Abstract In forested ecosystems, shrubs must succeed in persisting in low-light environments, while simultaneously having the ability to rapidly expand and occupy newly created canopy openings, yet little is known about the traits that make this possible. We hypothesize that shrub species that are abundant in the understory exhibit a specific set of functional traits that define their ability to persist during unfavorable periods and to rapidly exploit newly created habitats. We tested this by comparing field-measured functional traits such as biomass allocation, leaf display, crown morphology, and leaf traits, across individual size classes and two gap-forest environments of five shrub species. We observed significant differences in traits between species, size classes, and gap-forest environments. These differences were primarily related to biomass allocation traits, followed by leaf display, crown morphology, and leaf traits. Abundant shrubs like mountain maple (Acer spicatum) and hazelnut (Corylus cornuta) invested significantly more biomass in roots, had a larger total leaf area, and displayed leaves in a more efficient manner to intercept light. The high investment in root biomass can be interpreted as shrubs exploiting the persistence and colonization strategy through resprouting. Permanent sub-canopy status likely explains the importance of efficient leaf display, wherein abundant shrubs had a large leaf area with minimal support structures.

The stand-level risk of both defoliation and mortality due to the spruce budworm (Choristoneura fumiferana, Clem., SBW) is mainly linked to the quality of the resource such that stands dominated by the primary host species, balsam fir (Abies balsamea [L.] Mill.), are considered the most vulnerable. At the landscape scale, insect dispersal suggests that both configuration and resource availability should influence the onset and subsequent mortality of an outbreak. However, outbreak risk at this scale is yet to be quantified for the SBW, one of the most studied forest pest defoliators in the world. We aim to determine the risk of both SBW defoliation onset and resultant tree mortality at a landscape scale due to factors that have been associated with SBW defoliation at the stand scale. We used Cox proportional hazard models to quantify the relative risk of different landscape configuration and species composition patterns on defoliation onset and risk of mortality of the current SBW outbreak in Eastern Canada. In particular, we measured the risk associated with primary host proportion, relative amounts of secondary host and hardwood trees to fir trees, the structure of balsam fir in the landscape, and the structure of the forest overall. We show that defoliation onset is more related to the configuration of the landscape than to the species composition while the risk of ensuing mortality increases as the proportion of the primary host increases. Defoliation onset risk is increased by a more complex configuration of fir patches in the landscape. Mortality risk, however, is unaffected by fir configuration but is reduced where there is more black spruce (Picea mariana [Mill.] B.S.P.) or a decrease of balsam fir in the landscape. The usefulness of survival analysis for assessing risk for insect outbreaks has been overlooked and can quantify the effect of relevant factors to guide management and mitigation strategies. The results of our analysis suggest that forest management strategies should avoid clear cuts and the development of fir monocultures and instead maintain or increase the proportion of black spruce and improve forest connectivity across large forested landscapes to delay defoliation onset and reduce SBW-related mortality.

Natural disturbances drive forest dynamics and biodiversity at different spatial and temporal scales. Forests in the boreal biome are shaped by several types of disturbance, including fire, windthrow, and insect outbreaks, that vary in frequency, extent, severity, and specificity. In managed forests, disturbances also affect the amount and quality of available timber. Ecosystem management uses information on disturbance regimes as a guide to finding a balance between ecological, economic, and social viewpoints. In this chapter, we review current knowledge on disturbance regimes in boreal forests and discuss some implications for managing the impact and risk of disturbances in the context of forest ecosystem management and restoration.

As global climate conditions continue to change, disturbance regimes and environmental drivers will continue to shift, impacting global vegetation dynamics. Following a period of vegetation greening, there has been a progressive increase in remotely sensed vegetation browning globally. Given the many societal benefits that forests provide, it is critical that we understand vegetation dynamic alterations. Here, we review associative drivers, impacts, and feedbacks, revealing the complexity of browning. Concomitant increases in browning include the weakening of ecosystem services and functions and alterations to vegetation structure and species composition, as well as the development of potential positive climate change feedbacks. Also discussed are the current challenges in browning detection and understanding associated impacts and feedbacks. Finally, we outline recommended strategies.

Despite new knowledge in recent years, our understanding of the phenology of wood formation for various species growing in different environments remains limited. To enhance our knowledge of the tree growth dynamics of boreal tree species, we investigated the average seasonal, monthly, daily, and diel patterns of tree growth and water status from 11 years of observations with the 15 min and 1.5 µm resolved stem radial size variation data of 12 balsam fir (Abies balsamea (L.) Mill.) trees growing in a cold and humid boreal environment. Growth only occurred above an air temperature threshold of 9–10 °C, and the maximal growth rate over the year (23–24 June) was synchronous with the maximal day length (20–21 June) and not with the maximal air temperature, which occurred on average about 2 weeks later (4–5 July). Tree growth was mostly restricted by air temperature and solar radiation under these cold and wet boreal conditions, but our results also highlight a turgor-driven growth mechanism. Diel dynamics reveal that tree growth is minimal during the day when the stem dehydrates, and higher past midnight when the stem is fully rehydrated. This pattern suggests that carbon assimilation through photosynthesis occurs primarily during the day, while energy production and carbon allocation to woody tissues occur primarily at night via cellular respiration. Overall, our results show that the temporal patterns of the growth and water status of balsam fir growing in cold and humid boreal environments are controlled by a set of environmental factors that influence various physiological processes and mechanisms, many of which still need to be documented.

