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

 

Permanent landscape attributes such as topography (elevation [m]) and microtopography (local variation in elevation [cm]) can increase the risk of cold air drainage down-slopes and in microtopographic depressions, causing important temperature gradients that generate localized growing-season frosts. Since most studies on growing-season frosts are restricted to the northern parts of the boreal forest or to mountainous areas, their negative consequences on tree productivity at the boreal-temperate forest ecotone is often ignored. We quantified the intensity and probability of growing-season frosts at the boreal-temperate forest ecotone in regard to topographic and microtopographic landscape attributes, which were extracted from airborne LiDAR surveys. In situ air temperature was measured for two summers (2016–2017) with 252 temperature loggers installed in two 18-years-old spruce plantations established in both temperate and boreal mixedwood forests. Growing-season frosts were more intense and probable at the boreal mixedwood forest site compared to the temperate forest site. Still, at both sites, when growing-season frost occurred, air temperature could vary by 4 °C along the elevation gradient of 15 m, often reaching sub-zero values at low elevation while reaching above-freezing values at high elevation. The importance of microtopography on the risk of sub-zero temperatures increased where frost events were less likely to occur such as at the temperate forest and at high elevation. Considering that growing-season frosts can considerably reduce tree productivity, the effects of both topography and planting microsite should be considered when determining where to establish plantations in the landscape and to determine suitable frost-free planting microsites within a plantation.

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.

Increase in frost damage to trees due to earlier spring dehardening could outweigh the expected increase in forest productivity caused by climate warming. We quantified the impact of growing?season frosts on the performance of three spruce species (white, black, and Norway spruce) and various seed sources with different frost tolerance in two plantations, established on both sides of the eastern Canadian boreal?temperate forest ecotone. The objectives of this study were to determine (a) if spruce species and seed sources planted in sites far from their natural provenance would be less adapted to local site conditions, leading to increased frost damage and reduced height growth; (b) at which height above the ground growing?season frosts ceased to damage apical meristems; and (c) if height growth was best predicted by extreme climatic events (growing?season frosts) or by mean annual or summer temperature. At each site and for all spruce species and seed sources, we cross?sectioned spruce trees at different heights above the ground. Tree rings were cross?dated and screened for frost rings, which were then given a severity score based on cellular damage. Frost severity reduced height growth of all spruce species and provenances at both sites. Height growth of the non?native Norway spruce was the most reduced by frost severity and was the smallest species at both sites. Frost caused the highest growth reduction in white spruce at the boreal mixedwood site and had the least effect on black spruce at both sites. For all spruce species, height growth was affected up to 2 m above the ground. Model selection based on corrected Akaike's information criteria (AICc) identified that minimum temperature in May was by far the best climate variable predicting tree growth (AICc weight = 1), highlighting the importance of considering extreme climatic events, which are likely to increase in the future.

Climate warming-driven early leaf-out is expected to increase forest productivity but concurrently increases leaf exposure to spring frosts, which could reduce forests' net productivity. We hypothesized that due to their damaging effect on buds, spring frosts exert a stronger control on bud phenology than do growing degree-days. We monitored bud flush phenology of three white spruce seed sources (one local seed source from the boreal mixedwood forest and two seed sources from the temperate forest), one black spruce seed source originating from the boreal mixedwood forest and four nonlocal Norway spruce seed sources in 2016 and 2017 in two plantations located on both sides of the temperate-boreal mixedwood forest ecotone in eastern Canada (Quebec). We aimed to determine inter- and intraspecies variations in bud break timing and sensitivity to air temperature and photoperiod. We expected that bud break timing for boreal species and seed sources would be better synchronized with the decrease in frost probability than for nonlocal species and seed sources. We used mixed binomial regressions and AICc model selection to determine the best environmental variables predicting each transition from one stage of bud phenology to the next. At both plantation sites, white spruce bud flush began and ended earlier compared to black and Norway spruce. Buds of all spruce species were sensitive to frost probability for early phenological stages, whereas growing degree-days controlled the remaining stages. Photoperiod sensitivity was higher for white spruce compared to black and Norway spruce and reached its maximum in the temperate forest. At intraspecies level, the two southern white spruce seed sources opened their buds earlier than the local source and were more sensitive to photoperiod, which increased their exposure to spring frosts. Onset of spruce bud flush is driven by spring frosts and photoperiod, but once started, bud phenology responds to temperature. The high photoperiod sensitivity in white spruces could counterbalance climate warming and limit future premature leaf-out, whereas the low photoperiod sensitivity in black spruce should not restrain leaf-out advancement with climate warming. Our results call for adapting the temperature-driven hypotheses of ecophysiological models predicting leaf-out to include spring frost probability.

