In boreal ecosystems, phenological events display seasonal patterns. These patterns allow for the development of tissues during the short time window available for growth in cold climates. Primary and secondary growth, two expensive processes for plants, are supposedly modulated in time to optimize allocation of carbon to bud and woody tissues. We aimed to assess the phenology of primary and secondary meristems, testing their relationship over the closed black spruce stands of the commercially exploited forest region in Quebec, Canada.

Location

Quebec, Canada.

Time period

2002–2016.

Major taxa studied

Gymnospermae.

Methods

We combined weekly scaled field observations with Moderate Resolution Imaging Spectroradiometer (MODIS) time series of the normalized difference vegetation index (NDVI) to extract timings of photosynthesis and meristem growth in five black spruce [Picea mariana (Mill.) B.S.P.] stands located along a latitudinal gradient and to assess their relationship. We then tested empirical models based on geographical position and seasonal temperatures to predict wood phenology (i.e., the onset and ending of earlywood and latewood formation), and compared its spatial patterns with existing predictions of bud phenology for the same study area.

Results

Photosynthesis started at the beginning of May, 3 weeks before bud reactivation and the onset of wood growth. Latewood formation started in mid-July, after shoot elongation was completed. For wood phenology models, the residual standard error ranged from 1 week to 12 days. Growth dynamics spatialized across the boreal forest of Quebec varied with the transition between the subarctic and humid continental climate.

Main conclusions

Shoot elongation and latewood formation were temporally separated, providing evidence of a trade-off in structural carbon allocation between primary and secondary growth in trees. Spatial patterns of wood phenology predicted for the black spruce polygons are consistent with spatial patterns of bud phenology, demonstrating synchronized temporal dynamics of meristems at the regional scale.

Abstract 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.

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.

Spruce budworm (Choristoneura fumiferana (Clem)) is the main defoliator in the boreal forest of North America, and its outbreaks have major ecological and economic consequences and represent a challenge for forest management. Numerous studies have addressed the effects of this defoliator on mature trees, whereas the effects of spruce budworm on regeneration remain elusive. Furthermore, intensive exploitation practices during the last decades have left a large area of the Canadian boreal forest in an early development stage. In this context, it becomes vital to understand those factors affecting the severity of spruce budworm‐related defoliation on regeneration. Here, we determine the defoliation severity of black spruce and balsam fir seedlings in both mature pure black spruce and black spruce–balsam fir stands subjected to two different silvicultural treatments (clear‐cutting and partial cutting). Defoliation intensity varied between stand types, silvicultural treatments, species, and height classes. Seedlings in black spruce–balsam fir stands experienced twice the defoliation of those in pure black spruce stands (black spruce seedlings 10% vs. 23%; balsam fir seedlings 29% vs. 47%, respectively). Harvesting methods also influenced seedling defoliation. Under clear‐cutting, black spruce seedlings (24%) were three times as defoliated as black spruce seedlings in partial cutting stands (8%), whereas balsam fir seedlings in clear‐cutting plots experienced twice the defoliation (42%) of balsam fir seedlings in partial cutting plots (20%). The level of defoliation also increased with seedling height. This study will help silvicultural strategies adapt to the effects of natural disturbance regimes. As the intensity and severity of defoliator outbreaks are expected to increase under climate change, these results will help guide forest management strategies to select harvesting methods that will limit the effects of defoliation on conifer regeneration.

We investigated whether stand species mixture can attenuate the vulnerability of eastern Canada’s boreal forests to climate change and insect epidemics. For this, we focused on two dominant boreal species, black spruce [Picea mariana (Mill.) BSP] and trembling aspen (Populus tremuloides Michx.), in stands dominated by black spruce or trembling aspen (“pure stands”), and mixed stands (M) composed of both species within a 36 km2 study area in the Nord-du-Québec region. For each species in each stand composition type, we tested climate-growth relations and assessed the impacts on growth by recorded insect epidemics of a black spruce defoliator, the spruce budworm (SBW) [Choristoneura fumiferana (Clem.)], and a trembling aspen defoliator, the forest tent caterpillar (FTC; Malacosoma disstria Hübn.). We implemented linear models in a Bayesian framework to explain baseline and long-term trends in tree growth for each species according to stand composition type and to differentiate the influences of climate and insect epidemics on tree growth. Overall, we found climate vulnerability was lower for black spruce in mixed stands than in pure stands, while trembling aspen was less sensitive to climate than spruce, and aspen did not present differences in responses based on stand mixture. We did not find any reduction of vulnerability for mixed stands to insect epidemics in the host species, but the non-host species in mixed stands could respond positively to epidemics affecting the host species, thus contributing to stabilize ecosystem-scale growth over time. Our findings partially support boreal forest management strategies including stand species mixture to foster forests that are resilient to climate change and insect epidemics.

