Integrating repellent and attractant semiochemicals into a push–pull strategy for ambrosia beetles (Coleoptera: Curculionidae)

Non‐native ambrosia beetles (Coleoptera: Curculionidae), especially Xylosandrus compactus (Eichhoff), Xylosandrus crassiusculus (Motschulsky) and Xylosandrus germanus (Blandford), are destructive wood‐boring pests of trees in ornamental nurseries and tree fruit orchards. Previous studies have demonstrated the adults are repelled by verbenone and strongly attracted to ethanol. We tested a “push–pull” semiochemical strategy in Ohio, Virginia and Mississippi using verbenone emitters to “push” beetles away from vulnerable trees and ethanol lures to “pull” them into annihilative traps. Container‐grown trees were flood‐stressed to induce ambrosia beetle attacks and then deployed in the presence or absence of verbenone emitters and a perimeter of ethanol‐baited interception traps to achieve the following treatment combinations: (a) untreated control, (b) verbenone only, (c) ethanol only, and (d) verbenone plus ethanol. Verbenone and ethanol did not interact to reduce attacks on the flooded trees, nor did verbenone alone reduce attacks. The ethanol‐baited traps intercepted enough beetles to reduce attacks on trees deployed in Virginia and Mississippi in 2016, but not in 2017, or in Ohio in 2016. Xylosandrus germanus, X. crassiusculus and both Hypothenemus dissimilis Zimmermann and X. crassiusculus were among the predominant species collected in ethanol‐baited traps deployed in Ohio, Virginia and Mississippi, respectively. Xylosandrus germanus and X. crassiusculus were also the predominant species dissected from trees deployed in Ohio and Virginia, respectively. While the ethanol‐baited traps showed promise for helping to protect trees by intercepting ambrosia beetles, the repellent “push” component (i.e., verbenone) and attractant “pull” component (i.e., ethanol) will need to be further optimized in order to implement a “push–pull” semiochemical strategy.     

The capacity of conophthorin to enhance the attraction of two Xylosandrus species (Coleoptera: Curculionidae: Scolytinae) to ethanol and the efficacy of verbenone as a deterrent

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Influence of flood‐stress on ambrosia beetle host‐selection and implications for their management in a changing climate

• 1 Xylosandrus germanus (Blandford) is a key pest of ornamental nursery trees. Ethanol is the most attractive semiochemical known for X. germanus, and its emission from trees represents a primary host‐selection cue. Ethanol production is induced by a variety of abiotic and biotic stressors, which could thereby predispose trees to attack by ethanol‐responsive ambrosia beetles.
• 2 To better understand X. germanus host‐selection behaviour within ornamental nurseries, a series of experiments examined the influence of flood‐stress on the attractiveness and susceptibility of flowering dogwood Cornus florida L. Under field conditions, more X. germanus were attracted to experimentally flood‐stressed dogwoods than neighbouring nonflooded controls in 2009, 2010 and 2011. Flood‐stressed dogwoods were also preferentially attacked in 2009–2011, although no attacks occurred on any of the neighbouring nonflooded trees.
• 3 Solid‐phase microextraction‐gas chromatography‐mass spectrometry detected ethanol in stem tissue from flooded dogwoods but not nonflooded trees. Acetaldehyde, acetic acid and ethanol were also emitted from the outer bark of flooded dogwoods but not nonflooded trees.
• 4 These results demonstrate that X. germanus preferentially lands on and attacks physiologically‐stressed hosts, and further support the role of ethanol in mediating this interaction.
• 5 Attacks by X. germanus have previously been suspected to occur on trees viewed as ‘apparently‐healthy’, although the possibility of such trees being in apparently‐stressed at the time of attack cannot be ruled out given the results obtained in the present study. Minimizing the impact of stressors known to induce the production of ethanol should be the primary foundation of a management plan for X. germanus and other ethanol‐responsive ambrosia beetles.

