The spruce budworm (Choristoneura fumiferana; SBW) is a periodically outbreaking forest insect pest that affects the boreal forests of North America through extensive defoliation and tree mortality. Causes of widespread spatial synchrony of SBW outbreaks remain a key question in the ecology and management of this species. While the Moran effect (correlated favourable environmental conditions) and density-dependent dispersal (from epicentres of demographic explosions) have been proposed and supported as drivers of synchronised outbreaks, the relative contribution of long-distance dispersal is still poorly understood. In this study, we use a novel approach to distinguish resident from migrant moths and to assign migrants to likely source clusters with the goal of better characterising regional dispersal. First, we characterise the genetic diversity and structure of resident SBW larvae and three phenologically separated groups of moths over one flight season using Genotyping-by-Sequencing. Then, using a novel machine learning approach, we assign putative migrants to their likely source populations. We hypothesised that migrant moths and resident larvae would be genetically distinct and could be assigned to source populations. Our findings revealed complex patterns of moth dispersal and population differentiation within a single season, including two spatially overlapping genetic clusters. We observed subtle but significant genetic differences between resident larvae and migrant moths, supporting the hypothesis that long-distance dispersal contributes to outbreak dynamics and synchrony. These insights enhance our understanding of SBW population dynamics and suggest that effective management strategies, such as the Early Intervention Strategy (EIS), must account for the role of dispersal in mitigating the detrimental effects of major outbreaks.

Spruce budworm (SBW; Choristoneura fumiferana Clem.) outbreaks are an important natural disturbance in North America, killing trees over millions of hectares. We related 11 years of SBW defoliation in 87 plots in Gaspé Peninsula, Québec, to 23 stand, site, and climate variables. Defoliation was consistently ordered among host species: balsam fir > white spruce > black spruce. Within the relatively small 200 km2 study area, cluster analyses resulted in four and 10 clusters for balsam fir cumulative and current defoliation, respectively; variation in cumulative defoliation converged over 11 years. Current defoliation was significantly spatially autocorrelated among plots within stands, but autocorrelation weakened at distances >2500m. Cumulative defoliation was significantly related to insecticide spraying, minimum and maximum summer temperature, and interactions between SBW larvae per branch versus hardwood and white spruce basal area. Tree species, insecticide spraying, and number of defoliating SBW larvae were the main determinants of defoliation. Results showed much higher local spatial variability in current defoliation patterns than previous studies, but over the course of an outbreak, cumulative defoliation patterns converged. Cumulative defoliation patterns similar to these, assigned based on local defoliation severity, can be input into defoliation-based growth models to predict impacts on growth and survival.

The concepts of compensation and additive mortality form the ecological basis for understanding animal population responses to exploitation by humans. In the context of pest management, compensation is a density‐dependent response that allows populations to offset control‐related mortality, often via increased survival or reinvasion. Additive mortality, in contrast, accrues when a population's compensatory capacity is insufficient to offset losses, resulting in a net reduction in population size or growth rate. These concepts are rarely considered in forest insect pest management, which tends to emphasise short‐term plant protection over long‐term population control. We used published life table data for a major native forest insect defoliator, the spruce budworm ( Choristoneura fumiferana [Lepidoptera: Tortricidae]) to simulate the amount of additive mortality required to suppress an outbreak. Simulations also assessed how the failure to account for different compensatory responses could hinder successful control. Our results suggest that only relatively modest amounts of additive mortality (perhaps as low as approximately 8%–18%) may be needed to stop spruce budworm from outbreaking, with immigration being the strongest potential compensatory hindrance to outbreak suppression. Many of the compensatory responses that thwarted outbreak suppression in the past (e.g., low detection efficiency, immigration, indiscriminate killing of predators and parasitoids) have contemporary solutions that could increase additive mortality and thereby enhance the feasibility of population control strategies for native forest insect pests. Our results suggest that some native forest insect pests may require relatively little additive mortality to suppress outbreaks if compensation‐limiting strategies are used. Incorporating theoretical and strategic frameworks used in vertebrate population management could advance the development of native insect population control programmes.

