Spruce Budworm Research Articles

Biennial aerial application of Bacillus thuringiensis Berliner var. kurstaki is the most cost-effective approach of protection against spruce budworm (Choristoneura fumiferana [Clemens])

Key messageAerial application of Bacillus thuringiensis Berliner var. kurstaki (Btk) every second year to stands of white spruce (Picea glauca (Moench)Voss.), black spruce (Picea mariana Mill.) and balsam fir (Abies balsamea (L.) Mill.) is the most cost-effective spraying scenario for reducing the impact of spruce budworm (Choristoneura fumiferana Clemens) on wood production, providing a similar level of forest protection, but at lower cost, to the standard scenario currently used in which 50% of current year’s foliage is protected every year.ContextInsect outbreaks can have significant effects on forest productivity and various formulations of Bacillus thuringiensis Berliner var. kurstaki (Btk) are used to reduce their damage. In the Province of Québec, Canada, control programs aim to protect at least 50% of current-year foliage to limit tree mortality, but little information exists on the long-term cost-effectiveness of such programs.AimsOur goal was to evaluate the benefit/cost ratio and the efficacy of different Btk protection scenarios in reducing coniferous tree mortality and growth losses over a 11-year period. We hypothesized that less-intensive protection approaches (Btk applications every 2 or 3 years) may provide similar levels of protection but with higher cost-effectiveness ratios than the standard program currently used in Quebec.MethodsIn 2007, we established nineteen 100-ha experimental units in Quebec’s Côte-Nord region to determine the efficacy and cost-effectiveness of different Btk spraying scenarios for reducing tree mortality and volume losses in coniferous stands dominated by mature balsam fir trees (Abies balsamea (L.) Mill., with white spruce (Picea glauca (Moench) Voss.) and black spruce (P. mariana Mill.) as companion species. Tree mortality was monitored annually in three circular plots of 400 m2 within each experimental unit. Growth losses were evaluated using stem analyses.ResultsMortality was much higher in balsam fir than in black spruce and white spruce (respectively 74.4%, 13.8% and 5.9% in untreated stands) in all protection scenarios. The application of Btk every 2 years reduced balsam fir mortality to a level similar to the standard scenario (10.3% vs 7.15%, respectively) at a much lower cost. Growth losses have also been reduced but not to the same extent as in the standard scenario.ConclusionSpraying Btk every 2 years provides effective protection to balsam fir and is the most cost-effective scenario. A less intensive use of Btk would mitigate impact on non-target lepidoptera and allow protecting other areas, such as habitat of the woodland caribou, a threatened species which avoids disturbed areas.

On the characterization of patterning in spruce budworm time-series data

I outline the “definitional problem” in forest insect outbreak analytics and show how it is related to the “counting problem” in dendroentomology and the “forecasting problem” in forest insect population dynamics, through the ubiquitous presence of non-stationary complex periodicity. Using real-world examples from the spruce budworm ( Choristoneura fumiferana Clem.) system, I show that regardless how outbreak patterning is characterized—whether by peak impact, cycle frequency, interval duration, or interval severity—the distribution in pattern attributes appears to be extremely variable, regardless how the data are processed through definitional filters. I show that this extreme variability is an unavoidable and key feature of the system’s dynamics and argue that it needs to be viewed as an object of study, instead of a nuisance problem to be swept under the rug. The single biggest opportunity for rapid gains in spruce budworm predictive ecology is determining the environmental and ecological factors that separate high-intensity from low-intensity outbreak cycling.

