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Canada has a long and proud history in forest science, especially forest entomology. For over a hundred years, Canadians have been conducting research on forest insects to understand their ecology, measure their ecological and economic impacts, and find ways to prevent damage and implement control, among other areas of endeavour. In many cases this research has been influential globally. This collection of 14 papers involving 38 Canadian scientists is a centennial celebration of this history. Each paper reviews, and in some cases synthesizes, a major area of forest entomology science, including identification of major gaps and opportunities to help provide direction to future endeavours in these areas. David W. Langor, René I. Alfaro
From the early descriptive work, focussed on documenting the forest insect fauna
and the impacts of destructive species, Canadian forest entomology has passed
through several distinct phases, each triggered by new societal demands of
forests and of forest entomologists. We review the various stages that Canadian
forest entomology gone through in the last 100 years. Following the initial
descriptive and cataloguing phase, forest entomology entered a pest control or
forest protection phase, which eventually evolved into the integrated pest
management (IPM) era. Although IPM dominated the forest entomology discourse for
at least two decades, this approach is now considered to be narrow and
pest-centric and is being superseded by a more holistic approach where the
emphasis is on ensuring the health and sustainability of forests at landscape
levels. The new trends point away from the “command and control”
approach of attempting to eradicate pests or reducing pest damage, and towards
working with natural processes in the context of ecosystem management. We
indicate several areas where 21st century forest entomology could make a
contribution towards the sustainable management of Canadian forests.
Canada has a distinguished history of research in forest entomology. The number of peer-reviewed publications emanating from studies in forest entomology in Canada greatly increased following the Second World War. Much of the outstanding historical success in Canadian forest entomological research is attributable to the work of entomologists employed by the Canadian Forest Service, who authored the majority of studies until the mid 1970s and usually published them in The Canadian Entomologist. Since that time the majority of studies have been published by Canadian universities in a broad range of journals. Most early research in forest entomology in Canada was carried out by men, but since that time the proportion of forest entomological research carried out by women has increased significantly. The majority of research in central and eastern Canada focussed on defoliators and their natural enemies and host plants whereas the majority of research in western Canada examined bark beetles (Coleoptera: Curculionidae: Scolytinae) and their natural enemies and host plants. Although publications on defoliators and their natural enemies have occurred continuously throughout the historical development of forest entomology in Canada, the numbers of publications on wood borers (Coleoptera: Buprestidae, Cerambycidae) and bark beetles and their natural enemies have never been higher than they are presently.
The comparative ecology of conifer-feeding budworms in the genus Choristoneura Lederer (Lepidoptera: Tortricidae) in Canada is reviewed with emphasis on publications since 1980. Systematics and life history are updated and historical outbreak patterns and their current interpretation summarised. Recent evidence is analysed in the context of ecological interactions among three trophic levels; host plant, budworm herbivore, and natural enemies. The influence of weather and climate are viewed as modulating factors. The population behaviour of budworms is interpreted as the result of tri-trophic interactions that vary at different scales. The result of these multi-scale interactions is that despite shared phylogenetic constraints and common adaptations, different budworm species display different population behaviour because of specific ecological relationships with their respective hosts and natural enemies.
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.
Almost 100 years have passed since J.M. Swaine, the assistant entomologist in charge of Forest Insect Investigations, wrote, “Canadian bark-beetles: a preliminary classification, with an account of the habits and means of control”. The goal was to “put into the hands of practical foresters information of inestimable practical value… to prevent the continued loss of timber now being destroyed” by “the most insidious enemies of the forest”. In this paper, we celebrate Swaine’s pioneering work by summarising the foundational aspects of his early treatise of 1918: the “general habits” of bark beetles, classifications of their behaviour, causes of population increase, and mitigation tactics. In the founding text, Swaine identified all major Dendroctonus Erichson (Coleoptera: Curculionidae) bark beetles found in Canada, although details on life histories were scarce. We summarise current knowledge of the life histories and population dynamics of the spruce beetle, D. rufipennis Kirby; the Douglas-fir beetle, D. pseudotsugae Hopkins; the eastern larch beetle, D. simplex; and address the current range expansion of mountain pine beetle, D. ponderosae Hopkins. We review how aspects of global change, such as invasive species, have altered the population dynamics of certain bark beetles. Finally, we conclude with lessons from two of the many past contributors to bark beetle ecology in Canada, J.M. Swaine and H.A. Richmond.