(1) The eastern spruce budworm ( Choristoneura fumiferana , Clem.) is a native irruptive forest pest that defoliates spruce‐fir forests throughout North America's boreal zone. (2) Past studies suggest that successful spruce budworm population control requires high natural mortality from a variety of sources, including predators, especially from parasitoids and birds. While well represented in many different ecosystems, the role of generalist predatory spiders in these boreal systems remains largely unstudied. (3) To determine the identity and percentage of spiders preying upon spruce budworm, we hand‐collected spiders from balsam fir ( Abies balsamea ) in stands with relatively high spruce budworm densities from forests in insular Newfoundland and Labrador, Canada. (4) Using a spruce budworm‐specific TaqMan real‐time PCR assay, we successfully amplified spruce budworm DNA in 32% of collected spiders. After spider molecular barcoding, we found the web builders Grammonota angusta Dondale, Pityohyphantes (aff. subarcticus ), Dictyna brevitarsa Emerton and Estrandia grandaeva (Keyserling) represented 58% of the spiders feeding on spruce budworm, and the wandering hunter Philodromus rufus Dondale represented 11.8%. (5) Our molecular approach was an effective means with which to identify recently consumed prey and natural enemies in this boreal system.

Pesticides have been used in Canada since 1945 as part of large-scale aerial spray applications to control insect pests on forested lands. Some of the pesticides used historically were efficacious, nonselective, persistent, and have led to serious impacts on the environment. A well known, and extensively documented example is the large-scale aerial spray programs in New Brunswick, Canada. From 1952 to 1993, 97% of the 6.2 million ha of the forested lands of New Brunswick were treated with at least one application of one insecticide, the majority of which were applied to control outbreaks of eastern spruce budworm (Choristoneura fumiferana). The most well known insecticide was dichlorodiphenyltrichloroethane (DDT), applied from 1952 to 1968, which still persists in treated soils and adjacent water bodies, and caused the individual and cumulative ecosystem effects that can still be measured today. The insecticides that replaced DDT were nonpersistent and unlikely to be found today. However, during the years of application some of the insecticides were likely to have impacted local ecosystems to some degree. To aid future studies on the efficacy and environmental impact of these insecticides we created a digital spatial data set of known pesticide application in New Brunswick forestry from 1952 to 1993. The data set includes active ingredient, formulation, application rate, tank mix, aircraft type, and other ancillary information. The current version of the data is available on the New Brunswick Department of Natural Resources and Energy Development, GIS Open Data Page and in the supplemental material. Use of the data set for academic and educational purposes is encouraged, provided that both this data paper and the data source are properly cited; the Government of New Brunswick should be acknowledged as the data source (Open Government License http://www.snb.ca/e/2000/data-E.html).

The Early Intervention Strategy (EIS) was developed to control spruce budworm (SBW) populations before an outbreak develops. It uses an action threshold population density originally based on feeding fourth-instar larvae (L4) sampled in spring. But in practice, in all SBW management strategies, it is the density of overwintering second-instar larvae (L2) sampled in the previous fall that is used in decision making. Those densities serve to predict defoliation and plan control operations for the following summer. Therefore, there was a need to establish a relationship between L2 and L4 (apparent survival) by which to adequately express treatment thresholds in terms of L2 rather than L4. This was done with samples collected in each life stage in several sites in Quebec and New Brunswick and over three different periods between 1999 and 2022. We found that the distribution of L2 is more aggregated than that of L4, and thus that sample sizes needed for precise density estimation in that stage are larger than at the L4. We also found that apparent survival between L2 and L4 varies at low and high L2 densities for different reasons. For example, movement of larvae from the inner crown, bole, or non-host trees affect apparent survival at the lowest population densities. In contrast, at very high densities survival of populations during these early larval stages is reduced by crowding effects. At densities in the mid-range where control decisions need to be made, apparent survival is very close to 100% and varies little among years. The current threshold density is 3.5 L4 per 45-cm branch tip and was established as 7 L2 per 75-cm branch by assuming 50% mortality. Considering the results presented here, we recommend that the current EIS intervention threshold be set at 3.5 L2 per 75-cm branch. In this density, defoliation typically ranges between 15 and 30% in the following summer in the absence of treatment.