One century of carbon dynamics in the eastern Canadian boreal forest under various management strategies and climate change projections

Partial cutting has lower canopy removal intensities than clearcutting and has been proposed as an alternative harvesting approach to enhance ecosystem services, including carbon sequestration and storage. However, the ideal partial cutting/clearcutting proportion that should be applied to managed areas of the eastern Canadian boreal forest to enhance long-term carbon sequestration and storage at the landscape scale remains uncertain. Our study projected carbon dynamics over 100 years (2010–2110) under a portfolio of management strategies and future climate scenarios within three boreal forest management units in Quebec, Canada, distributed along an east–west gradient. To model future carbon dynamics, we used LANDIS-II, its Forest Carbon Succession extension, and several extensions that account for natural disturbances in the boreal forest (wind, fire, spruce budworm). We simulated the effects of several management strategies on carbon dynamics, including a business-as-usual strategy (clearcutting applied to more than 95 % of the annually managed area), and compared these projections against a no-harvest natural dynamics scenario. We projected an overall increase in total ecosystem carbon storage, mostly because of increased productivity and broadleaf presence under limited climate change. The drier Western region under climate scenario RCP8.5 was an exception, as stocks decreased after 2090 because of the direct negative effects of extreme climate change on coniferous species’ productivity. Under the natural dynamic scenario, our simulations suggest that the Quebec Forest in the Central and Western regions may act as a carbon sink, despite high fire-related carbon emissions, particularly under RCP4.5 and RCP8.5 Conversely, the eastern region periodically switched from carbon sink to source following SBW outbreaks, thus being a weak sink over the simulation period. Applying partial cutting to over 50 % of the managed forest area effectively mitigated the negative impacts of climate change on carbon balance, reducing differences in stand composition and carbon storage between naturally dynamic forests and those managed for timber. In contrast, clearcutting-based scenarios, including the business-as-usual approach, substantially reduced total ecosystem carbon storage— by approximately double (10 tC ha−1 yr−1) compared to partial cutting scenarios (<5 tC ha−1 yr−1). Clearcutting led to higher heterotrophic respiration due to the proliferation of fast-decomposing broadleaves, resulting in lower carbon accumulation compared to partial cuts. Our findings underscore the importance of balancing canopy removal intensities to increase carbon sequestration and storage while preserving other ecosystem qualities under climate change.

Budworms, beetles and wildfire: Disturbance interactions influence the likelihood of insect‐caused disturbances at a subcontinental scale

Abstract Irruptive forest insects are a leading biotic disturbance across temperate and boreal forests. Outbreaks of forest insects are becoming more frequent and extensive due to anthropogenic drivers (e.g. climate and land‐use), perhaps increasing the likelihood that forests will experience multiple insect‐caused disturbances. Across the fire‐prone Douglas‐fir forests of western North America, recent outbreaks of the western spruce budworm and Douglas‐fir beetle have impacted large expanses of forests, with a higher degree of overlap than expected in some ecoregions. Outbreaks of both insects are positively related to host availability and exhibit density‐dependent population dynamics that are affected by climate and weather. Here, we leverage data from aerial detection surveys, estimates of host availability, climate and weather, and categorized fire severity to describe the spatial overlap between western spruce budworm and Douglas‐fir beetle and assess: (1) how climate and host availability influence the biogeography of outbreaks; (2) how weather incites outbreaks; and finally, (3) how prior disturbances (fire and biotic) affect the subsequent outbreak likelihood of western spruce budworm and Douglas‐fir beetle. Models demonstrate that western spruce budworm and Douglas‐fir beetle share similar predisposing drivers of outbreaks. Outbreaks of both insects were more likely to occur following warm weather, but only beetle outbreaks were more likely following drought. When controlling for differences in outbreak distribution and inciting factors, results indicate that both prior fire and interspecific disturbance altered the likelihood of subsequent insect‐caused disturbance. Specifically, Douglas‐fir beetle outbreaks were more likely to occur for several years following low severity fire, but less likely otherwise. Prior defoliation, especially longer duration defoliation, increased the likelihood of beetle outbreak within stands and across the landscape. On the contrary, western spruce budworm outbreaks were less likely to occur following fire, while prior beetle activity dampened budworm outbreak likelihood for several years, and then eventually increased outbreak likelihood. Synthesis: Biotic–biotic disturbance interactions have the potential to amplify the incidence of insect‐caused disturbance across subcontinental scales. Our findings highlight the need for future work on mechanistic linkages between biotic disturbance agents as well as the ramifications for forest trajectories and function.

