AbstractBark beetles (Coleoptera: Curculionidae, Scolytinae) are a major threat to coniferous forests across much of the northern hemisphere, especially in a warmer and drier climate (Fettig et al., 2013). Control of bark beetle outbreaks to protect forests has been a recurring quest for more than a century, with varying success. In the 1970s and 1980s, considerable efforts were directed toward resolving controversies over the application of persistent pesticides as the principal method to manage outbreaks. Advances in research on pheromones of bark beetles and other behavioral compounds during these decades were incorporated into more ecologically benign approaches to managing stands. What emerged was integrated pest management. “Integrated pest management is a process of synthesis where all aspects of the pest-host system are studied and evaluated to provide the resource manager with an information base for decision-making. These aspects include the ecological and socioeconomic components of the system, its interrelations with other resources, treatment tactics to be used, and their effects on the pest and other components of the ecosystem. Evaluation of the decisions implemented is the end of the process and the beginning of a new one, refining the various components of the system to improve the decision support base for future decisions” (Stark & Waters, 1985, pp. 50–52).

Dendroctonus pseudotsugae is the most important forest insect pest of Douglas-fir Pseudotsuga menziesii in North America. Two subspecies of Douglas-fir beetle are recognized: D. pseudotsugae pseudotsugae, which inhabits southwestern Canada and western United States, and Dendroctonus pseudotsugae barragani, which occurs in northern Mexico. This study aimed to determine the effectiveness of 3-methylcylohex-2-en-1-one (MCH) in reducing aggregation of D. pseudotsugae barragani, and corresponding Douglas-fir mortality by this insect. Two field experiments were conducted: the first consisted of three doses of MCH in bubble caps (plus a control treatment) applied within 16 0.5 ha plots. The second consisted of three doses of MCH disrupt micro-flakes (plus a control treatment) applied within 16 1.0 ha plots. MCH bubble caps at 28.8 and 44.0 g AI/ha and MCH disrupt micro-flakes at 185.3 and 741 g AI/ha significantly reduced the aggregation of D. pseudotsugae barragani. When applying MCH as bubble caps, only the highest dose (44.0 g AI/ha) significantly reduced the number of successfully attacked trees. However, MCH micro-flake doses of 185.3 and 741 g AI/ha reduced the number of successfully attacked trees to zero.

AbstractThis paper is intended as an assessment of the state of knowledge and development of semiochemical methods for control of bark beetles (Coleoptera: Curculionidae) in western North America and as a roadmap for future research and development in this field. It is based on a keynote presentation given at the symposiumManaging Bark and Ambrosia Beetles with Semiochemicals, held in 2018 in Vancouver, British Columbia, Canada, at the Joint Meeting of the Entomological Society of America, the Entomological Society of Canada, and the Entomological Society of British Columbia. We describe currently available active ingredients, formulations, and release devices. Furthermore, we describe the most urgently needed new products and techniques, and constraints to their further development. Finally, we speculate about opportunities afforded by new advances in microbial synthesis of semiochemicals, streamlining of regulatory processing, and aerial application using unmanned aerial vehicles.

We summarize the status of semiochemical-based management of the major bark beetle species in western North America. The conifer forests of this region have a long history of profound impacts by phloem-feeding bark beetles, and species such as the mountain pine beetle (Dendroctonus ponderosae) and the spruce beetle (D. rufipennis) have recently undergone epic outbreaks linked to changing climate. At the same time, great strides are being made in the application of semiochemicals to the integrated pest management of bark beetles. In this review, we synthesize and interpret these recent advances in applied chemical ecology of bark beetles for scientists and land managers.

