°–66° N). Along the MAT gradient, we identified a threshold temperature (using segmented regression) of 4.9 ± 1.1°C, above which the response of xylem phenology to rising temperatures significantly decline. This threshold separates the Northern Hemisphere conifers into cold and warm thermal niches, with MAT and spring forcing being the primary drivers for the onset dates (estimated by linear and Bayesian mixed-effect models), respectively. The identified thermal threshold should be integrated into the Earth-System-Models for a better understanding of spring phenology in response to global warming and an improved prediction of global climate-carbon feedbacks.

In boreal landscapes, emphasis is currently placed on close‐to‐nature management strategies, which aim to maintain the biodiversity and ecosystem services related to old‐growth forests. The success of these strategies, however, depends on an accurate understanding of the dynamics within these forests. While moderate‐severity disturbances have recently been recognized as important drivers of boreal forests, little is known about their effects on stand structure and growth. This study therefore aimed to reconstruct the disturbance and postdisturbance dynamics in boreal old‐growth forests that are driven by recurrent moderate‐severity disturbances. We studied eight primary old‐growth forests in Québec, Canada, that have recorded recurrent and moderately severe spruce budworm (Choristoneura fumiferana [Clem.]) outbreaks over the 20th century. We applied an innovative dendrochronological approach based on the combined study of growth patterns and releases to reconstruct stand disturbance and postdisturbance dynamics. We identified nine growth patterns; they represented trees differing in age, size, and canopy layer. These patterns highlighted the ability of suppressed trees to rapidly fill gaps created by moderate‐severity disturbances through a single and significant increase in radial growth and height. Trees that are unable to attain the canopy following the disturbance tend to remain in the lower canopy layers, even if subsequent disturbances create new gaps. This combination of a low stand height typical of boreal forests, periodic disturbances, and rapid canopy closure often resulted in stands constituted mainly of dominant and codominant trees, similar to even‐aged forests. Overall, this study underscored the resistance of boreal old‐growth forests owing to their capacity to withstand repeated moderate‐severity disturbances. Moreover, the combined study of growth patterns and growth release demonstrated the efficacy of such an approach for improving the understanding of the fine‐scale dynamics of natural forests. The results of this research will thus help develop silvicultural practices that approximate the moderate‐severity disturbance dynamics observed in primary and old‐growth boreal forests., The analysis of radial growth patterns highlights that boreal old‐growth forests are resistant to recurrent and moderate‐severity disturbances, as understory trees can rapidly fill the resulting gaps. However, trees that are unable to attain the canopy following the disturbance then tend to remain in the lower canopy layers, even if a new disturbance reopens the canopy. Therefore, reaching the canopy represents a once‐in‐a‐lifetime opportunity, where success or failure depends on a relatively short window of time.

Bud and leaf development are important phenological events and help in defining the growing period of trees. Canopy greenness derived from PhenoCam has been used to investigate leaf phenology. Ques...

The erosion of old-growth forests in boreal managed landscapes is a major issue currently faced by forest managers; however, resolving this problem requires accurate surveys. The intention of our study was to determine if historic operational aerial forest surveys accurately identified boreal old-growth forests in Quebec, Canada. We first compared stand successional stages (even-aged vs. old-growth) in two aerial surveys performed in 1968 (preindustrial aerial survey) and 2007 (modern aerial survey) on the same 2200 km2 territory. Second, we evaluated the accuracy of the modern aerial survey by comparing its results with those of 74 field plots sampled in the study territory between 2014 and 2016. The two aerial surveys differed significantly; 80.8% of the undisturbed stands that were identified as “old-growth” in the preindustrial survey were classified as “even-aged” in the modern survey, and 60% of the stands identified as “old-growth” by field sampling were also erroneously identified as “even-aged” by the modern aerial survey. The scarcity of obvious old-growth attributes in boreal old-growth forests, as well as poorly adapted modern aerial survey criteria (i.e., criteria requiring high vertical stratification and significant changes in tree species composition along forest succession), were the main factors explaining these errors. It is therefore likely that most of Quebec’s boreal old-growth forests are currently not recognized as such in forest inventories, challenging the efficacy of sustainable forest management policies.