A number of insect species in a variety of families and orders damage early regenerating forests. Successful management of pests of regenerating forests requires detailed information on the natural history of the damaging organism, including the factors that increase risk, and a careful assessment of the risk mitigation options. Decision support systems, in the form of stand models capable of incorporating pest management options are required to guide decisions in terms of the expected yields under various pest management scenarios. In this paper we review research to date on the natural history and damage of the most important insect pests of regenerating forests in Canada and propose a framework for risk assessment. Canadian scientists have been active contributors to the research that advanced this field, from the early descriptive studies, to the development of practical tools to assist industry in managing pests of young forests.
The impacts of insect outbreaks on tree mortality, productivity, and stand development in Canada are reviewed, emphasising spruce budworm (Choristoneura fumiferana (Clemens), Lepidoptera: Tortricidae) and mountain pine beetle (Dendroctonus ponderosae Hopkins, Coleoptera: Curculionidae). Reduced growth and survival are a function of insect population and defoliation level. It is feasible to make short-term (annual) predictions of insect population and defoliation based upon sampling, but long-term, multi-year predictions are problematic. Given the historical record, it is expected that outbreaks will occur with relatively predictable frequency and basic host relationships and abiotic constraints will not change dramatically. However, the precision of predictions at fine scales is variable and reduced over time. Relationships between tree growth reduction, survival, and cumulative defoliation or beetle population level are available for major insect species. Understanding insect outbreak effects hinges on mortality, changes in interspecies competition, regeneration, and succession. Altered stand dynamics caused by insects can be interpreted for indicators such as wildlife habitat, old forest, riparian buffer cover, viewscapes, and connectivity. Anthropogenic changes are altering impacts via range expansions, northward shifts, and changes in forest composition. We can better understand effects of insect outbreaks and how best to ameliorate damage through a combination of empirical permanent plot studies, modelling, and manipulative experiments.
Symbiotic relationships between insects and fungi are known to cause tree mortality either through direct damage by larval feeding that can be facilitated by symbiotic fungi, or through insects vectoring pathogens directly to healthy trees. Within their native ranges, the impacts of many insect-fungus symbioses are restricted to weakened and declining trees; however, within the last century tree mortality caused by globally invasive insect–fungus associations has had a devastating impact on trees in both urban and natural forest ecosystems. Unfortunately, Canadian forests have been seriously affected by invasive organisms and an emerging threat is the expansion of a native bark beetle into the boreal forest of Alberta. This paper reviews the symbiotic relationships between selected invasive insects and pathogens that cause tree mortality within the urban and forested landscapes of Canada; it uses these case studies to illustrate potentially damaging new evolutionary trajectories.
Insects use semiochemicals to mediate important behaviours such as mating, oviposition, and foraging for resources. Chemical ecology research aims to identify these message-bearing chemicals and develop synthetic copies of semiochemicals for use in integrated pest management (IPM). There has been, and continues to be, an extensive research effort to understand the chemical ecology of various insects considered to be pests of forests in Canada. Canadian chemical ecology research has had an impact on IPM of forest insect pests in Canada and around the world. Canadian researchers have been involved in the identification of semiochemicals used by forest insects and the development and implementation of semiochemical-based management tactics for forest pest management. Semiochemicals have been incorporated into forest pest management for a variety of insect taxa primarily as tools to monitor and control forest insect pests in Canada. The goals of the current review are to: (1) highlight research conducted on semiochemical-based management of forest pests in Canada; (2) discuss the current and potential uses of semiochemicals in IPM of forest pests in Canada; and (3) evaluate potential areas for increased research and implementation of semiochemicals into the management of forest pests in Canada.