Large primary forest residuals can still be found in boreal landscapes. Their areas are however shrinking rapidly due to anthropogenic activities, in particular industrial-scale forestry. The impacts of logging activities on primary boreal forests may also strongly differ from those of wildfires, the dominant stand-replacing natural disturbance in these forests. Since industrial-scale forestry is driven by economic motives, there is a risk that stands of higher economic value will be primarily harvested, thus threatening habitats, and functions related to these forests. Hence, the objective of this study was to identify the main attributes differentiating burned and logged stands prior to disturbance in boreal forests. The study territory lies in the coniferous and closed-canopy boreal forest in Québec, Canada, where industrial-scale logging and wildfire are the two main stand-replacing disturbances. Based on Québec government inventories of primary forests, we identified 427 transects containing about 5.5 circular field plots/transect that were burned or logged shortly after being surveyed, between 1985 and 2016. Comparative analysis of the main structural and environmental attributes of these transects highlighted the strong divergence in the impact of fire and harvesting on primary boreal forests. Overall, logging activities mainly harvested forests with the highest economic value, while most burned stands were low to moderately productive or recently disturbed. These results raise concerns about the resistance and resilience of remnant primary forests within managed areas, particularly in a context of disturbance amplification due to climate change. Moreover, the majority of the stands studied were old-growth forests, characterized by a high ecological value but also highly threatened by anthropogenic disturbances. A loss in the diversity and functionality of primary forests, and particularly the old-growth forests, therefore adds to the current issues related to these ecosystems. Since 2013, the study area is under ecosystem-based management, which implies that there have been marked changes in forestry practices. Complementary research will be necessary to assess the capacity of ecosystem-based management to address the challenges identified in our study.

Climate change is projected to intensify the global hydrological cycle and increase temperatures. Excess N deposition could limit plant productivity because of a lack of plant–soil system efficiency with reduced water availability. However, the effects of soil warming and N?addition on tree water transport and water use remain poorly understood. This study aimed to quantify the effects of soil warming and N?addition associated with meteorological variables on the sap flux density and stem radius variation in black spruce (Picea mariana [Mill.] B.S.P.). From 2008 to 2014, we conducted a long?term experiment on mature trees growing in two stands (Bernatchez [BER] and Simoncouche [SIM]) at different latitudes to evaluate the effect of soil warming (H) and N?addition. During 2014, we investigated soil and plant water status between May and August, sap flux density from July to August and stem radius variation over the whole year. At the colder, northern site (BER), we detected a significant increase in sap flux density with the H?treatment, whereas no effect was observed at the warmer, southern site (SIM). At BER, diurnal sap flux density in H × N?addition treatment showed a significant effect in the afternoon near the peak of day compared with the other treatments. We found no significant effects of independent single or combined factors on soil and plant water status and stem radius variation at either site. Under climate change, black spruce at more northern sites could have a better water use and storage for wood productivity than those at more southern sites.

L’érosion des superficies des vieilles forêts boréales est actuellement l’un des enjeux majeurs de l’aménagement forestier; cependant, la résolution de ce problème nécessite des inventaires précises. Ainsi, l’objectif de cette étude était de déterminer si les précédents inventaires forestiers aériens identifiaient correctement les vieilles forêts dans les paysages boréaux du Québec, au Canada. Nous avons comparé les stades de succession (forêt équienne ou vieille forêt) de deux inventaires aériens réalisés en 1968 (inventaire aérien préindustriel) et en 2007 (inventaire aérien moderne) sur un territoire de 2200 km2. Nous avons aussi comparé les résultats de l’inventaire aérien moderne avec ceux obtenus à partir de 74 placettes de terrain échantillonnées entre 2014 et 2016. Les deux inventaires aériens étaient très incohérents : 80,8 % des peuplements non-perturbés identifiés comme « vieilles forêts » par l’inventaire préindustriel étaient classés comme « équiennes » par l’inventaire moderne et 60 % des placettes de terrain identifiées comme « vieilles forêts » étaient aussi classées « équiennes » par l’inventaire aérien moderne. Le manque d’attributs de vieilles forêts évidents ainsi que l’utilisation de critères inadaptés (c.-à-d. nécessitant une forte complexité verticale et d’importants changements de composition en espèces arborescentes durant la succession forestière) étaient les principaux éléments expliquant ces erreurs. Il est ainsi possible que la majorité des vieilles forêts boréales du Québec ne soient pas identifiées comme telles, limitant l’efficacité des stratégies de gestion durable.

The impact of traditional even-aged forest management on landscape age structure, tree composition, and connectivity has been well documented. Very little, however, is known about the impact on stand structural diversity. This study aims to compare the structural and abiotic characteristics of forest stands disturbed by clearcut logging and by stand-replacing fire in Quebec’s boreal landscapes. We hypothesized that unlike fire, logging specifically targeted stands having a higher economic value, i.e., merchantable volume, leaving altered forest characteristics on post-harvested landscapes. We compared two aerial forest surveys of a 2200 km2 study area, one survey completed before any logging activity (preindustrial survey; 1980s), and the second survey collected >10 years after logging activity (modern survey; 2000s). Forest stands at the time of the preindustrial survey were primary forests. We identified stands as either burned, logged, or left aside after forest management of the area (remaining stands) between the two surveys and compared their structural and abiotic characteristics using logistic regression. The structural and abiotic characteristics of burned and logged stands differed significantly. Relative to the burned stands, logged stands were older, denser, and marked by poorer drainage and a higher proportion of black spruce; therefore post-harvest and post-burn landscapes differed in terms of their structural diversities. Traditional even-aged forest management has significantly altered the boreal forest landscape by targeting specific stands having higher economic value and leaving behind stands of lower economic value. Remaining high economic stands should be protected, and a more balanced approach to harvesting must be used in the context of ecosystem-based management.