Key messageThe intra-annual variability in stem size of marginal beech population from south-eastern distribution range of beech in Poland is higher within than between populations.AbstractOne of the key issues of the distribution of tree species is their ability to track environmental changes. European beech (Fagus sylvaticaL.) is a species highly sensitive to extreme climatic events, because of its high phenotypic plasticity. In this study, we aim to determine the variability in stem size between and within marginal beech populations. Marginal populations of beech growing under uniform environmental conditions of provenance trial offer unique opportunity to detect adaptive differentiations driven by natural selection. In this work, we studied stem size variation recorded by automatic band dendrometers in four beech marginal populations growing in a common garden in the south-eastern distribution range of beech in Poland over the period 2016–2018. Strong climatic effects and weak provenance differences in seasonal stem size variation were observed. The provenances exhibited similar climate-related seasonal stem circumference variation. A high within-provenance variation was confirmed. Temperature of spring as well as temperature and precipitation of autumn were detected as key climatic parameters mostly for onset and end of stem size variation. Maximum stem size was mostly affected by the later end of its variation, which positively affected its duration. Climatic distance between beech provenances and provenance trial had a negligible effect on the variability in seasonal stem size variation between provenances. The evidence of weak inter-provenance and high intra-provenance variation in stem size changes observed in the south-eastern distribution range indicates that an individual-based approach could be a suitable strategy, when selecting for phenotypic plasticity.

Spruce budworm (SBW – Choristoneura fumiferana Clem.) is one of the most damaging defoliating insects in the coniferous forests of eastern North America. In Canada, the widely distributed balsam fir (Abies balsamea L. (Mill)) and black spruce (Picea mariana B.S.P. (Mill)) are its most important hosts. Defoliation by SBW reduces growth in the host trees and can lead to host mortality. Although SBW impacts on growth are well documented, much less is known about changes in wood properties resulting from defoliation. To address this knowledge gap, we sampled 36 SBW-infested stands located in the boreal forest of Quebec (Canada) to determine whether defoliation modifies the wood quality of affected trees. The selected stands had been subjected to one to four years of SBW defoliation. For both species, we assessed ring growth, wood density, and the anatomical characteristics of stem wood formed during the outbreak years. We determined that rings formed during the SBW outbreak had a significant and progressive loss of biomass production with a longer duration of defoliation. SBW significantly reduced latewood density in the second and third year of defoliation for black spruce and the third and fourth year for balsam fir. Average ring density was reduced only in black spruce and only after four years of defoliation. The observed changes in growth and density were associated with changes in anatomical features. While the cellular characteristics of the earlywood remained fairly constant, significant reductions occurred in latewood cell-wall thickness after three years of defoliation. Our study shows that not only do SBW outbreaks reduce annual radial growth, but the cellular characteristics in latewood cells are also modified momentarily. Thus, SBW outbreaks affect wood density and quality in both black spruce and balsam fir.

Old‐growth forests are optimal habitats for many woodpeckers, which are often themselves excellent indicators of deadwood‐associated biodiversity. Old‐growth forests are, however, heterogeneous ecosystems in terms of structure, composition, and deadwood characteristics, thus implying a varied use of these forests by woodpeckers. In boreal landscapes, old‐growth stands are threatened by forest harvesting; however, there is little information in regard to the consequences for biodiversity with the loss of specific types of old‐growth forests. This study aimed to assess how the black‐backed woodpecker (Picoides arcticus), a biodiversity indicator species associated with old‐growth forest attributes, uses different types of old‐growth forests for its foraging needs. We identified woodpecker foraging marks in 24 boreal old‐growth forest stands in eastern Canada that were dominated by black spruce (Picea mariana), located within the home range of eight black‐backed woodpeckers. We identified the various old‐growth forest types using a typology based on the structural attributes of old‐growth stands. We classified the sampled stands into four old‐growth forest types, corresponding to different successional stages (recent or old, relative to the onset of the old‐growth stage), composition (pure black spruce or mixed black spruce–balsam fir [Abies balsamea]), and productivity (ongoing paludification or not). The black‐backed woodpecker foraged in all types of old‐growth forests, but favored dense old‐growth forests that were not paludified and that showed a high temporal continuity (i.e., old‐growth dynamics probably started more than a century ago). The temporal continuity of the old‐growth state allows for the continuous supply of large, slightly decayed snags, the preferred foraging substrates of the black‐backed woodpecker. The old‐growth forest type most favored by this woodpecker is, however, also the forest type most often targeted first by logging operations. Protecting the biodiversity associated with recent deadwood in managed areas thus requires maintaining a sufficient area and density of dense, old‐growth black spruce‐dominated forests in managed areas.