The study of insect pathogens became established as a distinct discipline in the late 1940s. In the ~65 years that followed, forest pest management was the main theatre for the development and practice of insect pathology in Canada. Researchers from the federal government and academic institutions contributed to the growing discipline by acquiring foundational knowledge on taxonomy, mode of action, natural occurrence, and ecological role of key pathogens infecting forest pest insects, covering an array of fungi, Microsporidia, viruses, and bacteria. The ultimate goal was to develop pathogen-based alternatives to synthetic insecticides used in large-scale forest protection programmes throughout eastern Canada. That goal was achieved through the development of baculovirus-based products for control of gypsy moths (Lepidoptera: Erebidae), tussock moths (Lepidoptera: Erebidae), and various sawfly (Hymenoptera) species, which are now in the hands of private industry and poised for growing operational use. The second success was the development of products based on Bacillus thuringiensis Berliner (Bacillaceae), which have almost entirely replaced synthetic insecticides in forest protection. We review those successes and other key Canadian contributions to forest insect pathology within the context of emerging digital, molecular, and other technologies, and show how they have altered today’s face of forest pest management in Canada.
Biological control has been an important tactic in the management of Canadian forests for over a century, but one that has had varied success. Here, we review the history of biological control programmes using vertebrate and invertebrate parasitoids and predators against insects in Canadian forests. Since roughly 1882, 41 insect species have been the target of biological control, with approximately equal numbers of both native and non-native species targeted. A total of 161 species of biological control agents have been released in Canadian forests, spanning most major orders of insects, as well as mites and mammals. Biological control has resulted in the successful suppression of nine pest species, and aided in the control of an additional six species. In this review, we outline the chronological history of major projects across Canadian forests, focussing on those that have had significant influence for the development of biological control. The historical data clearly illustrate a rise and fall in the use of biological control as a tactic for managing forest pests, from its dominance in the 1940s and 1950s to its current low level. The strategic implementation of these biological control programmes, their degree of success, and the challenges faced are discussed, along with the discipline’s shifting relationship to basic science and the environmental viewpoints surrounding its use.
Chemical insecticides have been an important tool in the management of forest insect pests in Canadian forests. Aerial application of insecticides began in the 1920s and expanded greatly after World War II with the widespread adoption of DDT primarily for the suppression of spruce budworm, Choristoneura fumiferana Clemens (Lepidoptera: Tortricidae), and other defoliating insects. Significant progress was made in the development of new chemical insecticides and formulations including fenitrothion and tebufenozide, as well as technology for the application of insecticides against various insect pests. However, widespread opposition to the use of chemical insecticides in forest management has led to significant reductions in the number of insecticides registered for use in Canadian forests. Developments in the past 20 years have focussed on new insecticides, formulations, and technologies that seek to limit the impacts on non-target organisms and subsequent ecosystem effects. These developments have resulted in significant improvements in the management of traditional management targets, such as the spruce budworm (Choristoneura fumiferana (Clemens); Lepidoptera: Tortricidae) but also the management of invasive species, especially wood-boring beetles (Coleoptera: Buprestidae, Cerambycidae).
Outbreaks of insect pests periodically cause large losses of volume in Canada’s forests. Compounded with climate change, outbreaks create significant challenges for managing the sustainable delivery of ecosystem services. Current methods to monitor damage by these pests involve both field and aerial surveys. While relatively cost effective and timely, aerial survey consistency and spatial coverage may be insufficient for detailed monitoring across Canada’s vast forest-land base. Remote sensing can augment these methods and extend monitoring capabilities in time and space by incorporating knowledge of pest-host interactions and of how damage translates into a remote sensing signal for detection and mapping. This review provides a brief introduction to major forest insect pests in Canada (two bark beetles (Coleoptera: Curculionidae) and six defoliators) and the damage they cause, a synthesis of the literature involving aerial survey and remote sensing, and a discussion of how these two approaches could be integrated into future pest monitoring from a Canadian perspective. We offer some lessons learned, outline roles that remote sensing could serve in a management context, and discuss what ongoing and upcoming technological advances may offer to future forest health monitoring.
Canada is host to a number of native and introduced forest insects that negatively affect the goods and services provided by forests. Some insects affect forests on a fairly predictable, ongoing basis, while others have impacts that are intermittent, difficult to predict, and sometimes catastrophic. Economic analysis has provided important insights that have informed our understanding of the impact of forest pests and pest management. At the same time, interactions between forests, forest insects, forest management, and economic welfare are complex, and credible assessments may be time consuming and may not always provide results with the certainty sought by policy-makers. This paper reviews the contribution of economics to forest pest management in Canada and suggests future directions for this important field of research.