Increased greenhouse gas emissions are causing unprecedented climate change, which is, in turn, altering emissions and removals (referring to the oxidation of atmospheric CH4 by methanotrophs within the soil) of the atmospheric CH4 in terrestrial ecosystems. In the global CH4 budget, wetlands are the dominant natural source and upland soils are the primary biological sink. However, it is unclear whether and how the soil CH4 exchanges across terrestrial ecosystems and the atmosphere will be affected by warming and changes in precipitation patterns. Here, we synthesize 762 observations of in situ soil CH4 flux data based on the chamber method from the past three decades related to temperature and precipitation changes across major terrestrial ecosystems worldwide. Our meta-analysis reveals that warming (average warming of +2°C) promotes upland soil CH4 uptake and wetland soil CH4 emission. Decreased precipitation (ranging from −100% to −7% of local mean annual precipitation) stimulates upland soil CH4 uptake. Increased precipitation (ranging from +4% to +94% of local mean annual precipitation) accelerates the upland soil CH4 emission. By 2100, under the shared socioeconomic pathway with a high radiative forcing level (SSP585), CH4 emissions from global terrestrial ecosystems will increase by 22.8 ± 3.6 Tg CH4 yr−1 as an additional CH4 source, which may be mainly attributed to the increase in precipitation over 30°N latitudes. Our meta-analysis strongly suggests that future climate change would weaken the natural buffering ability of terrestrial ecosystems on CH4 fluxes and thus contributes to a positive feedback spiral.

Sapwood characteristics, such as sapwood area as well as thermal and hydraulic conductivity, are linked to species-specific hydraulic function and resource allocation to water transport tissues (xylem). These characteristics are often unknown and thus a major source of uncertainty in sap flow data processing and transpiration estimates because bulk rather than species-specific values are usually applied. Here, we analyzed the sapwood characteristics of fifteen common tree species in eastern North America from different taxonomic (i.e., angiosperms and gymnosperms) and xylem porosity groups (i.e., tracheid-bearing, diffuse- or ring-porous species) and we assessed how uncertainties in sapwood characteristics involved in sap flow calculations are propagated in tree water use estimates. We quantified their sapwood area changes with stem diameter (allometric scaling) and thermal conductivity. We combined these measurements with species-specific values of wood density and hydraulic conductivity found in the literature and assessed the role of wood anatomy in orchestrating their covariation. Using an example sap flow dataset from tree species with different xylem porosity, we assessed the sensitivity of tree water use estimates to sapwood characteristics and their interactions. Angiosperms (ring- and diffuse-porous species), with specialized vessels for water transport, showed a steeper relationship (scaling) between tree stem diameter and sapwood area in comparison to gymnosperms (tracheid-bearing species). Gymnosperms (angiosperms) were characterized by lower (higher) wood density and higher (lower) sapwood moisture content, resulting in non-significant differences in sapwood thermal conductivity between taxonomic and xylem porosity groups. Clustering of species sapwood characteristics based on taxonomic or xylem porosity groups and constraining these parameters could facilitate more accurate sap flow calculations and tree water use estimates. When combined with an increasing number of sap flow observations, these findings should improve tree- and landscape-level transpiration estimates, leading to more robust partitioning of terrestrial water fluxes.

Soil temperature (Tsoil) and soil water (θsoil) are fundamental variables that have an essential role in many processes in forest ecosystems, as well as influencing the tree species distribution and forest composition over time. We tested the Canadian Land Surface Scheme (CLASS) capacity to simulate Tsoil and θsoil in the boreal forest using a sixteen-year data set of daily measurements. Sensitivity analyses were also carried out to evaluate the impact of the thickness of organic layer (TOL), soil texture (percentage sand and clay: PS and PC), drainage parameter (DP), and water freezing point (FP) on simulated Tsoil and θsoil. Finally, the model was also calibrated with a combination of model parameters. CLASS well simulated Tsoil while its performance for θsoil varied by soil horizon and season. In winter particularly, soil liquid water was greatly underestimated because simulated Tsoil was below 0 °C. Nevertheless, simulated θsoil seasonal variation corresponded well with observations. Based on sensitivity analyses, TOL had an important effect on both Tsoil and θsoil. Although PS, PC and DP had almost no effect on Tsoil, their effects on θsoil were substantial. Tsoil increased throughout the year and θsoil increased during the winter with decreasing FP, yet the match between modelled θsoil and observations was not substantially improved. In general, Tsoil was well simulated by CLASS, except for the freezing during winter. Model calibration improves greatly both simulated Tsoil and θsoil, especially during winter in all soil layers. However, despite the model calibration, CLASS still requires improvement for modelling Tsoil and θsoil, hence emphasizing the need to review the equations governing these variables in CLASS.

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.

 

Balsam fir (Abies balsamea) is the most vulnerable species to the spruce budworm (Choristoneura fumiferana), one of the most devastating defoliators in the world. For decades, pest managers have advocated for reducing its abundance in the landscape to minimize losses to the spruce budworm (SBW). Although reduction of fir occurred during the endemic phase of the SBW cycle, there is little information about the extent to which this general principle (reduction of fir) was applied during an outbreak and whether it occurs at both stand and landscape levels. The objective of this paper is to compare the effect of insect and harvest disturbances on forest structure during the 1970–80s outbreak in Québec. We evaluate whether, (i) forest management activities targeted fir forests and whether patch size of host species influences management or SBW disturbance, (ii) SBW outbreaks and logging have similar or divergent effects on forest composition. Although data are from an earlier outbreak, they are at a scale rarely studied and will be useful in guiding decisions made at larger scales in the current and future outbreaks. Our results show that spruce was targeted preferentially by harvesting (up to 69% of plots) during the outbreak period, while it represented less than one third of plots defoliated by the SBW. On the other hand, fir stands represented up to 75% of plots that were defoliated by the SBW but less than 35% of plots that underwent harvesting. Harvesting targeted large blocks of spruce forest more than large blocks of fir-dominated forest while the opposite was observed for the SBW. In terms of regeneration, SBW tends to reduce fir and favor spruce recruitment, along with non-host species, whereas the opposite tendency was observed following harvesting. In terms of spatial organization of stands, our results support the suggestion that small stands of fir and large stands of spruce undergo the least SBW damage. Thus, in order to attenuate SBW impacts in the future, efforts should be made to ensure that spruce recruitment is favored and that its abundance increases at both the stand and landscape scale.