Abstract Narrow-spectrum insecticides are currently used to control populations of spruce budworm, Choristoneura fumiferana Clemens (Lepidoptera: Tortricidae), in eastern Canada. However, these could have nontarget impacts on other caterpillars – some of which may serve as alternative or alternate hosts to key parasitoids – that are also susceptible to control tactics. This study was conducted to determine how the insecticides, Bacillus thuringiensis variety kurstaki (Btk) and tebufenozide, used to control spruce budworm populations, impact caterpillar communities and associated parasitism rates. Post-treatment field sampling of caterpillars was conducted in 2018 and 2019 in New Brunswick, Canada, at sites treated with either Btk or tebufenozide and at control sites. Caterpillar species richness and abundance, community structure, and parasitism rates were assessed using molecular analyses for 659 collected caterpillars. We found that insecticide applications had no significant impact on abundance, species richness, or parasitism rate relative to the measurements made in the control sites. Nonetheless, a significantly higher caterpillar abundance and lower parasitism rate occurred in Btk-treated sites than in tebufenozide-treated sites. Overall, however, Btk and tebufenozide treatments did not negatively affect the non-budworm caterpillar community under the present conditions of low caterpillar densities, suggesting that parasitoids have alternative and alternate hosts after treatments that target the spruce budworm.

Pest density – plant damage relationships are essential guides for decision-making in integrated pest management. In this article, we established pest density – leaf damage relationships for the beech leaf-mining weevil, Orchestes fagi L. (formerly Rhynchaenus fagi, Coleoptera: Curculionidae), in its invasive range of Nova Scotia, Canada. Outbreaks of O. fagi cause tree-wide leaf necrosis in American beech (Fagus grandifolia Ehrh.), which can eventually result in tree mortality. In 2014 and 2016, we collected weekly samples in stands with American beech and assessed leaves for densities during different life stages (eggs, larvae, and pupae), population proxy measures (adult feeding damage, egg slits, and larval galleries), and percent necrosis. In general, feeding damage and leaf necrosis plateaued soon after the end of budburst, but before the larval mine expanded. This strongly suggested that leaf necrosis may be linked to damage caused by adults or by mine initiation rather than that caused by larval mine expansion and gallery development. The density of O. fagi per leaf for life stages and population proxies all significantly explained ∼42%–81% of the variation in end-of-season percent leaf necrosis. Results from this study provide a variety of relationships that could be used in both short- and long-term monitoring efforts for O. fagi.

Large-scale monitoring is used to track population trends for many ecologically and economically important wildlife species. Often, population monitoring involves professional staff travelling to collect data (i.e., conventional monitoring) or in efforts to reduce monitoring costs, by engaging volunteers (i.e., community science). Although many studies have discussed the advantages and disadvantages of conventional vs. community science monitoring, few have made direct, quantitative comparisons between these two approaches. We compared data quality and financial costs between contemporaneous and overlapping conventional and community science programs for monitoring a major forest pest, the spruce budworm ( Choristoneura fumiferanae Clem.). Although community science trapping sites were clumped around urban areas, abundance estimates from the programs were strongly spatially correlated. However, annual program expenditures were nearly four times lower in the community science versus the conventional program. We modelled a hypothetical hybrid model of the two programs, which provided full spatial coverage and potentially the same data, but at half the cost of the conventional program and with the added opportunity for public engagement. Our study provides a unique quantitative analysis of merits and costs of conventional versus community science monitoring. Our study offers insights on how to assess wildlife monitoring programs where multiple approaches exist.

