Spruce Budworm Populations Research Articles

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.

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.

Overwintering spruce budworm population density as predictor of following-year larval density and defoliation on balsam fir

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.

Nontarget impacts of insecticide-based population control of eastern spruce budworm (Lepidoptera: Tortricidae) on nontarget caterpillar communities and parasitism

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.

Not just for the birds: Spiders as natural enemies of spruce budworm (Choristoneura fumiferana, Clem.)

Abstract(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.

Spatiotemporal patterns of a structured spruce budworm diffusive model

We formulate and analyze a general reaction-diffusion equation with delay, inspired by age-structured spruce budworm population dynamics with spatial diffusion by matured individuals. The model has its particular feature for bistability due to the incorporation of a nonlinear birth function (Ricker's function) and a Holling type function of predation by birds. Here we establish some results about the global dynamics, in particular, the stability of and global Hopf bifurcation from the spatially homogeneous steady state when the maturation delay is taken as a bifurcation parameter. We also use a degree theoretical argument to identify intervals for the diffusion rate when the model system has a spatially heterogeneous steady state. Numerical experiments presented show interesting spatialtemporal patterns.

Modeling weather-driven long-distance dispersal of spruce budworm moths (Choristoneura fumiferana). Part 1: Model description

Long-term studies of insect populations in the North American boreal forest have shown the vital importance of long-distance dispersal to the maintenance and expansion of insect outbreaks. In this work, we extend several concepts established previously in an empirically-based dispersal flight model with recent work on the physiology and behavior of the adult eastern spruce budworm (SBW) moth, Choristoneura fumiferana (Clemens). An outbreak of defoliating SBW in Quebec, ongoing since the mid-2000s, already covers millions of hectares of forests in eastern Canada and threatens to spread into neighboring areas through annual summertime episodes of long-distance dispersal. Such flight events in favorable conditions frequently include billions of SBW moths dispersing in the warm atmospheric boundary layer, typically starting around sunset and often lasting through several hours of wind-driven transport over hundreds of kilometers. Successful SBW dispersal to possibly distant host forest areas depends acutely on the weather. Here we describe the components and results of SBW–pyATM, an open-source individual-based modeling framework developed in Python for the simulation of these weather-driven SBW dispersal events. Using seasonal SBW phenology results from BioSIM at known outbreak locations and high-resolution Weather Research and Forecasting (WRF) model output, we focus on modeling dispersal flights over two successive nights in July 2013 in southern Quebec. Our flight model closely reproduces the SBW spatial patterns and motions observed by weather surveillance radar over the St. Lawrence estuary. With SBW–pyATM we can estimate landing locations for both male and female SBW and the resulting spatial patterns of egg distribution, allowing us eventually to forecast future larval defoliation activity in new locations where immigration could help overcome local limitations on SBW populations. This information could then support forest management decisions where SBW outbreaks threaten valuable resources.

Dynamic Analysis of a Model for Spruce Budworm Populations with Delay

A class of delayed spruce budworm population model is considered. Compared with previous studies, both autonomous and nonautonomous delayed spruce budworm population models are considered. By using the inequality techniques, continuation theorem, and the construction of suitable Lyapunov functional, we establish a set of easily verifiable sufficient conditions on the permanence, existence, and global attractivity of positive periodic solutions for the considered system. Finally, an example and its numerical simulation are given to illustrate our main results.

Avian Predation in a Declining Outbreak Population of the Spruce Budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae).

Simple SummaryCages preventing access to birds were used to measure the rate of predation by birds in a spruce budworm population during the decline of an outbreak. Three species of budworm-feeding warblers were involved in this predation on larvae and pupae. It was found that bird predation is a very important source of mortality in declining spruce budworm populations, and that bird foraging behavior changes as budworm prey become rare at the end of the outbreak.The impact of avian predation on a declining population of the spruce budworm, Choristoneura fumifereana (Clem.), was measured using single-tree exclosure cages in a mature stand of balsam fir, Abies balsamea (L.), and white spruce, Picea glauca (Moench.) Voss. Bird population censuses and observations of foraging and nest-feeding activity were also made to determine the response of budworm-linked warblers to decreasing food availability. Seasonal patterns of foraging. as well as foraging success in the declining prey population was compared to similar information from birds observed in another stand where the spruce budworm population was rising. Avian predation was an important source of mortality between the 4th instar and moth emergence in the declining outbreak population. Mortality by predation increased from negligible to over 98% as budworm density dropped from 100 to <1 larva/kg of host foliage, over 3 years. Calculations based on nest-feeding activity and basic metabolic demands support these observed rates. Seasonal and yearly differences in predation rates observed between the two host-tree species correspond to equivalent shifts in bird foraging behavior in response to dropping insect density. In particular, a preference for searching on white spruce disappeared, although budworm-linked birds remained more efficient at finding food on this plant. The ability to change foraging behavior as prey density dropped differed between bird species.