     

Fungal pathogen and ethanol affect host selection and colonization success in ambrosia beetles

1. Ambrosia beetles exhibit broad host ranges but a narrow preference based on the condition of the host. Tissues infected by pathogens or containing ethanol can facilitate attacks by ambrosia beetles, although it still remains unclear how these factors interact.
2. The present study aimed to examine how (i) chestnut logs infected with the fungal pathogen Cryphonectria parasitica and treated with ethanol (i.e. baited with ethanol lure, soaked in ethanol or untreated) and (ii) hornbeam logs soaked in different ethanol concentrations (3–12.5%) affect host selection and colonization success of ambrosia beetles.
3. Ethanol‐soaked logs were more attractive to Anisandrus dispar than ethanol‐baited logs or untreated logs, although this difference was more evident in uninfected than infected logs. Increasing ethanol concentration in host tissues was differentially attractive to Xyleborinus saxesenii and Xylosandrus germanus. A nonlinear relationship was also documented between ethanol concentration and emergence of X. germanus adults.
4. Overall, the results obtained suggest that the presence of C. parasitica in chestnut logs can affect host selection in ambrosia beetles. In addition, the ethanol concentration in tree tissues affects host selection and colonization success, although the effect varies depending on the beetle species. This contrasting response could be a niche‐partitioning mechanism based on ethanol within host tissues.

     

Freeze stress of deciduous trees induces attacks by opportunistic ambrosia beetles

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Ambrosia beetle (Coleoptera: Curculionidae) responses to volatile emissions associated with ethanol-injected Magnolia virginiana

Xylosandrus germanus (Blandford) and other species of ambrosia beetles are key pests of ornamental nursery trees. A variety of laboratory- and field-based experiments were conducted in pursuit of improved monitoring strategies and to develop a trap tree strategy for ambrosia beetles. Traps baited with bolts prepared from Magnolia virginiana L. injected with ethanol caught five times more X. germanus than ethanol-baited traps. Basal stem injections of ethanol into M. virginiana induced more ambrosia beetle attacks than irrigating or baiting with ethanol, and no attacks occurred on water-injected trees. A positive correlation was also detected between concentration of injected ethanol and cumulative attacks. Solid phase microextraction-gas chromatography-mass spectrometry characterized bark emissions from ethanol- and water-injected M. virginiana at 1, 2, 10, and 16 d after treatment. Ethanol emission from injected trees steadily declined from 1 to 16 d after treatment, but was not emitted from water-injected trees. A variety of monoterpenes were also emitted in trace amounts from the ethanol- and water-injected trees. Antennal responses of X. germanus via gas chromatography-electroantennographic detection to volatiles from ethanol-injected M. virginiana occurred for ethanol, but not the various monoterpenes. X. germanus and other ambrosia beetles were also equally attracted to traps baited with ethanol alone compared with a synthetic mixture of ethanol plus various monoterpenes formulated to mimic ethanol-injected M. virginiana. Injecting concentrated solutions of ethanol into trees may be useful for establishing odor-based trap trees, which could aid with monitoring programs and/or potentially deflect ambrosia beetles away from valuable nursery stock.     

Ethanol‐injection induces attacks by ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) on a variety of tree species

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Conophthorin enhances the electroantennogram and field behavioural response of Xylosandrus germanus (Coleoptera: Curculionidae) to ethanol

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Phenological observations and trapping tactics for the granulate ambrosia beetle Xylosandrus crassiusculus (Coleoptera: Curculionidae,Scolytinae) in New Zealand

1. The granulate ambrosia beetle (GAB) Xylosandrus crassiusculus, was first formally detected in New Zealand in 2019. Since then, GAB has subsequently been found infesting numerous tree species in the Auckland region.
2. Flight intercept traps baited with ethanol lures were deployed from October 2019 to May 2021 at three sites in the Auckland region to ascertain the phenology of GAB in New Zealand. Two distinct peak flight periods were identified in early and late summer, while a smaller and inconsistent third peak was detected in early autumn. Logistical analysis of GAB captures in the ethanol-lured traps and degree-day (DD) accumulation indicated that 90% of flight activity is completed by 800 DD.
3. To assess monitoring tactics, flight intercept traps were baited with three different ethanol lures with varying release rates or ethanol-soaked or non-soaked wood bolts from three species of trees. A lure releasing 2 g ethanol per day was most effective at capturing GAB. Ethanol-soaked bolts were less effective than the lures.
4. Xylosandrus crassiusculus represents a significant risk for shrubs and trees native to New Zealand, as well as commercial horticultural and forestry trees. We recommend using ethanol-lured panel traps for monitoring purposes.