Spatial propagation in a delayed spruce budworm diffusive model

We investigate the spatial propagation in a delayed spruce budworm diffusive model where and represent, respectively, the incubation and the maturation delays for the spruce budworm. We find the minimal wave speed to determine the existence of traveling wave fronts of the model. More specifically, the model admits traveling wave fronts when ; the model has no traveling wave solutions when . The proofs are based on combining the upper and lower solutions with the approach of Wu and Zou's theorems, the limit arguments, and Laplace transform. The obtained results help us to understand the spreading patterns and the spreading speed of spruce budworm population.

Interfacial Ice Density Fluctuations Inform Surface Ice-Philicity.

The propensity of a surface to nucleate ice or bind to ice is governed by its ice-philicity─its relative preference for ice over liquid water. However, the relationship between the features of a surface and its ice-philicity is not well understood, and for surfaces with chemical or topographical heterogeneity, such as proteins, their ice-philicity is not even well-defined. In the analogous problem of surface hydrophobicity, it has been shown that hydrophobic surfaces display enhanced low water-density (vapor-like) fluctuations in their vicinity. To interrogate whether enhanced ice-like fluctuations are similarly observed near ice-philic surfaces, here we use molecular simulations and enhanced sampling techniques. Using a family of model surfaces for which the wetting coefficient, k, has previously been characterized, we show that the free energy of observing rare interfacial ice-density fluctuations decreases monotonically with increasing k. By utilizing this connection, we investigate a set of fcc systems and find that the (110) surface is more ice-philic than the (111) or (100) surfaces. By additionally analyzing the structure of interfacial ice, we find that all surfaces prefer to bind to the basal plane of ice, and the topographical complementarity of the (110) surface to the basal plane explains its higher ice-philicity. Using enhanced interfacial ice-like fluctuations as a measure of surface ice-philicity, we then characterize the ice-philicity of chemically heterogeneous and topologically complex systems. In particular, we study the spruce budworm antifreeze protein (sbwAFP), which binds to ice using a known ice-binding site (IBS) and resists engulfment using nonbinding sites of the protein (NBSs). We find that the IBS displays enhanced interfacial ice-density fluctuations and is therefore more ice-philic than the two NBSs studied. We also find the two NBSs are similarly ice-phobic. By establishing a connection between interfacial ice-like fluctuations and surface ice-philicity, our findings thus provide a way to characterize the ice-philicity of heterogeneous surfaces.

When do multiple pulses of environmental variation trigger tipping in an ecological system?

Climate change and anthropogenic impacts have a significant effect on natural ecosystems. As a response, tipping phenomena, i.e., abrupt qualitative changes in the dynamics of ecosystems, like transitions between alternative stable states, can be observed. We study such critical transitions, caused by an interplay between B-tipping, the rate of change of environmental forcing, and a rate-dependent basin boundary crossing. Instead of a slow trend of environmental change, we focus on pulses of variation in the carrying capacity in a simple ecological model, the spruce budworm model, and show how one pulse of environmental change can lead to tracking the current stable state or to tipping to an alternative state depending on the strength and the duration of the pulse. Moreover, we demonstrate that applying a second pulse after the first one, which can track the desired state, can lead to tipping, although its rate is slow and does not even cross the critical threshold. We explain this unexpected behavior in terms of the interacting timescales, the intrinsic ecological timescale, the rate of environmental change, and the movement of the basin boundaries separating the basins of attraction of the two alternative states.