Abstract Aim Microbial communities often vary spatially in how they assemble and knowledge is lacking about which factors determine the biogeography of host-associated microbiomes. Our aim is to assess the relative importance of spatial, environmental and host-associated factors on microbial community composition of an important defoliating insect. Location Boreal forests in eastern Canada (Quebec, Newfoundland and Labrador). Taxon The eastern spruce budworm (Choristoneura fumiferana) and its associated bacterial communities. Methods We characterized bacterial communities associated with spruce budworm larvae using bacterial 16S rRNA gene amplicon sequencing. We quantified how much of the variation in these bacterial communities could be explained by (1) environmental conditions, (2) the microbiome of foliage the larvae were eating, (3) host tree species and (4) spatial structure as quantified using Moran's Eigenvector Maps (MEMs). Results Budworm larval microbiomes varied significantly among sites and between host tree species. Larvae bacterial community structure was strongly correlated with the structure of bacterial communities taken from paired foliage samples. Spatial structure, foliage bacterial communities and host tree species collectively explained almost one-sixth of the variation in budworm bacterial communities while environmental conditions did not explain variation on their own. Main conclusions Lepidopteran microbiomes primarily originate from the foliage diet. However, subtle differences in microbial communities between larvae and foliage suggest that some bacteria establish and grow in the budworm microbiome, and that dispersal of bacteria from sources other than foliage as well as differences in environmental filtering between larval bodies and foliage play a role in the assembly of the budworm microbiome. While spatial location and spatial structure were also important drivers of spruce budworm bacterial community composition, none of the environmental variables we measured could explain the variation among sites, and identifying the drivers of this spatial variation remains an open question that will need to be addressed by future studies.

Species identification is a critical factor for obtaining accurate forest inventories. This paper compares the same method of tree species identification (at the individual crown level) across three different types of airborne laser scanning systems (ALS): two linear lidar systems (monospectral and multispectral) and one single-photon lidar (SPL) system to ascertain whether current individual tree crown (ITC) species classification methods are applicable across all sensors. SPL is a new type of sensor that promises comparable point densities from higher flight altitudes, thereby increasing lidar coverage. Initial results indicate that the methods are indeed applicable across all of the three sensor types with broadly similar overall accuracies (Hardwood/Softwood, 83–90%; 12 species, 46–54%; 4 species, 68–79%), with SPL being slightly lower in all cases. The additional intensity features that are provided by multispectral ALS appear to be more beneficial to overall accuracy than the higher point density of SPL. We also demonstrate the potential contribution of lidar time-series data in improving classification accuracy (Hardwood/Softwood, 91%; 12 species, 58%; 4 species, 84%). Possible causes for lower SPL accuracy are (a) differences in the nature of the intensity features and (b) differences in first and second return distributions between the two linear systems and SPL. We also show that segmentation (and field-identified training crowns deriving from segmentation) that is performed on an initial dataset can be used on subsequent datasets with similar overall accuracy. To our knowledge, this is the first study to compare these three types of ALS systems for species identification at the individual tree level.

Living trees in forests emit methane (CH4) from their stems. However, the magnitudes, patterns, drivers, origins, and biogeochemical pathways of these emissions remain poorly understood. We measured in situ CH4 fluxes in poplar stems and soils using static chambers and investigated the microbial communities of heartwood and sapwood by sequencing bacterial 16S, archaeal 16S, and fungal ITS rRNA genes. Methane emissions from poplar stems occurred throughout the sampling period. The mean CH4 emission rate was 2.7 mg m−2 stem d−1. Stem CH4 emission rate increased significantly with air temperature, humidity, soil water content, and soil CH4 fluxes, but decreased with increasing sampling height. The CO2 reduction and methylotrophic methanogenesis were the major methanogenic pathways in wood tissues. The dominant methanogen groups detected in stem tissues were Methanobacterium, Methanobrevibacter, Rice Cluster I, Methanosarcina, Methanomassiliicoccus, Methanoculleus, and Methanomethylophilaceae. In addition, three methanotrophic genera were identified in the heartwood and sapwood – Methylocystis, Methylobacterium, and Paracoccus. Overall, stem CH4 emissions can originate directly from the internal tissues or co-occur from soils and stems. The co-existence of methanogens and methanotrophs within heartwood and sapwood highlights a need for future research in the microbial mechanisms underlying stem CH4 exchange with the atmosphere.

Despite great concern for drought-driven forest mortality, the effects of frequent low-intensity droughts have been largely overlooked in the boreal forest because of their negligible impacts over the short term. In this study, we used data from 6,876 permanent plots distributed across most of the Canadian boreal zone to assess the effects of repeated low-intensity droughts on forest mortality. Specifically, we compared the relative impact of sequential years under low-intensity dry conditions with the effects of variables related to the intensity of dry conditions, stand characteristics, and local climate. Then, we searched for thresholds in forest mortality as a function of the number of years between two forest surveys affected by dry conditions of any intensity. Our results showed that, in general, frequent low-intensity dry conditions had stronger effects on forest mortality than the intensity of the driest conditions in the plot. Frequent low-intensity dry conditions acted as an inciting factor of forest mortality exacerbated by stand characteristics and environmental conditions. Overall, the mortality of forests dominated by shade-tolerant conifers was significantly and positively related to frequent low-intensity dry conditions, supporting, in some cases, the existence of thresholds delimiting contrasting responses to drought. In mixtures with broadleaf species, however, sequential dry conditions had a negligible impact. The effects of frequent dry conditions on shade-intolerant forests mainly depended on local climate, inciting or mitigating the mortality of forests located in wet places and dominated by broadleaf species or jack pine, respectively. Our results highlight the importance of assessing not only climate-driven extreme events but also repeated disturbances of low intensity. In the long term, the smooth response of forests to dry conditions might abruptly change leading to disproportional mortality triggered by accumulated stress conditions. Forest and wildlife managers should consider the cumulative effects of climate change on mortality to avoid shortfalls in timber and habitat.