     

Ambrosia beetles (Curculionidae: Scolytinae and Platypodinae) on Fagus crenata Blume: community structure,seasonal population trends and resource utilization patterns

Ambrosia beetles (many Scolytinae and all Platypodinae) are one of the most important insect pests for forestry worldwide, but little is known about the community structure of ambrosia beetles in terms of their vertical variations and resource utilization. We clarified the community structure and seasonal population trends of ambrosia beetles on 11 living and three newly dead Fagus crenata Blume trees using individual tube traps placed up to 10 m high from May to November in 2007 and 2008. We captured seven scolytine species (Ambrosiodmus lewisi (Blandford), Euwallacea validus (Eichhoff), Trypodendron proximum (Niisima), Xyleborinus saxeseni (Ratzeburg), Xyleborus atratus Eichhoff, Xylosandrus brevis (Eichhoff) and Xylosandrus germanus (Blandford)) and three platypodine species (Crossotarsus niponicus Blandford, Platypus hamatus Blandford and Platypus severini Blandford). The ambrosia beetles were suggested to breed at species‐specific height ranges, with the equal host resource use per individual among the species. Of the three major species, C. niponicus, P. hamatus and P. severini, two (C. niponicus and P. hamatus) had male‐biased sex ratios, which is considered a reproductive strategy to increase maternal fitness. Morphological characteristics of the mandibles may play an important role in the difference of sex roles on reproduction in the three major species.     

Species dependent influence of (-)-alpha-pinene on attraction of ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) to ethanol-baited traps in nursery agroecosystems

Field-based trapping experiments were conducted in Ohio in 2003, 2004, and 2008 to determine the influence of (-)-alpha-pinene on the attraction of exotic and native ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) to ethanol-baited traps. In 2003 and 2004, we determined the effect of adding an (-)-alpha-pinene ultrahigh release lure (UHR; 2 g/d at 20 degrees C) to traps baited with an ethanol UHR lure (0.39 g/d). FewerAnisandrus (Xyleborus) sayi (Hopkins) and Xyleborinus saxeseni (Ratzeburg) were collected in 2003 and 2004 from traps baited with ethanol UHR plus (-)-alpha-pinene UHR compared with ethanol UHR. (-)-alpha-Pinene also reduced the attraction of Xyloterinus politus (Say) to ethanol-baited traps in 2004. Total captures of Xylosandrus germanus (Blandford) in 2003 were higher in traps baited with ethanol UHR plus (-)-alpha-pinene UHR than in traps with ethanol UHR alone but not in 2004. In 2008, captures were compared among traps baited with eight combinations of ethanol and (-)-a-pinene at both UHR and low release (LR) rates. Release rates for ethanol LR and (-)-alpha-pinene LR were 0.027 and 0.0015 g/d, respectively. (-)-alpha-Pinene UHR and (-)-alpha-pinene LR reduced the attractiveness of ethanol UHR to A. sayi and X. saxeseni. Ethanol UHR was also more attractive than ethanol LR to A. sayi and X. germanus. These findings demonstrate traps baited with ethanol alone are more effective than ethanol plus (-)-alpha-pinene for monitoring ambrosia beetle flight activity in ornamental nurseries. Ethanol release rate is also an important consideration for monitoring purposes.     

Attraction of ambrosia beetles to ethanol baited traps in a Slovakian oak forest

The attractiveness of ultra high release ethanol lures to ambrosia beetles in Slovakian oak forests was tested from 2010 to 2012. A total of 24,705 specimens were captured during this three year period with Xyleborinus saxesenii (Ratzeburg, 1837) representing 49.28% (12,174 specimens) of the total. Other dominant species captured in the traps were Anisandrus dispar (F., 1792) (27.84%), Xyleborus monographus (F., 1792) (9.72%) and Trypodendron signatum (F., 1792) (6.04%). During this experiment, Xylosandrus germanus (Blandford, 1894) was detected for the first time in Slovakia with an increase in capture each year (19, 40 and 77 specimens, respectively). Flight period for ambrosia beetles in Slovakia occurs from the beginning of April through the end of September. This is the first time that ethanol baited traps were deployed in Slovakian oak forests and the lures were an effective tool for monitoring native and non-native ambrosia beetles.     