Outbreaking insects often undergo rapid population growth synchronously in multiple locations separated by large distances. Dispersal may play an essential role in synchronizing outbreaks over large geographic areas but its role in outbreak spread remains unclear. In our study, we used population genetics to assess how much dispersal contributes to the spread of a major forest pest, the spruce budworm (Choristoneura fumiferana Clemens). Using spatial analyses of genotypic data, we determined the extent to which dispersing individuals from epidemic populations in Quebec leave their genetic signature in adjacent, rising populations in New Brunswick and Maine. A lack of genetic differentiation between endemic and epidemic populations would indicate that outbreak spread depends on effective dispersal from epidemic to endemic regions. In contrast, endemic populations that are distinct from epidemic population would suggest that dispersal plays a reduced role in fomenting outbreak rise. An intermediate possibility with differentiation between epidemic and endemic population as a continuous spatial gradient might suggest a temporal lag in genetic differentiation that may take the form of a genetic travelling wave. PCA, cluster analyses, isolation by distance, and sPCA were used to characterize spatial genomic variation using 300 SBW larvae sampled in 2015 and genotyped at thousands of genome-wide SNPs. We found support for a genetic travelling wave pattern, matching the SBW density pattern. These results indicate that dispersal may trigger the transition of populations from the endemic to the epidemic state and thereby drive outbreak spread. By supplementing persisting endemic populations, migrants may drive populations to beyond levels that can be controlled by local biotic constraints. These results support the underlying justification of ongoing outbreak containment efforts in Atlantic Canada (i.e., the Early Intervention Strategy), although continued sampling as the outbreak progresses is needed confirm the temporal stability of the observed patterns.

The beech leaf-mining weevil, Orchestes fagi (L.), is native to Europe where it commonly attacks European beech. The weevil was discovered infesting American beech in Halifax and Cape Breton Island, Nova Scotia, Canada in 2012, but anecdotal reports of defoliated beech in the Halifax area as early as 2006 suggest it established 5–10 years prior to its discovery. Our objectives were to estimate the impact of O. fagi on American beech in forested sites and urban areas, as well as its economic impact on owners of residential properties with mature American beech. In 2014, we established fifteen plots in forested sites containing a total of 260 American beech at Sandy Lake, Oakfield, and Mount Uniacke (n = 5 plots per site), where weevil infestation levels were moderate, low, and nil, respectively. At the same time we recorded the degree of cankering by beech bark disease on the main stems of each tree. Plots were visited annually to record tree mortality (2014–2019) and percentage of leaves with larval mines or adult feeding (2016–2019). Between 2016 and 2019, the percentage of leaves mined by weevil larvae increased from 6 to 59% at Mount Uniacke and from 48 to 83% at Oakfield. During the same period, cumulative beech mortality increased from 35 to 48% at Mount Uniacke and from 10 to 70% at Oakfield. At Sandy Lake in 2016, 88% of the beech trees had died and there were too few living beech to collect a leaf sample in our plots so estimates of weevil damage (87% of leaves with mines) were obtained from life table plots in the same area. Tree mortality was associated with severity of cankering by beech bark disease only at Mount Uniacke, the site with the fewest years of defoliation by the leaf-mining weevil. We also surveyed residents of Halifax in 2016 and 2018 to determine the rate of beech mortality and costs of tree removal in urban residential areas in the same region (within 40 km) of the forest areas. Relative to the forested sites at Sandy lake and Oakfield, mortality rates were lower in urban areas (32% in 2016, 44% in 2018), even though signs of weevil defoliation had been apparent to residents as early as 2011–2012. Direct costs ($CAN) to property owners who hired arborists to remove dead beech trees averaged $1934 ($300–$6600) per resident in 2018. Options for mitigating the impact of O. fagi on American beech are briefly discussed.