Woodwasps in Sirex and related genera are well-represented in North American conifer forests, but the chemical ecology of native woodwasps is limited to a few studies demonstrating their attraction to volatile host tree compounds, primarily monoterpene hydrocarbons and monoterpene alcohols. Thus, we systematically investigated woodwasp-host chemical interactions in California’s Sierra Nevada and West Virginia’s Allegheny Mountains. We first tested common conifer monoterpene hydrocarbons and found that (−)-α-pinene, (+)-3-carene, and (−)-β-pinene were the three most attractive compounds. Based on these results and those of earlier studies, we further tested three monoterpene hydrocarbons and four monoterpene alcohols along with ethanol in California: monoterpene hydrocarbons caught 72.3% of all woodwasps. Among monoterpene hydrocarbons, (+)-3-carene was the most attractive followed by (−)-β-pinene and (−)-α-pinene. Among alcohols, ethanol was the most attractive, catching 41.4% of woodwasps trapped. Subsequent tests were done with fewer selected compounds, including ethanol, 3-carene, and ethanol plus (−)-α-pinene in both Sierra Nevada and Allegheny Mountains. In both locations, ethanol plus (−)-α-pinene caught more woodwasps than other treatments. We discussed the implications of these results for understanding the chemical ecology of native woodwasps and invasive Sirex noctilio in North America. In California, 749 woodwasps were caught, representing five species: Sirex areolatus Cresson, Sirex behrensii Cresson, Sirex cyaneus Fabricius, Sirex longicauda Middlekauff, and Urocerus californicus Norton. In West Virginia 411 woodwasps were caught representing four species: Sirex edwardsii Brullé, Tremex columba Linnaeus, Sirex nigricornis F., and Urocerus cressoni Norton.

Entomologists and silviculturists have long recommended management of stand basal area and/or mean tree diameter to mitigate the risk of mountain pine beetle (MPB) (Dendroctonus ponderosae Hopkins) outbreaks while simultaneously reducing wildfire risk. In recent decades, however, wildfire suppression and reduced harvests in western North America have created a forest landscape that is densely stocked and increasingly susceptible to bark beetle infestations, especially as the climate becomes warmer and drier. We examine the various MPB treatment options available to land managers, including insecticides, semiochemicals, sanitation, and silvicultural treatments, and describe their long-term consequences in terms of risk of future bark beetle outbreaks, wildfire, invasion by exotic weeds, loss of hydrologic values, and carbon sequestration. Paradoxically, the treatments that are most enduring and best preserve the ecosystem services of North American forests are ones that result in some thinning of these stands. We, therefore, propose a renewed focus on silvicultural treatments over large spatial scales, particularly in lodgepole pine (Pinus contorta Douglas ex Loudon), and recommend semiochemical treatments, which may not protect all trees, for the protection of high-value trees, especially for high-elevation pines that grow in smaller stands. Prophylactic insecticide applications should be reserved for situations where any tree mortality at all is unacceptable.

To develop safe and effective methods to protect whitebark pines, Pinus albicaulis Engelmann, and limber pines, Pinus flexilis James, from attack by mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae), we compared verbenone and verbenone plus green leaf volatiles (GLVs) for prevention of beetle attack. We used two strategies: area-wide protection where semiochemical-releasing flakes are dispersed over the forest floor, and individual tree tests where flakes are applied to tree trunks. The area-wide bioassays were conducted by applying verbenone- and GLV-releasing flakes without stickers to the forest floor on 0.81-ha plots dominated by whitebark pines in the State of Washington with four replicates. We conducted individual tree bioassays by applying the same formulations with stickers to whitebark and limber pines in Montana and Colorado, respectively. In all three situations, both verbenone-alone and verbenone plus GLVs significantly increased the proportion of trees escaping mass attack by beetles, but the two formulations were not significantly different from one another. Despite a lack of significance at a Bonferroni-adjusted α = 0.05, adding GLVs gave slightly greater absolute levels of tree protection in most cases. Monitoring traps placed in the area-wide treatments in Washington showed similar outcomes for numbers of beetles trapped: both treatments had significantly fewer beetles than controls, and they were not significantly different from one another. At peak flight, however, plots with GLVs combined with verbenone had roughly 40% fewer beetles than plots with verbenone alone. GLVs are considerably cheaper than verbenone, so tests of higher application rates may be warranted to achieve enhanced tree protection at reasonable cost.

The red turpentine beetle (RTB), Dendroctonus valens LeConte (Coleoptera: Curculionidae: Scolytinae), is a secondary pest of pines in its native range in North and Central America. Outbreaks and tree mortality attributed to RTB alone are rare in its native range. RTB was introduced into China in the early 1980s and spread rapidly from Shanxi Province to four adjacent provinces; it has infested over 500,000 ha of pine forest and has caused extensive tree mortality since 1999. We provide a historical background on RTB outbreaks, explanations for its invasive success, management options, and economic impacts of RTB in China. Genetic variation in RTB fungal associates, interactions between RTB and its associated fungi, behavioral differences in Chinese RTB, and other factors favoring RTB outbreaks are considered in an effort to explain the invasiveness of RTB in China. The promise of semiochemicals as a management tool is also discussed.