Although advance regeneration abundance and vigor are critical factors determining future forest composition and productivity, very few studies have focused on how they are affected by spruce budworm (SBW) outbreaks even though they affect millions of hectares of boreal forest on a cyclical basis. Post-SBW salvage logging is often used to reduce economic losses but could interact with the outbreak to affect advance regeneration. This study aims to determine the impact of SBW outbreaks and post-outbreak salvage logging on the defoliation of advance regeneration in mixed coniferous stands of northeastern Canada. Specifically, we assessed the effect of regeneration height and species (balsam fir or black spruce), as well as canopy composition, on the defoliation of advance regeneration. We then evaluated the effect of salvage logging on defoliation sustained by advance regeneration and compared it to the one observed in stands only affected by the SBW. Regeneration height and species, canopy composition and salvage logging all significantly affected defoliation and showed multiple interactions. Taller balsam fir seedlings were three times as defoliated as smaller ones, whereas it was 2.3 times for black spruce. Balsam fir seedlings were 15% more defoliated than black spruce. Seedlings of both species located beneath a balsam fir canopy were also more defoliated (>50% defoliation) than seedlings found under black spruce trees (about 26% defoliation). Salvage logging in black spruce-dominated stands resulted in a ?25% increase in defoliation of tall (2.5?m) black spruce regeneration when compared to non-harvested sites. We speculate that this could increase the fir component in spruce-dominated stands, leading to forests that are more susceptible to future SBW outbreaks. To protect spruce advance regeneration from increased defoliation, salvage harvesting of spruce-dominated stands may thus be delayed until the outbreak has subsided. Long-term studies are needed to determine whether a compositional change occurs or not, particularly in spruce-dominated stands. As a precautionary measure, changes in salvage logging practices may be implemented immediately to avoid potential problems such as decreased black spruce abundance and increased susceptibility to future SBW outbreaks.

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

Wildfires are a major driver of carbon stocks and ecosystem development in Canadian boreal forests, but there is little information on amounts and properties of the charcoal produced. Using data and samples available from a previous study, we determined amounts, depth distribution and chemical properties of visually-determined charcoal (>2 mm) in a boreal chronosequence in the Abitibi region of Quebec, Canada. Sites ranged from 24 to 2,355 years since fire (ysf) and originated from low- and high-severity soil burns (>5 or <5 cm organic horizon unburned, respectively). Two or three pits were sampled at 1-cm depth intervals from 20 jack pine (Pinus banksiana) sites (one low severity and 19 high severity) and 31 black spruce (Picea mariana) sites (12 low severity and 19 high severity). Site-level charcoal stocks ranged from 50 to 5,527 kg ha?1 with high within-site variability and lower stocks for the oldest sites. Depth distributions typically peaked around the organic-mineral interface, but some low-severity sites also had charcoal layers within the organic horizon. Means from 30 charcoal samples were 569 mg g?1 total C, 4.1 mg g?1 total N and 140 C/N (molar), with total C and C/N showing a trend of decline with time since fire, and total N showing an increase. Solid-state 13C CPMAS NMR spectra of nine samples showed high variability among the younger samples, but a trend to higher aromaticity for the older ones. A literature survey focusing on boreal forests similarly showed highly variable stocks and chemical properties of charcoal in organic horizon and upper mineral soil, with reduction of variance and lower stocks after several hundred years. This initial variation was also consistent with reports of highly variable temperatures and duration of charring in wildfires. Adding reports available for char production, and considering that most studies of char stocks and production are limited to the organic horizon (forest floor), suggests that initial production of charred material from boreal wildfires might be around 5–10 tons ha?1.