Characterizing forest structural changes in response to non-stand replacing disturbances using bitemporal airborne laser scanning data

Characterizing the extent, severity, and persistence of natural disturbances in forests is crucial in areas as large and heterogeneous as the Canadian boreal forest. Non-stand replacing (NSR) disturbances, in particular, can produce subtle and lagged impacts to forest canopy and structure with mechanisms that remain elusive, and they are challenging to discern using typical remote sensing approaches including aerial photointerpretation and spectral analysis of satellite imagery. Consequently, there is a need for timely and accurate information on the structural modifications due to NSR disturbances to inform proactive forest management practices. To address these needs, we leveraged a unique bitemporal airborne laser scanning (ALS) dataset to characterize changes in the forest structure caused by eastern spruce budworm (ESB, Choristoneura fumiferana (Clem.)), responsible for one of the greatest tree mortality in Canada. A range of infestation severity with varying impacts to forest structure are examined in a mixedwood boreal forest in Lac-Saint Jean, Quebec, Canada. We derived 14 ALS structural change metrics at 10 m spatial resolution, including height, cover, and gappiness 7 years apart (2014–2020). Six distinct structural responses to cumulative ESB infestations severity were identified using cluster analysis from the combination of the 14 change metrics, with canopy cover, the 75th and 25th height percentiles (p75-25) driving cluster separability. Canopy cover and p25 consistently decreased as cumulative infestation severity increased, whereas p75 showed greater variability across the landscape. Photointerpretation of aerial imagery over the same period confirmed the validity of the structural characterization. Further, we studied the role of initial forest structures in modulating the severity of the infestation and found that sparser canopies with cover <65% and shorter trees (p75 < 7.5 m, p25 < 2.5 m) were associated with less severe ESB infestations after 7 years, and controlling for underlying environmental factors. These findings showed the potential of bitemporal ALS data in characterizing structural changes due to ESB infestations at fine scale based on canopy cover and height, relevant for forest management strategies to better target current and future infestations.

Diversity, Stability, and the Forecast Challenge in Forest Lepidopteran Predictive Ecology: Are Multi-Scale Plant–Insect Interactions the Key to Increased Forecast Precision?

I report on long-term patterns of outbreak cycling in four study systems across Canada and illustrate how forecasting in these systems is highly imprecise because of complexity in the cycling and a lack of spatial synchrony amongst sample locations. I describe how a range of bottom-up effects could be generating complexity in these otherwise periodic systems. (1) The spruce budworm in Québec exhibits aperiodic and asynchronous behavior at fast time-scales, and a slow modulation of cycle peak intensity that varies regionally. (2) The forest tent caterpillar across Canada exhibits eruptive spiking behavior that is aperiodic locally, and asynchronous amongst regions, yet aggregates to produce a pattern of periodic outbreaks. In Québec, forest tent caterpillar cycles differ in the aspen-dominated northwest versus the maple-dominated southeast, with opposing patterns of cycle intensity between the two regions. (3) In Alberta, forest tent caterpillar outbreak cycles resist synchronization across a forest landscape gradient, even at very fine spatial scales, resulting in a complex pattern of cycling that defies simple forecasting techniques. (4) In the Border Lakes region of Ontario and Minnesota, where the two insect species coexist in a mixedwood landscape of hardwood and conifers, outbreak cycle intensity in each species varies spatially and temporally in response to host forest landscape structure. Much attention has been given to the effect of top-down agents in driving synchronizable population cycles. However, foliage loss, tree death, and forest succession at stem, stand, and landscape scales affect larval and adult dispersal success, and may serve to override regulatory processes that cause otherwise top-down-driven periodic, synchronized, and predictable population oscillations to become aperiodic, asynchronous, and unpredictable. Incorporating bottom-up effects at multiple spatial and temporal scales may be the key to making significant improvements in forest insect outbreak forecasting.

Response of stream habitat and microbiomes to spruce budworm defoliation: New considerations for outbreak management.