Parasitoid community responds indiscriminately to fluctuating spruce budworm (Lepidoptera: Tortricidae) and other caterpillars on balsam fir (Pinaceae)

AbstractThe world is astoundingly variable, and organisms – from individuals to whole communities – must respond to variability to survive. One example of nature’s variability is the fluctuations in populations of spruce budworm, Choristoneura fumiferana Clemens (Lepidoptera: Tortricidae), which cycle every 35 years. In this study, we examined how a parasitoid community altered its parasitism of budworm and other caterpillar species in response to these fluctuations. Budworm and other caterpillar species were sampled from balsam fir (Pinaceae) in three plots for 14 years in Atlantic Canada, then were reared to identify any emerging parasitoids. We found that the parasitoid community generally showed an indiscriminate response (i.e., no preference, where frequencies dictated parasitism rates) to changes in budworm frequencies relative to other caterpillar species on balsam fir. We also observed changes in topology and distributions of interaction strengths between the parasitoids, budworm, and other caterpillar species as budworm frequencies fluctuated. Our study contributes to the hypothesis that hardwood trees are a critical part of the budworm–parasitoid food web, where parasitoids attack other caterpillar species on hardwood trees when budworm populations are low. Taken together, our results show that a parasitoid community collectively alters species interactions in response to variable budworm frequencies, thereby fundamentally shifting food-web pathways.

Multitype bistability and long transients in a delayed spruce budworm population model

Spruce budworm is a major defoliator of forests in North America and its periodic outbreak can cause severe economic growth loss. In 2008, Vaidya and Wu proposed a delayed spruce budworm population model and studied the impact of physiological structure on outbreak control (Vaidya and Wu, 2008 [27]). In this paper, we revisit this model and study its global dynamics that has not been analyzed previously. By carefully dealing with the major difficulty caused jointly by the time delay and the nonlinearity, we give a rather complete stability analysis for the model, including global asymptotic stability, bistability, semi-stability and Hopf bifurcation. Moreover, we show that the time delay can induce interesting dynamics including multitype bistability and long transients.

From branch to bench: establishing wild spruce budworm populations into laboratory colonies for the exploration of local adaptation and plasticity

AbstractSpruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae), is a destructive defoliator found throughout the Nearctic boreal forest. This pest has a broad geographic range and shows regional variation in key life history traits. These population differences may represent important adaptations to local environmental conditions and reflect underlying genetic diversity. Existing laboratory colonies of spruce budworm do not capture this regional variation, so we established five new spruce budworm colonies from across its range to explore regional adaptations among spruce budworm populations within common garden experiments. We present methods for establishing new spruce budworm laboratory colonies from wild populations. We describe the process of flushing, rearing, and disease screening used on these new populations to produce healthy disease-free laboratory stocks.

Continent-wide population genomic structure and phylogeography of North America's most destructive conifer defoliator, the spruce budworm (Choristoneura fumiferana).

The spruce budworm, Choristoneura fumiferana, is presumed to be panmictic across vast regions of North America. We examined the extent of panmixia by genotyping 3,650 single nucleotide polymorphism (SNP) loci in 1975 individuals from 128 collections across the continent. We found three spatially structured subpopulations: Western (Alaska, Yukon), Central (southeastern Yukon to the Manitoba–Ontario border), and Eastern (Manitoba–Ontario border to the Atlantic). Additionally, the most diagnostic genetic differentiation between the Central and Eastern subpopulations was chromosomally restricted to a single block of SNPs that may constitute an island of differentiation within the species. Geographic differentiation in the spruce budworm parallels that of its principal larval host, white spruce (Picea glauca), providing evidence that spruce budworm and spruce trees survived in the Beringian refugium through the Last Glacial Maximum and that at least two isolated spruce budworm populations diverged with spruce/fir south of the ice sheets. Gene flow in the spruce budworm may also be affected by mountains in western North America, habitat isolation in West Virginia, regional adaptations, factors related to dispersal, and proximity of other species in the spruce budworm species complex. The central and eastern geographic regions contain individuals that assign to Eastern and Central subpopulations, respectively, indicating that these barriers are not complete. Our discovery of previously undetected geographic and genomic structure in the spruce budworm suggests that further population modelling of this ecologically important insect should consider regional differentiation, potentially co‐adapted blocks of genes, and gene flow between subpopulations.