Transpiration, a key component of the hydrological cycle, contributes greatly to the climate system by transferring large amount of water from soils to the atmosphere. Its correct representation within Land Surface Schemes in climate models is crucial to provide accurate and reliable climate projections. In this study, transpiration simulated by the Canadian Land Surface Scheme (CLASS) was compared to long-term observations of sap flow measurements in two boreal forest sites of eastern Canada dominated by balsam fir and black spruce. In general, CLASS adequately models daily transpiration during the growing season for most of the years at both sites. During the tree rehydration period (preceding the growing season), modeled transpiration was greatly underestimated because of overestimating the duration of the snowpack, the latter restricting transpiration. Moreover, CLASS did not capture the impact of extreme events on tree physiology and maintained high transpiration rates during a heat stress and a drought. During both observed and simulated drought events, transpiration modeled using CLASS was overestimated, due to insensitivity to substantial decreases in soil water content; modeled transpiration being strictly controlled by atmospheric variables (vapour pressure deficit and radiations). Thus, we also proposed and implemented a new equation that was able to increase the sensitivity of CLASS to decreasing soil water content. However, this equation needs to be further tested on different sites and tree species.

Transpiration, a key component of the hydrological cycle, contributes greatly to the climate system by transferring large amount of water from soils to the atmosphere. Its correct representation within Land Surface Schemes in climate models is crucial to provide accurate and reliable climate projections. In this study, transpiration simulated by the Canadian Land Surface Scheme (CLASS) was compared to long-term observations of sap flow measurements in two boreal forest sites of eastern Canada dominated by balsam fir and black spruce. In general, CLASS adequately models daily transpiration during the growing season for most of the years at both sites. During the tree rehydration period (preceding the growing season), modeled transpiration was greatly underestimated because of overestimating the duration of the snowpack, the latter restricting transpiration. Moreover, CLASS did not capture the impact of extreme events on tree physiology and maintained high transpiration rates during a heat stress and a drought. During both observed and simulated drought events, transpiration modeled using CLASS was overestimated, due to insensitivity to substantial decreases in soil water content; modeled transpiration being strictly controlled by atmospheric variables (vapour pressure deficit and radiations). Thus, we also proposed and implemented a new equation that was able to increase the sensitivity of CLASS to decreasing soil water content. However, this equation needs to be further tested on different sites and tree species.

Monitoring of forest response to gradual environmental changes or abrupt disturbances provides insights into how forested ecosystems operate and allows for quantification of forest health. In this chapter, we provide an overview of Smartforests Canada, a national-scale research network consisting of regional investigators who support a wealth of existing and new monitoring sites. The objectives of Smartforests are threefold: (1) establish and coordinate a network of high-precision monitoring plots across a 4400 km gradient of environmental and forest conditions, (2) synthesize the collected multivariate observations to examine the effects of global changes on complex above- and belowground forest dynamics and resilience, and (3) analyze the collected data to guide the development of the next-generation forest growth models and inform policy-makers on best forest management and adaptation strategies. We present the methodological framework implemented in Smartforests to fulfill the aforementioned objectives. We then use an example from a temperate hardwood Smartforests site in Quebec to illustrate our approach for climate-smart forestry. We conclude by discussing how information from the Smartforests network can be integrated with existing data streams, from within Canada and abroad, guiding forest management and the development of climate change adaptation strategies.

Field studies have shown that dense tree canopies and regular tree arrangements reduce noise from a point source. In urban areas, noise sources are multiple and tree arrangements are rarely dense. There is a lack of data on the association between the urban tree canopy characteristics and noise in complex urban settings. Our aim was to investigate the spatial variation of urban tree canopy characteristics, indices of vegetation abundance, and environmental noise levels. Using Light Detection and Ranging point cloud data for 2015, we extracted the characteristics of 1,272,069 public and private trees across the island of Montreal, Canada. We distinguished needle-leaf from broadleaf trees, and calculated the percentage of broadleaf trees, the total area of the crown footprint, the mean crown centroid height, and the mean volume of crowns of trees that were located within 100m, 250m, 500m, and 1000m buffers around 87 in situ noise measurement sites. A random forest model incorporating tree characteristics, the normalized difference vegetation index (NDVI) values, and the distances to major urban noise sources (highways, railways and roads) was employed to estimate variation in noise among measurement locations. We found decreasing trends in noise levels with increases in total area of the crown footprint and mean crown centroid height. The percentages of increased mean squared error of the regression models indicated that in 500m buffers the total area of the crown footprint (29.2%) and the mean crown centroid height (12.6%) had a stronger influence than NDVI (3.2%) in modeling noise levels; similar patterns of influence were observed using other buffers. Our findings suggest that municipal initiatives designed to reduce urban noise should account for tree features, and not just the number of trees or the overall amount of vegetation.

Climate models project warmer summer temperatures will increase the frequency and heat severity of droughts in temperate forests of Eastern North America. Hotter droughts are increasingly documented to affect tree growth and forest dynamics, with critical impacts on tree mortality, carbon sequestration, and timber provision. The growing acknowledgement of the dominant role of drought timing on tree vulnerability to water deficit raises the issue of our limited understanding of radial growth phenology for most temperate tree species. Here, we use well-replicated dendrometer band data sampled frequently during the growing season to assess the growth phenology of 610 trees from 15 temperate species over six years. Patterns of diameter growth follow a typical logistic shape, with growth rates reaching a maximum in June, and then decreasing until process termination. On average, we find that diffuse-porous species take 16–18 days less than other wood-structure types to put on 50% of their annual diameter growth. However, their peak growth rate occurs almost a full month later than ring-porous and conifer species (ca. 24 ± 4 days; mean ± 95% credible interval). Unlike other species, the growth phenology of diffuse-porous species in our dataset is highly correlated with their spring foliar phenology. We also find that the later window of growth in diffuse-porous species, coinciding with peak evapotranspiration and lower water availability, exposes them to a higher water deficit of 88 ± 19 mm (mean ± SE) during their peak growth than ring-porous and coniferous species (15 ± 35 mm and 30 ± 30 mm, respectively). Given the high climatic sensitivity of wood formation, our findings highlight the importance of wood porosity as one predictor of species climatic sensitivity to the projected intensification of the drought regime in the coming decades.