Defoliation by eastern spruce budworm is one of the most important natural disturbances in Canadian boreal and hemi-boreal forests with annual area affected surpassing that of fire and harvest combined, and its impacts are projected to increase in frequency, severity, and range under future climate scenarios. Deciding on an active management strategy to control outbreaks and minimize broader economic, ecological, and social impacts is becoming increasingly important. These strategies differ in the degree to which defoliation is suppressed, but little is known about the downstream consequences of defoliation and, thus, the implications of management. Given the disproportionate role of headwater streams and their microbiomes on net riverine productivity across forested landscapes, we investigated the effects of defoliation by spruce budworm on headwater stream habitat and microbiome structure and function to inform management decisions. We experimentally manipulated a gradient of defoliation among 12 watersheds during a spruce budworm outbreak in the Gaspésie Peninsula, Québec, Canada. From May through October of 2019-2021, stream habitat (flow rates, dissolved organic matter [DOM], water chemistry, and nutrients), algal biomass, and water temperatures were assessed. Bacterial and fungal biofilm communities were examined by incubating six leaf packs for five weeks (mid-August to late September) in one stream reach per watershed. Microbiome community structure was determined using metabarcoding of 16S and ITS rRNA genes, and community functions were examined using extracellular enzyme assays, leaf litter decomposition rates, and taxonomic functional assignments. We found that cumulative defoliation was correlated with increased streamflow rates and temperatures, and more aromatic DOM (measured as specific ultraviolet absorbance at 254 nm), but was not correlated to nutrient concentrations. Cumulative defoliation was also associated with altered microbial community composition, an increase in carbohydrate biosynthesis, and a reduction in aromatic compound degradation, suggesting that microbes are shifting to the preferential use of simple carbohydrates rather than more complex aromatic compounds. These results demonstrate that high levels of defoliation can affect headwater stream microbiomes to the point of altering stream ecosystem productivity and carbon cycling potential, highlighting the importance of incorporating broader ecological processes into spruce budworm management decisions.

Defoliation level interacts with tree species and soil richness to determine volume increment reduction and recovery from simulated spruce budworm attack

Defoliation and tree species determine growth during spruce budworm outbreaks, with site showing inconsistent effects. We determined effects of artificial defoliation treatments (0%, 50%, 100%, 100%+bud) for 3 years on volume increment of 240 balsam fir, black spruce, and white spruce trees, initially 7–10 years old and 2.3–4.1 m tall, on four soil drainage/richness classes. Current annual increment was significantly affected by species × site and site × defoliation interactions, with DBH as covariate. Specific volume increment (SVI) was significantly affected by a species × site × defoliation treatment interaction. Combining species and sites, volume reductions after 3 years were 21%–28%, 43%–51%, and 55%–66% for 50% defoliation, 100% defoliation, and 100%+bud. SVI was negatively, linearly related to cumulative defoliation. Marginal r2 and conditional r2 showed that 64% of variance of SVI was explained by site, species, cumulative defoliation, and DBH, but 90% was explained by fixed and random variables. After 3 years of defoliation, SVI of white spruce was 21%–23% greater than balsam fir and black spruce, and SVI on rich was 55%–79% greater than on poor sites. Cumulative defoliation was a good predictor of growth, and soil richness had a stronger effect on growth after defoliation than in previous studies.

Effects of spruce budworm defoliation on in-stream algal production and carbon use by food webs

Insect defoliation is a widespread cause of forest disturbance in Canada, and it has the potential to alter terrestrial carbon contributions to stream consumers and thus affect stream ecosystems. Although defoliation by spruce budworms ( Choristoneura fumiferana) is considered a natural disturbance with importance to forest succession, the magnitude and duration of outbreaks are increasing, and there is a growing need to understand their downstream consequences. To examine this, algal productivity and autochthonous (in-stream) energy supporting macroinvertebrate and fish consumers (using δ13C, δ15N) were examined in 12 streams in the Gaspé Peninsula, Québec, Canada, that ranged in watershed defoliation from spruce budworm. Hierarchical partitioning models were used to assess the contributions of watershed and stream variables to these endpoints. Total seasonal algal production ranged fourfold among sites, and cumulative defoliation was the strongest predictor. Consumers from streams with greater defoliation (cumulative or annual) had lower reliance on autochthonous carbon, despite increased algal production, and this effect was greater for brook trout than macroinvertebrates. These findings demonstrate for the first time that insect outbreaks—which alter forest structure and function—also similarly affect downstream aquatic ecosystems by causing changes in the production and contributions of carbon sources. This suggests that insect control strategies and forest management decisions should also consider implications for downstream aquatic ecosystems.