Previous year outbreak conditions and spring climate predict spruce budworm population changes in the following year

We determined effects of local spruce budworm (Choristoneura fumiferana Clem.; SBW) population level, proximity to sites with high SBW populations, insecticide spray, and environmental variables on SBW populations from 2014 to 2018, the outbreak initiation period in northern New Brunswick, Canada. SBW second instar larvae (L2) per branch data collected at 1100–2000 sample points per year were used to create annual interpolated population rasters. Fishnet sample points extracted from these rasters were overlaid with georeferenced layers of 46 possible predictor variables including forest composition, climate, topography, site quality, and insecticide treatment. Results showed that local SBW population in the previous year, proximity to sites with high SBW populations, and early spring climate were consistently the most important predictors over the 5 study years. Simultaneous autoregressive models were used to address spatial autocorrelation when forecasting the SBW L2 population, and a linear mixed effects model was fit to aggregate data for 2015–2018. The models reduced spatial dependence in the residuals, and explained 68–79% of variance in annual L2 levels and 53% of variance over the 4 years combined. Sensitivity analysis showed that locations with 5–10 more SBW L2 per branch than observed values, or 20–40 km closer to high population sites in the previous year could have up to 24 more L2 in the current year. Cumulative degree days in April helped to estimate the upper and lower bounds of the population. Expansion and retraction of SBW outbreak initiation were mathematically described. Understanding which variables influence SBW outbreak initiation and population level assists in design of small area target-specific insecticide spray applications and helps focus SBW L2 sampling on predicted outbreak hot spots.

A Conceptual Framework for the Spruce Budworm Early Intervention Strategy: Can Outbreaks be Stopped?

The spruce budworm, Choristoneura fumiferana, Clem., is the most significant defoliating pest of boreal balsam fir (Abies balsamea (L.) Mill.) and spruce (Picea sp.) in North America. Historically, spruce budworm outbreaks have been managed via a reactive, foliage protection approach focused on keeping trees alive rather than stopping the outbreak. However, recent theoretical and technical advances have renewed interest in proactive population control to reduce outbreak spread and magnitude, i.e., the Early Intervention Strategy (EIS). In essence, EIS is an area-wide management program premised on detecting and controlling rising spruce budworm populations (hotspots) along the leading edge of an outbreak. In this article, we lay out the conceptual framework for EIS, including all of the core components needed for such a program to be viable. We outline the competing hypotheses of spruce budworm population dynamics and discuss their implications for how we manage outbreaks. We also discuss the practical needs for such a program to be successful (e.g., hotspot monitoring, population control, and cost–benefit analyses), as well as the importance of proactive communications with stakeholders.

The Impact of Moth Migration on Apparent Fecundity Overwhelms Mating Disruption as a Method to Manage Spruce Budworm Populations

Aerial applications of a registered formulation of synthetic spruce budworm female sex pheromone were made in 2008, 2013 and 2014 to disrupt mating in populations of this forest insect pest in Quebec, Canada. Each year, the applications resulted in a 90% reduction in captures of male spruce budworm moths in pheromone-baited traps. A commensurate reduction in mating success among virgin females held in individual cages at mid-crown of host trees was also obtained. However, there was no reduction in the populations of eggs or overwintering larvae in the following generation (late summer and fall). The failure of this approach as a viable tactic for spruce budworm population reduction could have resulted from considerable immigration of mated females, as evidenced by high rates of immigration and emigration that caused steep negative relationships between apparent fecundity and the density of locally emerged adults.

Dynamics and Management of Rising Outbreak Spruce Budworm Populations

Management of spruce budworm, Choristoneura fumiferana (Clem.), outbreak spread requires understanding the demographic processes occurring in low, but rising populations. For the first time, detailed observations were made in the early stages of outbreak development. We sampled populations over a three-year period in both treated and untreated populations in the Lower St-Lawrence region of Quebec, Canada, and measured the density-dependence of survival and population growth rates, and the impact of natural enemies and insecticides. Insecticides tested were Bacillus thuringiensis (Berliner 1915) and tebufenozide. We recorded strong density-dependence of survival between early larval stages and adult emergence, explained largely by the variation of natural enemy impacts and overcrowding. We also observed inverse density-dependence of apparent fecundity: net immigration into lower-density populations and net emigration from the higher, linked to a threshold of ~25% defoliation. Because of high migration rates, none of the 2013 treatments reduced egg populations at the end of summer. However lower migration activity in 2014 allowed population growth to be reduced in treated plots. This evidence lends support to the conclusion that, for a budworm population to increase to outbreak density, it must be elevated via external perturbations, such as immigration, above a threshold density of ~4 larvae per branch tip (L4). Once a population has increased beyond this threshold, it can continue growing and itself become a source of further spread by moth migration. These findings imply that populations can be brought down by insecticide applications to a density where mortality from natural enemies can keep the reduced population in check, barring subsequent immigration. While we recognize that other factors may occasionally cause a population to exceed the Allee threshold and reach outbreak level, the preponderance of immigration implies that if all potential sources of significant numbers of moths are reduced on a regional scale by insecticide applications, a widespread outbreak can be prevented, stopped or slowed down by reducing the supply of migrating moths.