Changes in symbiont assemblages can affect the success and impact of invasive species, and may provide knowledge regarding the invasion histories of their vectors. Bark beetle symbioses are ideal systems to study changes in symbiont assemblages resulting from invasions. The red turpentine beetle (Dendroctonus valens) is a bark beetle species that recently invaded China from its native range in North America. It is associated with ophiostomatalean fungi in both locations, although the fungi have previously been well-surveyed only in China. We surveyed the ophiostomatalean fungi associated with D. valens in eastern and western North America, and identified the fungal species using multi-gene phylogenies. From the 307 collected isolates (147 in eastern North America and 160 in western North America), we identified 20 species: 11 in eastern North America and 13 in western North America. Four species were shared between eastern North America and western North America, one species (Ophiostoma floccosum) was shared between western North America and China, and three species (Grosmannia koreana, Leptographium procerum, and Ophiostoma abietinum) were shared between eastern North America and China. Ophiostoma floccosum and O. abietinum have worldwide distributions, and were rarely isolated from D. valens. However, G. koreana and L. procerum are primarily limited to Asia and North America respectively. Leptographium procerum, which is thought to be native to North America, represented >45% of the symbionts of D. valens in eastern North America and China, suggesting D. valens may have been introduced to China from eastern North America. These results are surprising, as previous population genetics studies on D. valens based on the cytochrome oxidase I gene have suggested that the insect was introduced into China from western North America.

Mountain pine beetle, Dendroctonus ponderosae Hopkins, is a primary agent of forest disturbance in western North America. Episodic outbreaks occur at the convergence of favorable forest age and size class structure and climate patterns. Recent outbreaks have exceeded the historic range of variability of D. ponderosae-caused tree mortality affecting ecosystem goods and services at broad spatial scales. At the same time, significant advances have occurred in our understanding of the chemical ecology of D. ponderosae and in the development of semiochemical-based tools and tactics to protect trees from mortality attributed to D. ponderosae. We synthesize related information relevant to the ecology and management of D. ponderosae and identify factors limiting the effectiveness and utility of semiochemical treatments, areas of continuing research and fields for which further work is needed.

The red turpentine beetle (RTB), Dendroctonus valens LeConte (Coleoptera: Curculionidae: Scolytinae), is a secondary pest of pines in its native range in North and Central America. Outbreaks and tree mortality attributed to RTB alone are rare in its native range. RTB was introduced into China in the early 1980s and spread rapidly from Shanxi Province to four adjacent provinces; it has infested over 500,000 ha of pine forest and has caused extensive tree mortality since 1999. We provide a historical background on RTB outbreaks, explanations for its invasive success, management options, and economic impacts of RTB in China. Genetic variation in RTB fungal associates, interactions between RTB and its associated fungi, behavioral differences in Chinese RTB, and other factors favoring RTB outbreaks are considered in an effort to explain the invasiveness of RTB in China. The promise of semiochemicals as a management tool is also discussed.

To assess effects of low-intensity fire, we combined two silvicultural prescriptions with prescribed fire in the California Cascade Range. In the first treatment, two 100-ha stands were thinned to reduce density while retaining old-growth structural characteristics, yielding residual stands with high structural diversity (HSD). Two other 100-ha plots were thinned to minimize old growth structure, producing even-aged stands of low structural diversity (LSD), and one 50-ha split-plot from each treatment was burned. In addition, two 50 ha old-growth Research Natural Areas (RNA) were selected as untreated reference plots, one of which was also burned. Fire treatments profoundly altered mite assemblages in the short term, and forest structure modification likely exacerbated that response. Sampling conducted two years following treatment confirmed a continuing decline in oribatid mite abundance. Oribatid species richness and assemblage heterogeneity also declined, and community dominance patterns were disrupted. Oribatid responses to fire were either more intense or began earlier in the LSD treatments, suggesting that removal of old-growth structure exacerbated mite responses to fire. Prostigmatids recovered quickly, but their populations nonetheless diminished significantly in burned split-plots. Mite assemblage responses to prescribed fire were continuing nearly two years later, with no clear evidence of recovery.