A dynamic outbreak distribution model (DODM) for an irruptive folivore: The western spruce budworm

Forest disturbances caused by irruptive insect outbreaks have become more frequent due to the non-stationary and warming climate that favours insect development and survival, as well as forest management practices that increase host tree susceptibility. However, predicting and managing the impacts of these highly mobile pests is a challenge due to their uncertain distributions. Dynamic species distribution models (DSDM) acknowledge the non-stationary environment and species’ ecological niches and facilitate niche modeling by matching species observational data relative to environmental data, leading to better outcomes and reducing over-prediction in conservation efforts. The western spruce budworm (WSB; Choristoneura freemani Freeman) is a native forest pest with irruptive dynamics characterized by periodic landscape-scale outbreaks. WSB feeds primarily on Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) foliage and can cause extensive defoliation throughout western North American forests. Using decades-long presence and absence records of WSB outbreaks detected via aerial surveys and environmental variables, we developed dynamic outbreak distribution models (DODMs) to predict the distribution of outbreaks over time. Our results showed that DODMs provided robust temporal trends of WSB distributions while generating predictions that support documented northward shifts in epidemic populations. We found that in addition to the environmental variables known to affect WSB survival, phenological synchrony between bud burst and the emergence of larvae in spring was the most important predictor of outbreak distributions. We also identified high-risk areas for future WSB outbreaks and predicted range expansions and contractions under different climate change scenarios. Our study highlights the importance of using presence-absence data and incorporating temporal trends in environmental variables for better assessment of forest pest potential outbreak distributions. These findings have implications for proactive management strategies to prevent outbreaks and mitigate the impacts of WSB on forest ecosystems.

Mass Trapping Lepidopteran Pests with Light Traps, with Focus on Tortricid Forest Pests: What If?

The management of Lepidopteran pests with light traps (LTs) is often achieved by luring adults to death at light sources (light trap-based mass trapping, or LTmt). Large-scale LTmt programs against agricultural pests initiated in the late 1920s in the United States were phased out in the 1970s, coinciding with the rise of pheromone-based management research. The interest in LTmt has surged in recent years with the advent of light emitting diodes, solar power sources, and intelligent design. The first step in implementing LTmt is to identify a trapping design that maximizes the capture of target pests and minimizes the capture of non-target beneficial insects-with a cautionary note that high captures in LTs are not equivalent to the feasibility of mass trapping: the ultimate objective of LTmt is to protect crop plants from pest damage, not to trap adults. The captures of egg-carrying females in light traps have a greater impact on the efficiency of LTmt than the captures of males. When LTmt is defined as a harvesting procedure, the biomass of females in LTs may be viewed as the best estimator of the mass trapping yield; biomass proxy has universal application in LTmt as every living organism can be defined on a per weight basis. While research has largely focused on agricultural pests, an attempt is made here to conceptualize LTmt as a pest management strategy in forest ecosystems, using spruce budworm as a case study. The mass trapping of female budworms is impossible to achieve in endemic populations due to the large spatial scale of forest landscapes (implying the deployment of a prohibitively large number of LTs); in addition, ovipositing female budworms do not respond to light sources at a low density of conspecifics. The light-based mass trapping of female budworms may provide a realistic management option for geographically isolated forest stands heavily infested with budworms, as a tool to prevent tree mortality. Somehow unexpectedly, however, one factor obscuring the feasibility of LTmt is as follows: the complex ('unknowable') economic valuation of forest stands as opposed to agricultural landscapes.

Limited Differences in Insect Herbivory on Young White Spruce Growing in Small Open Plantations and under Natural Canopies in Boreal Mixed Forests.