The current research on the dynamics of tree ring formation in conifers has provided new insights into how rate and duration xylem-cell production and development control the size of the xylem conduits leading to the formation of earlywood and latewood. So far, the physiology behind wood formation processes and the associated kinetics has rarely been considered, leading to the impossibility to grasp the drivers of wood density changes along the tree-rings. Despite the importance of wood density for carbon sequestration and tree hydraulics, little is known about the factors controlling variations in wood density across the tree ring, i.e. micro-density, at the intra-annual scale. We first developed a process-based mechanistic model that simulates the development of conifer tracheids from a simple sugar signal that we discuss together with the main kinetics and environmental variables leading to the formation of micro-density in black spruce, the main conifers species in the boreal forest of Canada. At the beginning of the growing season, low sugar availability in the cambium results in slow wall deposition that allows for a lengthier enlargement time thus producing large cells with thin walls (i.e. earlywood). In late summer and early autumn, high sugar availability produces narrower cells with thick cell walls (i.e. latewood). Wood formation dynamics had an indirect effect on micro-density. Micro-density increased under longer periods of cell wall deposition and shorter durations of enlargement. Cell diameter indirectly affected micro-density via cell wall thickness, which was the most important parameter affecting micro-density. Cell traits experienced the joint action of enlargement and secondary wall deposition in shaping the intra-annual patterns of tree rings. Our results point to the predictive power of a simple sugar signal. During the growing season, the amount of carbon allocated to wood formation largely influences the duration of cell differentiation, thus modulating cell diameter, cell wall thickness and by result tree-ring micro-density.

Wood density is the product of carbon allocation for structural growth and reflects the trade-off between mechanical support and water conductivity. We tested a conceptual framework based on the assumption that micro-density depends on direct and indirect relationships with endogenous and exogenous factors. The dynamics of wood formation, including timings and rates of cell division, cell enlargement, and secondary wall deposition, were assessed from microcores collected weekly between 2002 and 2016 from five black spruce stands located along a latitudinal gradient in Quebec, Canada. Cell anatomy and micro-density were recorded by anatomical analyses and X-ray measurements. Our structural equation model explained 80% of micro-density variation within the tree-ring with direct effects of wall thickness (σ = 0.61), cell diameter (σ = -0.51), and photoperiod (σ = -0.26). Wood formation dynamics had an indirect effect on micro-density. Micro-density increased under longer periods of cell-wall deposition and shorter durations of enlargement. Our results fill a critical gap in understanding the relationships underlying micro-density variation in conifers. We demonstrated that short-term responses to environmental variations could be overridden by plastic responses that modulate cell differentiation. Our results point to wood formation dynamics as a reliable predictor of carbon allocation in trees.

Spruce budworm (SBW) outbreaks are a major disturbance in North American forests. In Quebec, Canada, the recent history of SBW outbreaks at the local scale is well-known. Studies at the Québec scale nonetheless remain rare despite the need to better understand the dynamics of SBW outbreaks at a larger scale. This study aimed to reconstruct the spatiotemporal dynamics of SBW outbreaks during the 20th century across the insect's range in southern Quebec. To this end, we sampled 83 stands throughout southern Quebec. These stands were selected according to their age and the presence of black, white, and red spruce. In fact, spruce, unlike balsam fir, survives the SBW outbreaks and can record them in these growth rings. In each stand, cores were taken from 20 spruce trees. The dendrochronological series of more than 1,600 trees were analyzed, and we identified, through the k-means grouping of stands, the spatial patterns of tree growth for the three previously documented 20th century SBW outbreaks. The outbreaks were not homogeneous across the distribution range of the insect. Two groups of stands showed early- (1905–1930) and late-century (1968–1988) outbreaks of high severity and a mid-century (1935–1965) outbreak of moderate severity. This pattern is explained mainly by the presence of the insect within the balsam fir–yellow birch and balsam fir–white birch bioclimatic domains, areas where outbreaks tend to be most severe because of the abundance of balsam fir, the main SBW host species. However, these two models differ in terms of the duration of outbreaks. A third, more northern, cluster of stands experienced lower severity outbreaks over the 20th century, a pattern explained by a lower proportion of balsam fir trees in these landscapes. Our study shows that, on the one hand, these three groups of stands are defined by outbreaks of specific duration (an outbreak period beginning when more than 20% of the trees are affected and ending when

