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.

Summary• 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.

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.

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Bark beetles (Coleoptera: Curculionidae, Scolytinae) have specialized feeding habits, and commonly colonize only one or a few closely related host genera in their geographical ranges. The red turpentine beetle, Dendroctonus valens LeConte, has a broad geographic distribution in North America and exploits volatile cues from a wide variety of pines in selecting hosts. Semiochemicals have been investigated for D. valens in North America and in its introduced range in China, yielding apparent regional differences in response to various host volatiles. Testing volatiles as attractants for D. valens in its native and introduced ranges provides an opportunity to determine whether geographic separation promotes local adaptation to host compounds and to explore potential behavioral divergence in native and introduced regions. Furthermore, understanding the chemical ecology of host selection facilitates development of semiochemicals for monitoring and controlling bark beetles, especially during the process of expansion into new geographic ranges. We investigated the responses of D. valens to various monoterpenes across a wide range of sites across North America and one site in China, and used the resulting information to develop an optimal lure for monitoring populations of D. valens throughout its Holarctic range. Semiochemicals were selected based on previous work with D. valens: (R)-(+)-alpha-pinene, (S)-(-)-alpha-pinene, (S)-(-)-beta-pinene, (S)-(+)-3-carene, a commercially available lure [1:1:1 ratio of (R)-(+)-alpha-pinene:(S)-(-)-beta-pinene:(S)-(+)-3-carene], and a blank control. At the release rates used, (+)-3-carene was the most attractive monoterpene tested throughout the native range in North America and introduced range in China, confirming results from Chinese studies. In addition to reporting a more effective lure for D. valens, we present a straightforward statistical procedure for analysis of insect trap count data yielding cells with zero counts, an outcome that is common but makes the estimation of the variance with a Generalized Linear Model unreliable because of the variability/mean count dependency.

Abstract. 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.

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.

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.