In managed boreal forests, both plantations and natural regeneration are used to re-establish a cohort of conifer trees following harvest or disturbance. Young trees in open plantations generally grow more rapidly than under forest canopies, but more rapid growth could be compromised by greater insect damage. We compared insect damage on white spruce (Picea glauca (Moench) Voss, Pinaceae) growing in plantations with naturally regenerated trees under mature forest canopies in boreal forests (Québec, Canada). We selected ten sites in the naturally regenerated forest and in small, multispecies plantations and sampled ten young trees of 2.5-3 m (per site) in late summer 2020 and again in early and late summer 2021. We compared overall rates of herbivory, galls (adelgids), damage by the spruce budworm (Choristoneura fumiferana, Clemens), and defoliation from sawflies. Overall, insect herbivory damage remained at similarly low levels in both habitats; an average of 9.3% of expanding shoots were damaged on forest trees and 7.7% in plantation trees. Spruce budworm damage increased from 2020 to 2021 and remained higher in under-canopy trees, but damage rates were negligible at this early stage of the outbreak (1.5% in forest vs. 0.78% of buds damaged on plantation trees). While damage due to galls was higher in plantations, the overall low level of damage likely does not pose a significant impact on the growth or mortality of young trees.

Lepidopteran scales in lake sediments as a reliable proxy for spruce budworm outbreak events in the boreal forest of Eastern Canada

Characterizing disturbance regimes over long time scales is paramount for describing and identifying their variability. The most important biotic disturbance in the eastern Canadian boreal forest is the defoliation caused by the eastern spruce budworm, a moth of the insect order Lepidoptera. Lepidopteran scales have recently been used to reconstruct spruce budworm population fluctuations throughout the Holocene. However, this novel proxy has yet to be compared to an independent proxy. This study aimed to determine whether lepidopteran scales found in the surface sediments of boreal lakes tracked large spruce budworm populations, that is, outbreaks, using yearly aerial surveys (1967–present) of spruce budworm defoliation as an independent proxy. Scales were extracted (1 cm resolution) from the top 20 cm of 210Pb-dated sediment cores recovered from nine lakes. To identify significant abundance peaks of scales in the time series, we removed background noise using a modified version of CharAnalysis. A 100-year smoothing window width combined with a 60th percentile threshold yielded the highest true positive and true negative occurrences, and the lowest false positive and false negative occurrences, with values of 0.69 and 0.70 for Cohen’s Kappa and Matthews correlation coefficient, respectively. Our findings demonstrate that lepidopteran scales are a suitable proxy for identifying spruce budworm outbreaks in the sediment record enabling the reconstruction of budworm and other lepidopteran species outbreak dynamics at millennial timescales.

Density‐dependence and competition between Tranosema rostrale and Elachertus cacoeciae, two parasitoids of the spruce budworm

AbstractCompetition between parasitoids for available hosts is common in insects. These interactions are often negative and are believed to reduce the overall impact of top‐down population regulation. In the spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae), two parasitoid species are very common and often interact during the period of low‐density populations between outbreaks: the koinobiont endoparasitoid Tranosema rostrale (Brishke) (Hymenoptera: Ichneumonidae) and the idiobiont ectoparasitoid Elachertus cacoeciae (Howard) (Hymenoptera: Eulophidae). Data collected from field‐implanted larvae of spruce budworm exposed in summer during a 35‐year period between 1987 and 2022 were used to estimate the response of these two parasitoids to the density of their spruce budworm host, and the nature and importance of competitive interactions between them. It was found that both parasitoids are most effective at very low host population density, and that a strong, density‐dependent, and negative correlation of frequencies exists between them. This negative correlation is evident both among and within years and indicates that the parasitoids have developed seasonal history and host choice behaviors that tend to minimize the negative impact of direct competition on their respective performance. It appears that the koinobiont T. rostrale manages, through those adaptive strategies, to avoid direct competition inside a larval host with the competitively superior idiobiont E. cacoeciae.