In an attempt to improve semiochemical-based treatments for protecting forest stands from bark beetle attack, we compared push-pull versus push-only tactics for protecting lodgepole pine (Pinus contorta Douglas ex Loudon) and whitebark pine (Pinus albicaulis Engelm.) stands from attack by mountain pine beetle (Dendroctonus ponderosae Hopkins) in two studies. The first was conducted on replicated 4.04-ha plots in lodgepole pine stands (California, 2008) and the second on 0.81-ha plots in whitebark pine stands (Washington, 2010). In both studies, D. ponderosae population levels were moderate to severe. The treatments were 1) push-only (D. ponderosae antiaggregant semiochemicals alone); 2) push-pull (D. ponderosae antiaggregants plus perimeter traps placed at regular intervals, baited with four-component D. ponderosae aggregation pheromone); and 3) untreated controls. We installed monitoring traps baited with two-component D. ponderosae lures inside each plot to assess effect of treatments on beetle flight. In California, fewer beetles were collected in push-pull treated plots than in control plots, but push-only did not have a significant effect on trap catch. Both treatments significantly reduced the rate of mass and strip attacks by D. ponderosae, but the difference in attack rates between push-pull and push-only was not significant. In Washington, both push-pull and push-only treatments significantly reduced numbers of beetles caught in traps. Differences between attack rates in treated and control plots in Washington were not significant, but the push-only treatment reduced attack rates by 30% compared with both the control and push-pull treatment. We conclude that, at these spatial scales and beetle densities, push-only may be preferable for mitigating D. ponderosae attack because it is much less expensive, simpler, and adding trap-out does not appear to improve efficacy.

DISRUPT Micro‐Flake Verbenone Bark Beetle Anti‐Aggregant flakes (Hercon Environmental, Inc., Emigsville, Pennsylvania) were applied in two large‐scale tests to assess their efficacy for protecting whitebark pinePinus albicaulisEngelm. from attack by mountain pine beetleDendroctonus ponderosaeHopkins (Coleoptera: Scolytinae) (MPB). At two locations, five plots of equivalent size and stand structure served as untreated controls. All plots had early‐ to mid‐outbreak beetle populations (i.e. 7.1–29.2 attacked trees/ha). Verbenone was applied at 370 g/ha in both studies. Intercept traps baited with MPB aggregation pheromone were placed near the corners of each plot after the treatment in order to monitor beetle flight within the plots. Trap catches were collected at 7‐ to 14‐day intervals, and assessments were made at the end of the season of stand structure, stand composition and MPB attack rate for the current and previous years.Applications of verbenone flakes significantly reduced the numbers of beetles trapped in treated plots compared with controls at both sites by approximately 50% at the first collection date.The applications also significantly reduced the proportion of trees attacked in both Wyoming and Washington using the proportion of trees attacked the previous year as a covariate in the model for analysis of current year attack rates; in both sites, the reduction was ≥ 50%.The flake formulation of verbenone appears to have promise for area‐wide treatment by aerial application when aiming to control the mountain pine beetle in whitebark pine forests.

The black twig borer, Xylosandrus compactus (Eichhoff) (Coleoptera: Curculionidae: Scolytinae), is a pest of coffee and many endemic Hawaiian plants. Traps baited with chemical attractants commonly are used to capture ambrosia beetles for purposes of monitoring, studying population dynamics, predicting outbreaks, and mass trapping to reduce damage. The objectives of this research were to optimize trapping systems for X. compactus and other ambrosia beetles such as Xylosandrus crassiusculus (Motschulsky) and Xyleborinus saxesenii (Ratzeburg) by comparing efficacy of several attractants, repellents, and trap types. The ability of certain chemicals to act as beetle repellents and thus interfere with trap catch was tested for purposes of protecting host plants from attack. Potential attractants and application methods tested were as follows: ethyl alcohol pouch delivery system, ethyl alcohol vial delivery system, α-pinene in Eppendorf tubes, eugenol bubblecaps, ginger oil bubblecaps, manuka oil bubblecaps, phoebe oil bubblecaps, and an unbaited control. Potential repellents tested were limonene and verbenone. Ethyl alcohol vials were as attractive as ethyl alcohol sleeves, and were more effective than traps baited with eugenol and α-pinene. Japanese beetle traps were more effective for black twig borer trapping than Lindgren funnel traps, and were easier to deploy. Verbenone and limonene significantly reduced trap catch of Xylosandrus compactus and X. crassiusculus, suggesting that they may be effective for reducing attraction to host plants. These results show the importance of developing a combination of several monitoring techniques to enhance management procedures for the black twig borer.