Globally, increasing drought-induced tree mortality rates under climate change are projected to have far-reaching effects on forest ecosystems. Among these forest systems, the boreal forest is considered a ‘tipping element’ of the Earth's climate system. This forest biome plays a critical role in ecosystem services, structures and functions while being highly sensitive to drought stress. Although process-based models are important tools in ecological research, very few have yet been developed that integrate advanced physiological mechanisms to simulate drought-induced mortality in boreal forests. Accordingly, based on the process-based TRIPLEX model, this study introduces the new TRIPLEX-Mortality submodule for the Canadian boreal forests at the stand level, that for the first time successfully incorporates two advanced drought-induced physiological mortality mechanisms (i.e., hydraulic failure and carbon starvation). To calibrate and validate the model, 73 permanent sample plots (PSPs) were selected across Canada's boreal forests. Results confirm a good agreement between simulated mortality and mortality observations (R2=0.79; P<0.01; IA=0.94), demonstrating good model performance in simulating drought-induced mortality in boreal forests. Sensitivity analysis indicated that parameter sensitivity increased as drought intensified, and the shape parameter (c) for calculating percentage loss of conductivity (PLC) was the most sensitive parameter (average SI = -3.51) to simulate tree mortality. Furthermore, the results of model input sensitivity analysis also showed that the model can capture changes in mortality under different drought scenarios. Consequently, our model is suitable for simulating drought-induced mortality in boreal forests while also providing new insight into improving model simulations for tree mortality and associated carbon dynamics in a progressively warmer and drier world.

Although the spatiotemporal dynamics of spruce budworm outbreaks have been intensively studied, forecasting outbreaks remains challenging. During outbreaks, budworm-linked warblers (Tennessee, Cape May, and bay-breasted warbler) show a strong positive response to increases in spruce budworm, but little is known about the relative timing of these responses.

Most songbird species show some degree of fidelity to their previous breeding location, especially after successful reproduction. However, species associated with highly dynamic food sources (e.g., outbreaking insects) may have to adopt more flexible strategies. Three species (Tennessee Warbler, Leiothlypis peregrina; Cape May Warbler, Setophaga tigrina; and Bay-breasted Warbler, S. castanea ) show strong numerical responses to spruce budworm (Choristoneura fumiferana-SBW) outbreaks. These species, referred to as "budworm-linked warblers", might track SBW larvae through extensive natal dispersal. Then, the superabundance of food during outbreaks would be expected to lead to high breeding productivity which, in turn, should promote breeding site fidelity. Here, we aimed to determine whether budworm-linked warblers were faithful to their previous year's breeding season location and, if so, whether their probability of return was influenced by habitat characteristics such as the density of SBW larvae, stand structure, or landscape structure. We hypothesized that return rate of budworm-linked warblers will be high, as reported in other species of New World warblers, and we predicted that among habitat characteristics, return rate will increase with the density of SBW larvae. We banded 117 budworm-linked warblers (94 % being males) in 75 study plots distributed along a gradient of SBW density and searched for returning individuals within 50 m of their capture sites using song playbacks. Contrary to our hypothesis, resighting rate was very low (0-10.5%). This relative "infidelity" suggests that breeding dispersal of budworm-linked warblers was relatively extensive. Only habitat proportion within an 8-km radius had an important (negative) effect on the probability of resighting Bay-breasted Warbler. Budworm-linked warblers did not exhibit strong site fidelity as adults, but instead performed breeding dispersal movements, presumably to track SBW outbreaks. This strategy may reflect strong spatiotemporal variations in the density SBW larvae. © 2021 by the author(s). Published here under license by the Resilience Alliance.

Exposure to allergenic tree pollen is an increasing environmental health issue in urban areas. However, reliable, well-documented, peer-reviewed data on the allergenicity of pollen from common tree species in urban environments are lacking. Using the concept of ‘riskscape’, we present and discuss evidence on how different tree pollen allergenicity datasets shape the risk for pollen-allergy sufferers in five cities with different urban forests and population densities: Barcelona, Montreal, New York City, Paris, and Vancouver. We also evaluate how tree diversity can modify the allergenic risk of urban forests. We show that estimates of pollen exposure risk range from 1 to 74% for trees considered to be highly allergenic in the same city. This variation results from differences in the pollen allergenicity datasets, which become more pronounced when a city’s canopy is dominated by only a few species and genera. In an increasingly urbanized world, diverse urban forests offer a potentially safer strategy aimed at diluting sources of allergenic pollen until better allergenicity data is developed. Our findings highlight an urgent need for a science-based approach to guide public health and urban forest planning.

Purpose of Review

Forest managers have long suggested that forests can be made more resilient to insect pests by reducing the abundance of hosts, yet this has rarely been done. The goal of our paper is to review whether recent scientific evidence supports forest manipulation to decrease vulnerability. To achieve this goal, we first ask if outbreaks of forest insect pests have been more severe in recent decades. Next, we assess the relative importance of climate change and forest management–induced changes in forest composition/structure in driving these changes in severity.

Recent Findings

Forest structure and composition continue to be implicated in pest outbreak severity. Mechanisms, however, remain elusive. Recent research elucidates how forest compositional and structural diversity at neighbourhood, stand, and landscape scales can increase forest resistance to outbreaks. Many recent outbreaks of herbivorous forest insects have been unprecedented in terms of duration and spatial extent. Climate change may be a contributing factor, but forest structure and composition have been clearly identified as contributing to these unprecedented outbreaks.