This chapter revisits the hypothesis put forward in 2007 that there is a close relationship between the structure (species composition and their distribution in the overstory) of mature boreal forest stands and the occurrence—as well as the periodicity, synchronicity, and amplitude—of insect outbreaks within the forest. New data from original dendroecological and paleoecological studies of spruce budworm outbreaks over the last 10 000 years strengthen this hypothesis. The variation in spatial and temporal dynamics of outbreaks is the reflection of the changes in forest structure driven by fire and insect outbreaks. There is a close inverse relationship between fire frequency and outbreak frequency over time. High fire frequency favors the presence of non-host species of the insect while low fire frequency favors the development of fire intolerant species such as balsam fir, the preferred host of the budworm, which influences outbreak frequency and impact. This would explain why the outbreaks of the 20th century were so widespread, even spreading further north due to climate warming.

Significance Forest trees can live for hundreds to thousands of years, and they play a critical role in mitigating global warming by fixing approximately 15% of anthropogenic CO2 emissions annually by wood formation. However, the environmental factors triggering wood formation onset in springtime and the cellular mechanisms underlying this onset remain poorly understood, since wood forms beneath the bark and is difficult to monitor. We report that the onset of wood formation in Northern Hemisphere conifers is driven primarily by photoperiod and mean annual temperature. Understanding the unique relationships between exogenous factors and wood formation could aid in predicting how forest ecosystems respond and adapt to climate warming, while improving the assessment of long-term and high-resolution observations of global biogeochemical cycles., Wood formation consumes around 15% of the anthropogenic CO2 emissions per year and plays a critical role in long-term sequestration of carbon on Earth. However, the exogenous factors driving wood formation onset and the underlying cellular mechanisms are still poorly understood and quantified, and this hampers an effective assessment of terrestrial forest productivity and carbon budget under global warming. Here, we used an extensive collection of unique datasets of weekly xylem tissue formation (wood formation) from 21 coniferous species across the Northern Hemisphere (latitudes 23 to 67°N) to present a quantitative demonstration that the onset of wood formation in Northern Hemisphere conifers is primarily driven by photoperiod and mean annual temperature (MAT), and only secondarily by spring forcing, winter chilling, and moisture availability. Photoperiod interacts with MAT and plays the dominant role in regulating the onset of secondary meristem growth, contrary to its as-yet-unquantified role in affecting the springtime phenology of primary meristems. The unique relationships between exogenous factors and wood formation could help to predict how forest ecosystems respond and adapt to climate warming and could provide a better understanding of the feedback occurring between vegetation and climate that is mediated by phenology. Our study quantifies the role of major environmental drivers for incorporation into state-of-the-art Earth system models (ESMs), thereby providing an improved assessment of long-term and high-resolution observations of biogeochemical cycles across terrestrial biomes.