Influence of a Foliar Endophyte and Budburst Phenology on Survival of Wild and Laboratory-Reared Eastern Spruce Budworm, Choristoneura fumiferana on White Spruce (Picea glauca)

A manipulative field study was carried out to determine whether the foliar endophyte fungus, Phialocephala scopiformis DAOM 229536, decreased the performance of eastern spruce budworm, Choristoneura fumiferana larvae developing on white spruce trees. Overwintered second-instar budworm larvae from a laboratory colony or from a wild population were placed on endophyte positive or negative trees one or two weeks before budburst. The presence of the endophyte in the needles reduced the survival of C. fumiferana from both a wild population and a laboratory colony. Survival for budworm juveniles up to pupation and to adult emergence was 13% and 17% lower, respectively, on endophyte positive trees. The endophyte did not influence the size or sex of survivors and budworm survival was not influenced by any two- or three-way interactions. Budworm survival was higher for wild than for laboratory-reared budworm and for budworm placed on trees a week before budburst. This may be the first field study to demonstrate the efficacy of an endophytic fungus against wild individuals of a major forest insect pest. The efficacy of the endophyte at low larval densities suggests that it could be a useful tactic to limit spruce budworm population growth in the context of an early intervention strategy.

Insights into the Structure of the Spruce Budworm (Choristoneura fumiferana) Genome, as Revealed by Molecular Cytogenetic Analyses and a High-Density Linkage Map.

Genome structure characterization can contribute to a better understanding of processes such as adaptation, speciation, and karyotype evolution, and can provide useful information for refining genome assemblies. We studied the genome of an important North American boreal forest pest, the spruce budworm, Choristoneura fumiferana, through a combination of molecular cytogenetic analyses and construction of a high-density linkage map based on single nucleotide polymorphism (SNP) markers obtained through a genotyping-by-sequencing (GBS) approach. Cytogenetic analyses using fluorescence in situ hybridization methods confirmed the haploid chromosome number of n = 30 in both sexes of C. fumiferana and showed, for the first time, that this species has a WZ/ZZ sex chromosome system. Synteny analysis based on a comparison of the Bombyx mori genome and the C. fumiferana linkage map revealed the presence of a neo-Z chromosome in the latter species, as previously reported for other tortricid moths. In this neo-Z chromosome, we detected an ABC transporter C2 (ABCC2) gene that has been associated with insecticide resistance. Sex-linkage of the ABCC2 gene provides a genomic context favorable to selection and rapid spread of resistance against Bacillus thuringiensis serotype kurstaki (Btk), the main insecticide used in Canada to control spruce budworm populations. Ultimately, the linkage map we developed, which comprises 3586 SNP markers distributed over 30 linkage groups for a total length of 1720.41 cM, will be a valuable tool for refining our draft assembly of the spruce budworm genome.

Do natural enemies explain fluctuations in low-density spruce budworm populations?

Understanding the causal pathways through which forest insect outbreaks are triggered is important for resource managers. However, detailed population dynamics studies are hard to conduct in low-density, pre-outbreak populations because the insects are difficult to sample in sufficient numbers. Using laboratory-raised larvae installed in the field across a 1,000km east-west gradient in Québec (Canada) over an 11-yr period, we examined if parasitism and predation were likely to explain fluctuations in low-density spruce budworm (Choristoneura fumiferana; SBW) populations. Parasitism rates by the two main larval parasitoid species, Elachertus cacoeciae and Tranosema rostrale, peaked during different years. This suggests that temporal fluctuations in overall parasitism were partly buffered by compensatory dynamics among parasitoid species. Still, spatial covariance analyses indicate that the residual interannual variation in parasitism rates was substantial and correlated over large distances (up to 700km). On the other hand, interannual variation in predation rates was not spatially correlated. Piecewise structural equationmodels indicate that temporal variation in parasitism and predation does not influence temporal variation in wild SBW abundance. Spatially, however, SBWs installed in warmer locations tended to show higher parasitism rates, and these higher rates correlated with lower wild SBW population levels. Overall, the results indicate that large-scale drops in parasitism occur and could potentially contribute to SBW population increases. However, during the period covered by this study, other factors such as direct effects of weather on SBW larval development or indirect effects through host tree physiology or phenology were more likely to explain large-scale variation in wild SBW populations.