Annual monitoring of mortality agents in the course of a spruce budworm (Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae)) population cycle is essential to understanding the factors governing the rise and collapse of outbreaks. To date, assessments of causes of budworm mortality have relied on laboratory rearing of field-collected larvae, followed by visual identification of emerging parasitoids and/or microscopic analysis of pathogens in larval carcasses. Although this approach has provided vital information on the abundance and identity of mortality agents, the procedure is labor-intensive and has limits in terms of accuracy. To overcome these shortcomings, we developed a molecular identification tool that makes use of real-time quantitative PCR (qPCR) and TaqMan® technologies. The tool relies on taxon-specific molecular variants (single nucleotide polymorphism [SNP] markers) found in mitochondrial (COI) and nuclear (28S rDNA) genes, for parasitoids, and in the nuclear SSU rDNA gene for microsporidian pathogens; these are then used as molecular signatures targeted by qPCR primers and TaqMan probes. Thus, the design of several sets of primers and probes deployed in multiplex format enables the identification of natural enemies via a molecular sorting process, bypassing barcode sequencing. Crude budworm DNA extracts are processed through a first module that detects dipteran and hymenopteran parasitoids, and microsporidian infections. Positive samples are then processed for species determination using three additional modules, enabling the identification of 20 common natural enemies of the spruce budworm. The tool has been fully validated using DNA samples from all comprised taxa, and both its sensitivity and accuracy compared favorably with the rearing-based method in an analysis of field-collected budworms. Using this tool, sample processing can be completed within two days, does not require larval rearing, provides accurate species identification, and can be conducted by technical staff without extensive molecular biology or insect taxonomy training.

Spruce budworm (Choristoneura fumiferana Clem.) defoliation of balsam fir (Abies balsamea (L.) Mill.) has been shown to be less severe in stands with higher hardwood (broadleaved trees) content during both building and declining phases of budworm outbreaks. Reduced defoliation associated with forest composition could be explained by habitat fragmentation and/or natural enemy hypotheses, which posit higher early-instar larval dispersal loss or more intense parasitism in hardwood-softwood mixed stands than in pure balsam fir stands, respectively. We carried out field studies in 27 hardwood, mixedwood, and softwood plots with varied balsam fir-hardwood composition to assess effects of forest composition on: 1) first- and second-instar larval dispersal loss, and 2) stage-specific spruce budworm density and parasitism rates. Results indicated that increasing hardwood content increased second-instar (L2) dispersal losses. Stand type significantly affected dispersal loss of L2 larvae, both measured directly for three years and sampled on balsam fir seedlings. Post-hoc pair-wise comparisons indicated that L2 dispersal loss was less in softwood than in mixedwood or hardwood stand types based on measurements on ground traps and less in softwood and mixedwood than in hardwood stands based on sampling budworm larvae on regeneration. This suggested that high early-instar larval dispersal loss is a plausible explanation for the decreased spruce budworm density and balsam fir defoliation associated with increasing hardwood content. Stand type did not significantly affect parasitism rates.

Abstract The egg parasitoid Trichogramma spp. (Hymenoptera: Trichogrammatidae) is a widely used biocontrol agent against lepidopteran pests. Historically, Trichogramma were deployed either by plane or by using cardboard cards on which parasitized eggs are glued and manually installed at sites. Plane deployment is costly and card installation is time consuming, but the use of Trichogramma has been shown to be efficient against several pests. In 2016 and 2017, a research project investigated the potential use of unmanned aerial system for distributing Trichogramma as biocontrol agents against two major pests: an agricultural pest of maize, the European corn borer, Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae), and a forest pest, the eastern spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera:Tortricidae). Exposure duration of parasitized eggs to field conditions (temperature, predation, etc.) in maize fields influenced the Trichogramma's emergence rate, suggesting that timing of parasitoid releases with their emergence is essential. Although parasitism of naturally occurring eggs in maize fields could not be compared due to the low density of the European corn borer, parasitism of sentinel eggs by Trichogramma was more prominent in plots with unmanned aircraft systems (UAS)-releases compared to control plots. For spruce budworm, treatment with Trichogramma increased egg parasitism and there was no difference between the deployment by UAS and by Trichocards. We discuss these results in the context of pest biology and management. We also discuss the advantages and shortcomings of both methods and offer insights into where future work might go to further leverage the use of UAS in managing these important pests.