Summary

Current research supports using silviculture to create pest-resistant forest landscapes. However, the precise mechanisms by which silviculture can increase resistance remains uncertain. Further, humans tend to more often create pest-prone forests due to political, economic, and human resistance to change and a short-sighted risk management perspective that focuses on reactive rather than proactive responses to insect outbreak threats. Future research efforts need to focus on social, political, cultural, and educational mechanisms to motivate implementation of proven ecological solutions if pest-resistant forests are to be favoured by management.

Outbreaks of eastern spruce budworm (Choristoneura fumiferana; hereafter SBW) are a major natural disturbance in coniferous forests of eastern North America. These outbreaks provide a superabundant source of food for insectivorous birds. Three species, referred to as budworm-linked warblers, exhibit strong positive numerical responses to early increases of SBW density: Tennessee Warbler (Leiothlypis peregrina), Cape May Warbler (Setophaga tigrina), and Bay-breasted Warbler (S. castanea). Their abundance increases even before defoliation is visible from aerial surveys. Budworm-linked warblers may detect new epicenters of SBW outbreaks through natal dispersal, as this movement is typically much more extensive than subsequent (breeding dispersal) movements. Our main objectives were, thus, (1) to determine whether sudden increases in the abundance of budworm-linked warblers could be used to detect early stages of SBW outbreaks, and (2) to examine age-specific responses of budworm-linked warblers to local and landscape-level habitat characteristics, in order to investigate the potential role of natal dispersal in the detection of new epicenters. To do so, we estimated the abundance of each species of budworm-linked warbler in 75 study plots sampling a gradient of SBW density and related them to 7 stand variables and landscape metrics with generalized additive mixed models. We also compared the responses of yearling (second-year; SY) and older (after-second-year; ASY) individuals to the density of SBW larvae and habitat variables at different spatial scales. We captured 31 Tennessee Warblers, 27 Cape May Warblers, and 57 Bay-breasted Warblers. The abundance of all three species of budworm-linked warblers increased with SBW larval density, but the numerical response of Bay-breasted Warbler was initiated earlier and it varied with age. SY individuals tended to be associated with stands supporting lower larval densities than ASY individuals and, as suggested by other authors, Bay-breasted Warbler appeared to be more efficient at exploiting SBW larvae at low density. For that reason, this species represents an early indicator of stands undergoing SBW outbreaks and we propose to use its abundance as an indicator to orient labour-intensive ground surveys of SBW larvae.

Aims

In the boreal mixedwood, fire initiates forest succession; however, over time other disturbances such as insect outbreaks cause pulses of mortality and opportunities for recruitment of shade?intolerant species.

Questions

What are the respective roles of time since fire and insect outbreaks in driving directional, vs cyclical and retrogressive succession? Do assessments from direct measurements and chronosequence approaches converge? We hypothesise that the chronosequence approach will accurately characterise large?scale compositional patterns especially in younger stands and that direct measurements will better describe small?scale, non?directional changes in succession.

Location

Boreal mixed wood of northwestern Quebec (79°1? W, 48°30? N).

Methods

We sampled 469 plots over an 18?year interval (in 1991 and 2009) in mixed hardwood/conifer forests to observe in situ the changes in stands having originated from seven fires covering a 249?year chronosequence.

Results

The combination of the remeasurement and chronosequence analyses validates predictions of forest succession. Our results indicate that time since last fire is the dominant factor explaining forest succession for the first 150 years after fire and globally at the scale of the entire forest, although smaller?scale disturbances such as spruce budworm outbreaks can change the course of forest succession, especially at stand scales.

Conclusion

While time since fire is the dominant driver of forest succession in younger forests, secondary disturbances, such as spruce budworm outbreaks, can advance or reverse forest succession in older forests. This study also serves as a validation of the overall patterns described in spatial chronosequence approaches at the landscape level in fire?dominated systems but highlights that local succession may deviate from the overall pattern especially in older forests or in forests affected by non?stand re?initiating intermediate disturbances.

On vegetation-covered land surfaces, tree transpiration, compared to soil and canopy evaporation, is a major process that sends large amounts of water back to the atmosphere. While the driving forces of tree transpiration have been studied over a range of tree species across an array of ecosystems, no work has been done on balsam fir and black spruce in the humid boreal forest of eastern Canada.We thus studied the relationships between environmental variables and sap flow velocity (as a proxy for transpiration) for these two boreal tree species located at two forest sites in Quebec, Canada over multiple growing seasons (2004 to 2013 for balsam fir and 2006 to 2009 for black spruce). Our results showed that daily sap flow had a strong non-linear relationship with vapour pressure deficit (VPD) for both species. Sap flow was also strongly correlated to solar radiation (Rad) for both species although with slightly weaker relationships than for VPD. Other variables such as daily maximum temperature and precipitation explain a smaller portion of the variance in sap flow while soil water content (SWC) and wind speed had almost no effect. An analysis of the relationships between sap flow and VPD/Rad on an hourly basis over multiple years showed strong diel hysteresis for both species. Contrary to what has been previously proposed, the magnitude of this hysteresis does not seem to relate to the degree of iso/anisohydricity. Finally, our investigation of sap flow relationships to environmental variables during a drought period at the balsam fir site showed that sap flow was only slightly reduced despite a significant decrease in SWC. On the other hand, VPD and Rad remained the main drivers of sap flow. This study emphasizes that VPD and Rad are indeed the major drivers of transpiration during the growing season as well as during drought in humid boreal region.