In boreal landscapes, emphasis is currently placed on close-to-nature management strategies, which aim to maintain the biodiversity and ecosystem services related to old-growth forests. The success of these strategies, however, depends on an accurate understanding of the dynamics within these forests. This study aims to reconstruct the disturbance and post-disturbance dynamics in boreal old-growth forests that are driven by recurrent moderate-severity disturbances. We studied eight old-growth forests in Quebec, Canada, that has recorded recurrent and moderate to severe spruce budworm (Choristoneura fumiferana [Clem.]) outbreaks over the 20th century. To reconstruct the disturbance history and the post-disturbance dynamics of the study sites, we used dendrochronological data and k-means clustering to identify growth releases and growth patterns in the studied trees. We identified nine growth patterns; these patterns represented trees differing in age, size, and canopy layer, and indicated different tree histories. Spruce budworm outbreaks caused recurrent moderate-severity disturbances within the study sites. The canopy gaps created by these disturbances were filled mainly by understorey trees, which responded by single and significant increases in radial growth and height. In contrast, overstorey trees had little influence on gap filling; thus, trees were mostly from the dominant and codominant canopy layers. Our study underlines the resistance of boreal old-growth forests to recurrent and moderate-severity disturbances, as understorey trees can rapidly fill the resulting gaps. However, trees that are unable to attain the canopy following the disturbance then tend to remain in the lower canopy layers. Therefore, reaching the canopy represents a once-in-a-lifetime opportunity, where success or failure depends on a relatively short window of time. This gap-filling dynamic produces, however, a vertical structure that is often similar to the expected structure in even-aged forests. Our results highlight the efficacy of identifying tree growth patterns to reconstruct stand disturbance dynamics and contribute to developing closer to nature forest management strategies.

New insights into the intra-annual dynamics of tree-ring formation can improve our understanding of tree-growth response to environmental conditions at high-resolution time scales. Obtaining this information requires, however, a weekly monitoring of wood formation, sampling that is extremely time-intensive and scarcely feasible over vast areas. Estimating the timing of cambial and xylem differentiation by modeling thus represents an interesting alternative for obtaining this important information by other means. Temporal dynamics of cambial divisions can be extracted from the daily tree-ring growth rate computed by the Vaganov–Shashkin (VS) simulation model, assuming that cell production is tightly linked to tree-ring growth. Nonetheless, these predictions have yet to be compared with direct observations of wood development, i.e., via microcoring, over a long time span. We tested the performance of the VS model by comparing the observed and predicted timing of wood formation in black spruce [Picea mariana (Mill.)]. We obtained microcores over 15 years at 5 sites along a latitudinal gradient in Quebec (Canada). The measured variables included cell size and the timing of cell production and differentiation. We calibrated the VS model using daily temperature and precipitation recorded by weather stations located on each site. The predicted and observed timing of cambial and enlarging cells were highly correlated (R 2 = 0.8); nonetheless, we detected a systematic overestimation in the predicted timing of cambial cells, with predictions delayed by 1–20 days compared with observations. The growth rate of cell diameter was correlated with the predicted growth rate assigned to each cambial cell, confirming that cell diameter developmental dynamics have the potential to be inferred by the tree-ring growth curve of the VS model. Model performances decrease substantially in estimating the end of wood formation. The systematic errors suggest that the actual relationships implemented in the model are unable to explain the phenological events in autumn. The mismatch between the observed and predicted timing of wood formation in black spruce within our study area can be reduced by better adapting the VS model to wet sites, a context for which this model has been rarely used.

Research Highlights: Radial growth patterns of trees growing in old-growth boreal forests in eastern Canada can be grouped into a small number of simple patterns that are specific to different old-growth forest types or successional stages. Background and Objectives: Identifying the main radial growth trends in old-growth forests could help to develop silvicultural treatments that mimic the complex dynamics of old-growth forests. Therefore, this study aimed to identify the main radial growth patterns and determine how their frequencies change during forest succession in old-growth forests, focusing on boreal landscapes in eastern Canada. Materials and Methods: We used dendrochronological data sampled from 21 old-growth stands in the province of Quebec, Canada. Tree-ring chronologies were simplified into chronologies of equal length to retain only primary growth trends. We used k-means clustering to identify individual growth patterns and the difference in growth-pattern frequency within the studied stands. We then used non-parametric analyses of variance to compare tree or stand characteristics among the clusters. Results: We identified six different growth patterns corresponding to four old-growth forest types, from stands at the canopy breakup stage to true old-growth stands (i.e., when all the pioneer cohort had disappeared). Secondary disturbances of low or moderate severity drove these growth patterns. Overall, the growth patterns were relatively simple and could be generally separated into two main phases (e.g., a phase of limited radial increment size due to juvenile suppression and a phase of increased radial increment size following a growth release). Conclusions: The complexity of old-growth forest dynamics was observed mainly at the stand level, not at the tree level. The growth patterns observed in true old-growth forests were similar to those observed following partial or stem-selection cuts in boreal stands, thus, these silvicultural treatments may be effective in mimicking old-growth dynamics.