Increased unpredictability in spruce budworm outbreaks following habitat loss and landscape fragmentation

The spruce budworm (SBW) is the insect defoliator responsible for the greatest tree mortality in North America. Fragmentation and loss of habitat are expected to affect the movement of SBW through forested landscapes during periods of dispersal thereby modifying outbreak severity and frequency.We developed a spatially explicit tri-trophic model by combining a forest landscape model and a system of difference equations to evaluate how the interactions between the SBW, its hosts and its parasitoids are disrupted by habitat loss and fragmentation. We designed landscapes with varying quantities of habitat and fragmentation levels to measure their influence on population dynamics and outbreak severity and frequency.Our model simulations show that landscapes with higher proportion of host tree habitat were also characterised by greater SBW average densities, average cumulative defoliation and outbreak severity, which agrees with earlier studies. However, the bottom-up control by tree hosts breaks down as their proportion in the landscape decreased or when they formed isolated patches. Indeed, we found that landscapes with a low proportion of habitat fragmented into relatively large distant patches produced greatest variability in average SBW abundance and cumulative defoliation between simulation runs. In landscapes with higher habitat proportion, increased isolation between patches produced asynchronous outbreaks. Moreover, in these landscapes, parasitoids occupy fewer habitat cells than the SBW, suggesting that they were not able to efficiently track their prey. Together, these results indicate that increased distance between habitat patches can disrupt trophic control of SBW leading to increased unpredictability and asynchrony in outbreak dynamics.For continued effective management of SBW using current deterministic decision-making tools, we recommend prioritising the maintenance of habitat connectivity in forested landscapes.

Evaluation of branch sampling, ocular assessments, and aerial surveys for estimating spruce budworm defoliation

We compared three methods for estimating current-year spruce budworm ( Choristoneura fumiferana (Clem.)) defoliation from 2014 to 2021 using a network of 99 permanent sample plots in central Gaspé Peninsula, Québec. Percent current-year defoliation was measured by assessing shoots from mid-crown branches, ocular ratings of all individual trees using binoculars, and provincial government aerial surveys. Ocular survey defoliation differed from branch sample defoliation in 5–6 out of 7 years, consistently underestimating defoliation, across the full range of defoliation severity observed. Nested mixed-effect models for fir–spruce combined, balsam fir, white spruce, and black spruce ocular survey defoliation bias resulted in marginal R2 of 0.40, 0.47, 0.82, and 0.86, respectively. Current defoliation severity and its interaction with previous year defoliation and weather conditions significantly affected ocular survey bias. Correspondence of aerial survey estimates and mean plot defoliation occurred in only 43% of all plot-years and ranged from 14%–58% in individual years. Differences between aerial survey defoliation and plot values mainly resulted from assigning an adjacent class (e.g., light &lt;30% assigned as moderate 31%–70% defoliation) or misplaced defoliation polygon boundaries, suggesting that assignment of aerial survey defoliation to plots or specific ground areas needs ground truth sampling.

Long-insert sequence capture detects high copy numbers in a defence-related beta-glucosidase gene βglu-1 with large variations in white spruce but not Norway spruce

Conifers are long-lived and slow-evolving, thus requiring effective defences against their fast-evolving insect natural enemies. The copy number variation (CNV) of two key acetophenone biosynthesis genes Ugt5/Ugt5b and βglu-1 may provide a plausible mechanism underlying the constitutively variable defence in white spruce (Picea glauca) against its primary defoliator, spruce budworm. This study develops a long-insert sequence capture probe set (Picea_hung_p1.0) for quantifying copy number of βglu-1-like, Ugt5-like genes and single-copy genes on 38 Norway spruce (Picea abies) and 40 P. glauca individuals from eight and nine provenances across Europe and North America respectively. We developed local assemblies (Piabi_c1.0 and Pigla_c.1.0), full-length transcriptomes (PIAB_v1 and PIGL_v1), and gene models to characterise the diversity of βglu-1 and Ugt5 genes. We observed very large copy numbers of βglu-1, with up to 381 copies in a single P. glauca individual. We observed among-provenance CNV of βglu-1 in P. glauca but not P. abies. Ugt5b was predominantly single-copy in both species. This study generates critical hypotheses for testing the emergence and mechanism of extreme CNV, the dosage effect on phenotype, and the varying copy number of genes with the same pathway. We demonstrate new approaches to overcome experimental challenges in genomic research in conifer defences.