Defoliation can enhance tree water status by reducing canopy transpiration under drought. During long-lasting insect outbreaks however, this effect can be transient as reduced foliage affects not only transpiration but also the entire soil-plant-atmosphere continuum. In this study, we investigated the effects of defoliation and vapor pressure deficit (VPD) on plant and soil water status in balsam fir and black spruce defoliated by spruce budworm, Choristoneura fumiferana (Clemens). We sampled 48 fir trees and 36 spruce trees subjected to differing severities of defoliation. In May–September 2014 and 2015, we monitored the relative shoot water content (RWC) and soil volumetric water content (VWC), and midday shoot water potential (?md, only in 2015). We applied linear mixed models (LMMs) to assess changes in RWC, ?md, and VWC to defoliation and VPD and we ran structural equation models (SEM) to determine the causal relationships between the measured variables in relation to defoliation and VPD. In LMMs models, defoliation and VPD, as individual factors, reduced ?md in both balsam fir and pooled species models but did not affect RWC. Defoliation alone increased VWC in balsam fir and in pooled models. We observed no interaction between VPD and defoliation on tree water status, but significant effect on VWC (in balsam fir and pooled models), indicating that both factors had independent and additive effects on plants but not on soil. However, in SEM models, RWC was negatively correlated to defoliation, suggesting a hydraulic safety margin. Under conditions of multiple-years of natural defoliation during a spruce budworm outbreak, the decrease in ?md reflects the amount of internal water capacitance that could be caused by both a lower ?md due to larval feeding and a negative feedback between defoliation and xylem vulnerability.

Spruce budworm (SBW) outbreaks are a major natural disturbance in boreal forests of eastern North America. During large?scale infestations, aerial spraying of bacterial insecticides is used to suppress local high?density SBW populations. While the primary goal of spraying is the protection of wood volume for later harvest, it should also maintain carbon stored in trees. This study provides the first quantitative analysis of the efficacy of aerial spraying against SBW on carbon dynamics in balsam fir, spruce, and mixed fir–spruce forests. In this study, we used the TRIPLEX?Insect model to simulate carbon dynamics with and without spray applications in 14 sites of the boreal forest located in various regions of Québec. We found that the efficacy of aerial spraying on reducing annual defoliation was greater in the early stage (<5 yr since the outbreak began) of the outbreak than in later (5–10 yr since the outbreak began) stage. Our results showed that more net ecosystem productivity is maintained in balsam fir (the most vulnerable species) than in either spruce or mixed fir–spruce forests following spraying. Also, average losses in aboveground biomass due to the SBW following spraying occurred more slowly than without spraying in balsam fir forests. Our findings suggest that aerial spraying could be used to maintain carbon in conifer forests during SBW disturbances, but that the efficacy of spray programs is affected by host species and stage of the SBW outbreak.

Phenology has become a field of growing importance due to the increasingly apparent impacts of climate change. However, the time-consuming, subjective and tedious nature of traditional human field observations have hindered the development of large-scale phenology networks. Such networks are rare and rely on time-lapse cameras and simplistic color indexes to monitor phenology. To automatize rapid, detailed and repeatable analyzes, we propose an Artificial Intelligence (AI) framework based on machine learning and computer vision techniques. Our approach extracts multiple ecologically-relevant indicators from time-lapse digital photography datasets. The proposed framework consists of three main components: (i) a random forest model to automatically select relevant images based on color information; (ii) a convolutional neural network (CNN) to identify and localize open tree buds; and (iii) a density-based spatial clustering algorithm to cluster open bud detections across the time-series. We tested this framework on a dataset including thousands of black spruce and balsam fir tree images captured using our phenological camera network. The performed experiments showed the efficiency of the proposed approach under challenging perturbation factors, such as significant image noise. Our framework is exceedingly faster and more accurate than human analysts, reducing the time-series processing time from multiple days to under an hour. The proposed methodology is particularly appropriate for large-scale and long-term analyzes of ecological imagery datasets. Our work demonstrates that the use of computer vision and machine learning methods represents a promising direction for the implementation of national, continental, or even global plant phenology networks.

1. As major alterations are occurring in climate and pest ranges, it is imperative to evaluate their combined contribution to tree mortality in order to propose mitigation measures and limit losses in forest productivity. The objective of this study was to explore the association between declines in tree growth resulting from climatic and biotic (spruce budworm) disturbances, and their interactions on tree mortality of two dominant tree species, Abies balsamea and Picea mariana, of the eastern North?American boreal forest.
2. We disentangle the influences of abiotic and biotic components on growth through a combination of model?data comparison techniques. First, we characterized the variability in tree growth and mortality in the study area using a network of tree?ring width measurements collected from living and dead trees. Subsequently, a bioclimatic simulation model was used to estimate the past annual, nonlinear, responses of stand?level net primary production (NPP) to climate variability (period 1902–2012). From these two data sources, we defined the biotic stress events as the variance in the tree?ring data unexplained by the bioclimatic forest growth simulation.
3. Throughout the 20th century, two periods of adverse climatic conditions preceded spruce budworm outbreaks episodes and induced tree mortality. Climatic stress events were associated with cold springs, warmer than average summers. We found that past stress history in interaction with tree characteristics and species predisposed trees to mortality. In addition, co?occurring events (climatic and biotic) increased the severity of mortality episodes.
4. Synthesis. Our study challenges the belief that spruce budworm outbreak is the primary driver of broad?scale tree mortality in eastern boreal forest. Rather, tree mortality is the result of cumulative events that combine unfavourable conditions for growth, resulting in loss of tree vigour and subsequently, mortality. Co?occurrence of stresses in the future may lead to more severe episodes of mortality, as extreme climatic events become more frequent.