Old-growth forests play a major role in conserving biodiversity, protecting water resources, sequestrating carbon, and these forests are indispensable resources for indigenous societies. To preserve the ecosystem services provided by these boreal ecosystems, it becomes necessary to develop novel silvicultural practices capable of emulating the natural dynamics and structural attributes of old-growth forests. The success of these forest management strategies depends on developing an accurate understanding of natural regeneration dynamics. Our goal was therefore to identify the main patterns and the drivers involved in the regeneration dynamics of old-growth forests, placing our focus on boreal stands dominated by black spruce (Picea mariana(L.) Mill.) and balsam fir (Balsam fir(L.) Mill.) in eastern Canada. We sampled 71 stands in a 2200 km2study area located within Quebec’s boreal region. For each stand, we noted tree regeneration (seedlings and saplings), structural attributes (diameter distribution, deadwood volume, etc.), and abiotic (topography and soil) factors. We observed that secondary disturbance regimes and topographic constraints were the main drivers of balsam fir and black spruce regeneration. Furthermore, the regeneration dynamics of black spruce appeared more complex than those of balsam fir. We observed distinct phases of seedling production first developing within the understory, then seedling growth when gaps opened in the canopy, followed by progressive canopy closure. Seedling density, rather than the sapling density, had a major role in explaining the ability of black spruce to fill the canopy following a secondary disturbance. The density of balsam fir seedlings and saplings was also linked to the abundance of balsam fir trees at the stand level. This research helps explain the complexity of old-growth forest dynamics where many ecological factors interact at multiple temporal and spatial scales. This study also improves our understanding of ecological processes within native old-growth forests and identifies the key factors to consider when ensuring the sustainable management of old-growth boreal stands.

International audience; The spatial and temporal distributions of plant hormones during tree-ring formation are beginning to be elucidated. Plant hormones play specific roles in tree-ring formation depending on the developmental zone in which they operate. Tree rings are the result of the seasonal activity of the vascular cambium, the secondary meristem of woody angiosperms and gymnosperms. During tree-ring formation, a combination of endogenous and environmental factors affects cambial division, cell differentiation and maturation; this, in turn, affects wood quantity and quality. Among the endogenous factors affecting tree-ring formation, plant hormones are recognized as determinant players in regulating many aspects of the features and fate of each xylem cell. Most of our knowledge regarding the roles of plant hormones on tree-ring formation comes from herbaceous plants, although an increasing interest involves the analysis of the hormonal patterns in tree-species. This paper reviews the state of knowledge of the role of plant hormones during tree-ring formation by focusing on experiments performed on woody species. An overview of the main plant hormones and their main activities during radial tree growth will be followed by discussion of their role in each tree-ring developmental stage and in the overall tree-ring seasonal pattern.

Old-growth stands are considered as key components of boreal forest diversity and their preservation is largely integrated into management plans. However, while the differences between old-growth and young forests have largely been studied, little is known about the diversity of boreal old-growth forests. In managed landscapes, the efficacy of old-growth conservation plans may be reduced depending on how these old-growth forests are considered: as a single, homogeneous and steady-state forest type or as multiple, diverse and dynamic forest types. To fulfil this gap, our objectives were: (1) to create a typology of old-growth boreal structures; (2) to observe how these structures are influenced by environmental and temporal parameters; and (3) to elaborate a succession model of old-growth structural dynamics along temporal and environmental gradients. Seventy-one mature and overmature stands were sampled within a 2200 km2 territory situated in Eastern Canada. Cluster analysis divided the sampled stands into two even-aged types, three transition old-growth types and six true old-growth types. Slope, minimum time since last fire and organic horizon depth were the three environmental and temporal parameters influencing the old-growth structures. Paludification-related productivity decline was present in only one old-growth forest type, while the other sites remained productive. These results allowed the creation of three succession models of the dynamics of old-growth stands in the boreal forest of eastern Canada. Boreal stands can undergo numerous structural changes once the old-growth succession process is initiated. An increase in structural diversity when the true old-growth stage is reached, coupled with a variety of secondary disturbance characteristics, favours multiple pathways of structural evolution of these ecosystems over time. Therefore, forest management planning should incorporate this complexity to improve the preservation of old-growth forests in managed territories.