Insect outbreaks can cover vast geographic areas making it onerous to cost-effectively monitor populations to address management or ecological questions. Community science (or citizen science), which entails engaging the public to assist with data collection, provides a possible solution to this challenge for the spruce budworm ( Choristoneura fumiferana Clemens), a major defoliating pest in North America. Here, we lay out the Budworm Tracker Program, a contributory community science program developed to help monitor spruce budworm moths throughout eastern Canada. The program outsources free pheromone trap kits to volunteers who periodically check and collect moths from their traps throughout the budworm flight period, then return them in a prepaid envelope to the organizers. Over three years, the program engaged an average of 216–375 volunteers and yielded a data return rate of 68%–89%, for a total of 16 311–54 525 moths per year. Volunteer retention among years was 71%–89%. Data from this program offer compelling evidence for the range of long-distance moth dispersal. Although our program was designed for spruce budworm, this template could easily be adapted for forestry, urban forestry, and agricultural systems to monitor any of the numerous organisms for which there is an established trapping method.

Abstract Spatial synchrony is a common characteristic of spatio‐temporal population dynamics across many taxa. While it is known that both dispersal and spatially autocorrelated environmental variation (i.e., the Moran effect) can synchronize populations, the relative contributions of each, and how they interact, are generally unknown. Distinguishing these mechanisms and their effects on synchrony can help us to better understand spatial population dynamics, design conservation and management strategies, and predict climate change impacts. Population genetic data can be used to tease apart these two processes as the spatio‐temporal genetic patterns they create are expected to be different. A challenge, however, is that genetic data are often collected at a single point in time, which may introduce context‐specific bias. Spatio‐temporal sampling strategies can be used to reduce bias and to improve our characterization of the drivers of spatial synchrony. Using spatio‐temporal analyses of genotypic data, our objective was to identify the relative support for these two mechanisms to the spatial synchrony in population dynamics of the irruptive forest insect pest, the spruce budworm (Choristoneura fumiferana), in Quebec (Canada). AMOVA, cluster analysis, isolation by distance, and sPCA were used to characterize spatio‐temporal genomic variation using 1,370 SBW larvae sampled over four years (2012–2015) and genotyped at 3,562 SNP loci. We found evidence of overall weak spatial genetic structure that decreased from 2012 to 2015 and a genetic diversity homogenization among the sites. We also found genetic evidence of a long‐distance dispersal event over >140 km. These results indicate that dispersal is the key mechanism involved in driving population synchrony of the outbreak. Early intervention management strategies that aim to control source populations have the potential to be effective through limiting dispersal. However, the timing of such interventions relative to outbreak progression is likely to influence their probability of success.

Eruptive insect pests have significant impacts on the structure and function of forest ecosystems. Outbreaks of spruce budworm (Choristoneura fumiferana) (Clem.), for instance, occur approximately every 35-40 years, resulting in the loss of millions of hectares of spruce-fir forests in eastern North America. In addition to the density-dependent relationships that drive insect outbreaks, rising populations often coincide with drought events, which are expected to increase in both frequency and intensity in response to climate change. However, as populations approach the eruptive phase, consequences of intraspecific competition may outweigh the benefit of host water stress. The objective of our study was to quantify defoliation and insect performance responses to the interactive effects of drought and density of spruce budworm. To test for these interactions, we established a manipulative field experiment in a mature, balsam fir-dominated forest stand using a combination of single-tree rainout shelters and sleeve-caged insect larvae at four different densities: 0, 25, 50, or 100 individuals. Defoliation of 1-year old shoots, but not current-year shoots, significantly increased in response to higher insect densities. Density also had a significant, negative effect on budworm percent survival, although the total number of recovered adults remained highest in the high density treatment. Adult female body mass was significantly reduced in response to increased density, but only on droughted trees. Lastly, male wing length was significantly decreased in response to increased density. Overall, our results demonstrate that across a broad range of outbreak densities, rain exclusion had a minor impact. Accordingly, we anticipate that as insect pest populations approach epidemic levels, the influence of density on defoliation, insect survival, and body condition is likely to outweigh the impact of moderate drought stress.