Understanding and measuring forest resistance and resilience have emerged as key priorities in ecology and management, particularly to maintain forest functioning. The analysis of the factors involved in a forest’s ability to cope with disturbances is key in identifying forest vulnerability to environmental change. In this study, we apply a procedure based on combining pathway analyses of forest composition and structure with quantitative indices of resistance and resilience to disturbances. We applied our approach to boreal forests affected by a major spruce budworm outbreak in the province of Quebec (Canada). We aimed to identify the main patterns of forest dynamics and the environmental factors affecting these responses. To achieve this goal, we developed quantitative metrics of resistance and resilience. We then compared forests with different pre-disturbance conditions and explored the factors influencing their recovery following disturbance. We found that post-outbreak forest dynamics are determined by distinct resistance and resilience patterns according to dominant species and stand composition and structure. Black spruce forests are highly resistant to spruce budworm outbreaks, but this resistance is conditioned by the length of the defoliation period, with long outbreaks having the potential to lead the system to collapse. In contrast, balsam fir forests easily change to a different composition after outbreaks but are highly resilient when mixed with hardwood species. Overall, the severity of the disturbance and the tree species affected are the main drivers contributing to boreal forest resistance and resilience. Our procedure is valuable to understand post-disturbance dynamics of a broad range of communities and to guide management strategies focused on enhancing the resistance and resilience of the system.

Climate change is predicted to alter relationships between trophic levels by changing the phenology of interacting species. We tested whether synchrony between two critical phenological events, budburst of host species and larval emergence from diapause of eastern spruce budworm, increased at warmer temperatures in the boreal forest in northeastern Canada. Budburst was up to 4.6 ± 0.7 days earlier in balsam fir and up to 2.8 ± 0.8 days earlier in black spruce per degree increase in temperature, in naturally occurring microclimates. Larval emergence from diapause did not exhibit a similar response. Instead, larvae emerged once average ambient temperatures reached 10°C, regardless of differences in microclimate. Phenological synchrony increased with warmer microclimates, tightening the relationship between spruce budworm and its host species. Synchrony increased by up to 4.5 ± 0.7 days for balsam fir and up to 2.8 ± 0.8 days for black spruce per degree increase in temperature. Under a warmer climate, defoliation could potentially begin earlier in the season, in which case, damage on the primary host, balsam fir may increase. Black spruce, which escapes severe herbivory because of a 2?week delay in budburst, would become more suitable as a resource for the spruce budworm. The northern boreal forest could become more vulnerable to outbreaks in the future.

Tree seedlings planted in abandoned agricultural fields interact with herb communities through competition, tolerance, and facilitation. In addition, they are subject to herbivory by small mammals, deer or invertebrates. To increase the success of forest restoration in abandoned fields and reduce management costs, we should determine which species are tolerant to or facilitated by herbaceous vegetation and those which require protection from competition and predation. Eight native tree species were planted in plots covered by herbaceous vegetation, plots where herbaceous vegetation was removed, and plots where seedlings were surrounded by an organic mulch mat. Half of the seedlings were protected against small mammal damage. Results showed that two non-pioneer and moderately shade-tolerant species (yellow birch and red oak) were inhibited by herbaceous vegetation. Birch species were particularly affected by small mammal predation. No effects of predation or herbaceous competition were observed for conifer species. Rather, herbaceous vegetation had a positive effect on the survival and the height growth of tamarack (Larix laricina). None of the tested herb communities had a stronger competitive effect on tree growth than another. Restoration of abandoned fields using multi-tree species should be designed at the seedling scale rather than at the site scale to account for different tree responses to predation and competition as well as variable site conditions. An approach resembling precision agriculture is proposed to lower costs and any potential negative impact of more intensive vegetation management interventions. © 2018 by the authors.

Natural disturbances are fundamental to forest ecosystem dynamics and have been used for two decades to improve forest management, notably in the boreal forest. Initially based on fire regimes, there is now a need to extend the concept to include other types of disturbances as they can greatly contribute to forest dynamics in some regions of the boreal zone. Here we review the main descriptors—that is, the severity, specificity, spatial and temporal descriptors and legacies, of windthrow and spruce bud worm outbreak disturbance regimes in boreal forests—in order to facilitate incorporating them into a natural disturbance-based forest management framework. We also describe the biological legacies that are generated by these disturbances. Temporal and spatial descriptors characterising both disturbance types are generally variable in time and space. This makes them difficult to reproduce in an ecosystem management framework. However, severity and specificity descriptors may provide a template upon which policies for maintaining post harvesting and salvage logging biological legacies can be based. In a context in which management mainly targets mature and old-growth stages, integrating insect and wind disturbances in a management framework is an important goal, as these disturbances contribute to creating heterogeneity in mature and old-growth forest characteristics.

L’apprentissage automatique, une branche importante de l’intelligence artificielle, est de plus en plus mis en application dans le domaine des sciences comme l’écologie forestière. Ici, nous examinons et faisons le point sur trois méthodes d’apprentissage automatique généralement utilisées incluant l’apprentissage par arbre de décision, le réseau de neurones artificiels et la machine à vecteurs de support, ainsi que leurs applications au niveau de quatre aspects différents de l’écologie forestière au cours de la dernière décennie. Ces applications incluent : (i) les modèles de répartition des espèces, (ii) les cycles de carbone, (iii) l’évaluation et la prédiction des dangers et (iv) d’autres applications en gestion forestière. Alors que les approches d’apprentissage automatique sont utiles au niveau de la classification, de la modélisation et de la prédiction en recherche dans le domaine de l’écologie forestière, le développement accru des technologies d’apprentissage automatique est limité par le manque de données pertinentes et le « seuil relativement plus élevé » des applications. Cependant, l’utilisation combinée d’algorithmes multiples et de communication et coopération améliorées entre les chercheurs en écologie et les concepteurs d’apprentissage automatique présente toujours des défis importants et des tâches en vue de l’amélioration de la recherche écologique à l’avenir. Nous laissons entendre que les applications futures d’apprentissage automatique en écologie deviendront un outil de plus en plus intéressant pour les écologistes face aux « données massives » et que les écologistes auront accès à plus de types de données tel que le son et la vidéo dans un proche avenir ce qui ouvrira probablement de nouvelles avenues en matière de recherche en écologie forestière. [Traduit par la Rédaction]

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.