Natural disturbances have a major impact on boreal forest landscape dynamics, and although fire history is well documented at the Holocene scale, spruce budworm (Choristoneura fumiferana (Clemens)) (SBW) dynamics have only been known for the last three centuries. This is likely due to the difficulty in using and interpreting existing indicators (cephalic head capsules and feces). In this methodological study, we present an original approach using lepidopteran wing scales to reconstruct insect abundance. We analyzed two sediment cores from the boreal forest in central Quebec and extracted wing scales at every stratigraphic level. The required quantity of sediment for paleoecological analysis is relatively small given the large quantity of wing scales produced by Lepidoptera and their small size. Scales are well preserved due to their chitinous structure and their great variety of shapes offer a high potential for taxonomic identification. A statistical model based on the shape of scales of the three major epidemic lepidopterans in Quebec discriminated 68% of SBW scales. This indicator allows a more efficient and more precise reconstruction of SBW history with respect to the use of cephalic head capsules or feces.

Tree-related microhabitats (TreM) and deadwood are two forest attributes providing essential resources for biodiversity conservation and ecosystem services. Old-growth forests are generally defined by a high abundance and diversity of TreM and deadwood, but little is known about TreM and deadwood dynamics once the old-growth stage is reached, in particular in the boreal biome. In this context, knowledge on TreM and deadwood dynamics in old-growth forest stands is necessary to better understand how these forests contribute to biodiversity and ecosystem services. The aim of this study is thus to determine how TreM, and deadwood abundance and diversity vary within boreal old-growth forests. To reach this objective, we surveyed TreM and deadwood attributes, as well as structural and abiotic attributes, in 71 boreal old-growth forests situated in Quebec, Canada. We used hierarchical clustering analysis to identify TreM and deadwood abundance and diversity patterns in the studied stands. We identified five clusters of TreM and deadwood characteristics, which corresponded to three stages of old-growth forest succession: canopy break-up (beginning of the old-growth stage), transition old-growth stage (replacement of the first cohort by old-growth cohorts) and true old-growth stage (first cohort all or almost all gone). The peak in TreM richness and diversity was reached at the transition old-growth stage, whereas the peak for deadwood richness and diversity was reached at the true old-growth stage. Overall, true old-growth forests were defined by a combination of moderate to high TreM density and high deadwood volume, but these values significantly varied among stands depending on past secondary disturbances, stand structure and its composition (black spruce [Picea mariana Mill.] dominated vs mixed black spruce – balsam fir [Abies balsamea (L.) Mill.]). These results therefore underscore the importance of considering old-growth forests as dynamic rather than static ecosystems, as the composition of tree- and deadwood-related microhabitats in the same old-growth stand may markedly change over time. At landscape scale, these results also imply that the mosaic of habitats present in old-growth forests can vary greatly from one location to another, highlighting the importance of maintaining a diversity of old-growth forest structure and composition.

There is a growing need for understanding the timing of wood formation in relation to the dynamics of bud phenology at wide geographical scale. This study analysed the relationships between long-term chronologies of xylem growth and the timing of plant phenology detected by Normalized Difference Vegetation Index (NDVI) in five permanent stands across the latitudinal distribution of black spruce [ Picea mariana (Mill.) BPS] in the boreal forest of Quebec, Canada. Xylogenesis was studied weekly from April to October for thirteen years (2002–2014) on anatomical sections derived by wood microcores. The timing of the growing season detected by remote sensing was extracted from MODerate resolution Imaging Spectroradiometer (MODIS) 250 m 16-days NDVI data. The NDVI time-series were fitted using a double-logistic curve. Phenological chronologies from remote sensing and xylem phenology showed a latitudinal trend. The models correlating the data inferred from satellite sensors and the spring observations of xylem phenology were significant ( p