Novel genotypes often arise during biological invasions, but their role in invasion success has rarely been elucidated. Here we examined the population genetics and behavior of the fungus, Leptographium procerum, vectored by a highly invasive bark beetle, Dendroctonus valens, to determine whether genetic changes in the fungus contributed to the invasive success of the beetle-fungal complex in China. The fungus was introduced by the beetle from the United States to China, where we identified several novel genotypes using microsatellite markers. These novel genotypes were more pathogenic to Chinese host seedlings than were other genotypes and they also induced the release of higher amounts of 3-carene, the primary host attractant for the beetle vector, from inoculated seedlings. This evidence suggests a possible mechanism, based on the evolution of a novel genotype during the two or three decades since its introduction, for the success of the beetle-fungal complex in its introduced region.

*Recent studies have investigated the relationships between pairs or groups of exotic species to illustrate invasive mechanisms, but most have focused on interactions at a single trophic level. *Here, we conducted pathogenicity tests, analyses of host volatiles and fungal growth tests to elucidate an intricate network of interactions between the host tree, the invasive red turpentine beetle and its fungal associates. *Seedlings inoculated with two strains of Leptographium procerum isolated from Dendroctonus valens in China had significantly longer lesions and higher mortality rates than seedlings inoculated with other fungal isolates. These two strains of L. procerum were significantly more tolerant of 3-carene than all other fungi isolated there, and the infection of Chinese pine (Pinus tabuliformis) seedlings by these two strains enhanced the production and release of 3-carene, the main attractant for D. valens, by the seedlings. *Our results raise the possibility that interactions among the fungal associates of D. valens and their pine hosts in China may confer advantages to these strains of L. procerum and, by extension, to the beetles themselves. These interactions may therefore enhance invasion by the beetle-fungal complex.

Bacterial communities are known to play important roles in insect life histories, yet their consistency or variation across populations is poorly understood. Bacteria associated with the bark beetle Dendroctonus valens Le Conte from eight populations, ranging from Wisconsin to Oregon, were evaluated and compared. We used the culture-independent technique of denaturing gradient gel electrophoresis to visualize bacterial diversity, or individual operational taxonomic units (Os), from individual beetles. One-way analysis of similarities was used to test for differences of bacterial communities between sites. Analysis of community profiles showed that individual beetles on average contained 10 OTUs, with frequency of association from 2 to 100% of beetles. OTU sequences most closely matched beta- and gamma-proteobacteria, and one each matched Bacilli and Actinobacteria. Several OTUs were particularly abundant, most notably an Actinobacterium from 100% and two Proteobacteria from 60% of beetles sampled. Some OTUs were similar to previously described bacteria with known biochemical capabilities and ecological functions, suggesting that some bacterial associates of D. valens may contribute to its ability to exploit a resource low in nutrients and high in defensive compounds. There were significant differences of bacterial communities between sites. The strength of these differences was positively correlated with distance between sites, although additional unexplained factors also contribute to the variation.

We tested a new formulation of verbenone, an antiaggregation pheromone of the mountain pine beetle (Dendroctonus ponderosae Hopkins) (Coleoptera: Curculionidae: Scolytinae), for area-wide protection of lodgepole pine (Pinus contorta Douglas ex Loudon) stands in the western United States. Helicopter applications of verbenone-releasing laminated flakes were made at the rate of 370g of active ingredient/ha to two sites, one in California and one in Idaho, during summer 2005. Each site consisted of five 20.23-ha treated plots and five matching 20.23-ha untreated plots. We assessed D. ponderosae flight into study plots using traps baited with aggregation pheromones and we tallied D. ponderosae attack rates on P. contorta trees in treated and control plots before and after application. There were no significant differences between numbers of D. ponderosae trapped on treated and control plots. However, a significantly smaller proportion of P. contorta trees was attacked in treated plots than in control plots at both sites; the attack rate in untreated stands was roughly three times that of treated stands in both California and Idaho, even at this low application rate. Furthermore, attack rate in 2004 was a significant explanatory variable for the response in 2005 regardless of treatment in both California and Idaho. There was no significant treatment effect at either site on attraction of Temnochila chlorodia (Coleoptera: Trogositidae) Mannerheim, a key predator of D. ponderosae, to the prey aggregation pheromone.