Three main hypotheses have been postulated over the past century to explain the outbreaking population dynamics of eastern spruce budworm, Choristoneura fumiferana (Clemens). The Silviculture Hypothesis first arose in the 1920s, with the idea that outbreaks were driven by forestry practices favoring susceptible softwood species. In the 1960s, it was proposed that populations were governed by Multiple Equilibria, with warm weather conditions releasing low-density populations from the regulatory control of natural enemies. Dispersal from outbreak foci, or “epicenters,” was seen as causing widespread outbreaks that eventually collapsed following resource depletion. However, in the 1980s, following the re-analysis of data from the 1940s outbreak in New Brunswick, this interpretation was challenged. The alternative Oscillatory Hypothesis proposed that budworm population dynamics were governed by a second-order density-dependent process, with oscillations being driven by natural enemy–victim interactions. Under this hypothesis, weather and resource availability contribute to secondary fluctuations around the main oscillation, and weather and moth dispersal serve to synchronize population cycles regionally. Intensive, independent population studies during the peak and declining phases of the 1980s outbreak supported the principal tenet of the Oscillatory Hypothesis, but concluded that host plant quality played a more important role than this hypothesis proposed. More recent research on the early phase of spruce budworm cycles suggests that mate-finding and natural-enemy-driven Allee effects in low-density populations might be overcome by immigration of moths, which can facilitate the onset of outbreaks. Even more recent research has supported components of all three hypotheses attempting to explain spruce budworm dynamics. In the midst of a new rising outbreak (2006-present), we discuss the evolution of debates surrounding these hypotheses from a historic perspective, examine gaps in current knowledge, and suggest avenues for future research (e.g., intensive studies on low-density populations) to better understand and manage spruce budworm populations.

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.

Spruce budworm (Choristoneura fumiferana Clem.; SBW) outbreaks are one of the dominant natural disturbances in North America, having killed balsam fir (Abies balsamea (L.) Mill.) and spruce (Picea sp.) trees over tens of millions of hectares. Responses to past SBW outbreaks have included the aerial application of insecticides to limit defoliation and keep trees alive, salvage harvesting of dead and dying trees, or doing nothing and accepting the resulting timber losses. We tested a new ‘early intervention strategy’ (EIS) focused on suppressing rising SBW populations before major defoliation occurs, from 2014 to 2018 in New Brunswick, Canada. The EIS approach included: (1) intensive monitoring of overwintering SBW to detect ‘hot spots’ of low but rising populations; (2) targeted insecticide treatment to prevent spread; and (3) proactive public communications and engagement on project activities and results. This is the first attempt of area-wide (all areas within the jurisdiction of the province of New Brunswick) management of a native forest insect population. The project was conducted by a consortium of government, forest industry, researchers, and other partners. We developed a treatment priority and blocking model to optimize planning and efficacy of EIS SBW insecticide treatment programs. Following 5 years of over 420,000 ha of EIS treatments of low but increasing SBW populations, second instar larvae (L2) SBW levels across northern New Brunswick were found to be considerably lower than populations in adjacent Québec. Treatments increased from 4500 ha in 2014, to 56,600 ha in 2016, and to 199,000 ha in 2018. SBW populations in blocks treated with Bacillus thuringiensis or tebufenozide insecticide were consistently reduced, and generally did not require treatment in the subsequent year. Areas requiring treatment increased up to 2018, but SBW L2 populations showed over 90% reductions in that year. Although this may be a temporary annual decline in SBW population increases, it is counter to continued increases in Québec. Following 5 years of tests, the EIS appears to be effective in reducing the SBW outbreak.

Our review highlights research during the past century focussed on the population ecology of outbreak-prone insect defoliators in Canadian forests. Based on reports from national and provincial surveys that began in the 1930s, there have been at least 106 insect defoliators reported to outbreak, most of which are native Lepidoptera, Hymenoptera (sawflies), or Coleoptera (in order of frequency from most to least). Studies comparing life-history traits of outbreak versus non-outbreak species to better understand why certain species are more outbreak-prone indicate several traits especially common among outbreak species, including egg clustering and aggregative larval feeding. There have been at least 50 time-series studies examining the spatiotemporal population behaviour of 12 major defoliator species. These studies provide evidence for both regular periodicity and spatial synchrony of outbreaks for most major species. Life-table studies seeking to understand the agents causing populations to fluctuate have been carried out for at least seven outbreak species, with the majority identifying natural enemies (usually parasitoids) as the major driver of outbreak collapse. Our review concludes with several case studies highlighting the impact and historical underpinnings of population studies for major defoliator species and a discussion of potential avenues for future research.

Defoliation by spruce budworm (Choristoneura fumiferana Clem.) on balsam fir (Abies balsamea (L.) Mill.) is more severe in fir than in mixed fir-hardwood stands. Previous studies assumed that defoliation in fir-hardwood stands was reduced in proportion to percent hardwood regardless of outbreak severity. We tested the influence of stand composition on defoliation during the first 5 years of a spruce budworm outbreak near Amqui, Quebec, by sampling 27 fir-hardwood plots selected to represent three percent hardwood basal area classes (0%–25%, 40%–65%, and 75%–95%). Balsam fir defoliation was significantly lower (p < 0.001) as hardwood content increased, but the relationship varied with overall defoliation severity each year. Annual plot defoliation in fir-hardwood plots, estimated using: (1) defoliation in pure fir plots and the assumption that defoliation in fir-hardwood plots was reduced in proportion to percent hardwood; (2) a generalized linear mixed-effects model with defoliation in pure fir plots, percent hardwood, and interaction as fixed-effects; and (3) Random Forests prediction incorporating 11 predictor variables, resulted in r = 0.77, 0.87, and 0.92 versus measured defoliation, respectively. Average defoliation severity in softwood plots and percent hardwood content were the most important variables in Random Forests analysis. Data on average defoliation level in softwood stands, as an indicator of overall outbreak severity, improves prediction of balsam fir defoliation in mixed stands.

We investigated the potential for human-mediated range expansion of an exotic beech leaf-mining weevil, Orchestes fagi (Linnaeus) (Coleoptera: Curculionidae: Curculioninae: Rhamphini) (formerly known as Rhynchaenus fagi) on timber or firewood, which for eight to nine months of the year may harbour adults in diapause. In both relatively low-density and high-density populations, adults were found on the base, middle, and upper boles of the primary host, American beech (Fagus grandifolia Ehrhart; Fagaceae), as well as red maple (Acer rubrum Linnaeus; Sapindaceae) and red spruce (Picea rubens Sargent; Pinaceae) in the vicinity. Comparatively few individuals were found on tree branches, or in the moss, duff, or soil collected beneath beech trees. Overwintering adults appeared to favour parts of trees with relatively high bark roughness. Our study suggests that, between the months of July through May, any woody stems near areas having O. fagi outbreaks are likely to harbour adults. Moreover, as all of the trees studied are common sources of timber or firewood, the harvest and transport of wood from these areas may facilitate outbreak spread; this may explain the multiple, distantly distributed populations of O. fagi that have been reported in eastern Nova Scotia, Canada in recent years.