Suitability of California bay laurel and other species as hosts for the non‐native redbay ambrosia beetle and granulate ambrosia beetle

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Ovarian development of Agrilus planipennis: effects of age and mating status and influence on attraction to host volatiles

Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is a major pest of ash trees, Fraxinus spp. (Oleaceae), in North America. This study investigated the timing of reproductive development in female beetles and the influence of female reproductive maturity on attraction to host volatiles. Based on dissections of females of increasing age, females with access to males for mating, and thus presumed mated, developed mature eggs only after 18–24 days. In contrast, female beetles reared without access to males, and thus unmated, did not develop mature eggs at any age. Chemical analysis of cuticular hydrocarbons detected the contact sex pheromone, 9‐methyl‐pentacosane, in cohorts of females which were 8–9 days old and older, supporting previous research that this compound signals sexual maturity to males. Results from field‐trapping bioassays demonstrated that stage of female reproductive maturity influenced their attraction to host volatiles: females caught on traps baited with foliar volatiles contained eggs and ovarioles that were significantly less developed than those on traps baited with bark sesquiterpenes. However, our results revealed that females with immature stages of ovarioles and undeveloped eggs, such as those observed in unmated females, were rarely ever caught on traps baited with either of the host volatile lures. Further research on host compounds attractive to immature females is critical for early detection and possible control of A. planipennis populations during the extended pre‐oviposition period.     

Fungal,host and non-host volatiles modify attraction of the walnut twig beetle,Pityophthorus juglandis,to pheromone lures

1. Thousand Cankers Disease (TCD) of walnut trees is caused by the pathogenic fungus Geosmithia morbida vectored by the walnut twig beetle (WTB) Pityophthorus juglandis. Monitoring efforts for WTB rely on pheromone-baited traps, but lures are likely effective at attracting beetles only over short distances. Fungal-derived kairomones may increase the efficacy of current lures, while additional volatiles may repel beetles from valuable trees.
2. The objective of this study was to determine the extent to which fungal, host and non-host volatiles modify the attraction of WTB to pheromone-baited traps. A trapping study that combined fungal, host-associated and non-host compounds with WTB-pheromone lures was conducted over three years in black walnut plantations experiencing a TCD outbreak in Walla Walla, WA.
3. Traps baited with pheromone and G. morbida volatiles (i.e., isoamyl and isobutyl alcohol) consistently attracted more WTB, while other fungal volatiles inconsistently increased attraction compared to those baited with pheromone lure alone. This is the first field study that demonstrates fungal volatiles can increase the attraction of a bark beetle to its pheromone in a hardwood system.
4. One fungal (benzyl alcohol) and two additional volatiles (limonene, piperitone) repelled WTB from pheromone-baited traps. Although limonene is known to repel WTB, this is the first demonstration that benzyl alcohol and piperitone repel a bark beetle.
5. Fungal volatiles may increase the efficacy of monitoring efforts and may play an important role in management tactics for WTB, especially in detecting the introduction and establishment of nascent populations and protecting trees from colonizing beetles.

     

Test of nonhost angiosperm volatiles and verbenone to protect trap trees for Sirex noctilio (Hymenoptera: Siricidae) from attacks by bark beetles (Coleoptera: Scolytidae) in the northeastern United States

Sirex noctilio F. (Hymenoptera: Siricidae) is an invasive woodwasp, currently established in northeastern North America. In other regions of the world, stressed trap trees are used to monitor populations of S. noctilio and to provide inoculation points for the biological control nematode Deladenus siricidicola Bedding. However, the operational use of trap trees for S. noctilio in North America may be compromised by the large community of native organisms that inhabit stressed and dying pine trees. Common bark beetles such as Ips pini (Say) and Ips grandicollis (Eichhoff) (Coleoptera: Scolytidae) could potentially compete with S. noctilio and associates for resources on trap trees, possibly reducing the efficacy of trap trees as habitats for the woodwasp. In an attempt to develop a technology to mitigate this potential issue, three common semiochemical interruptants--conophthorin, green leaf volatile mix, and verbenone--were tested for effectiveness in reducing arrivals of I. pini and I. grandicollis on trap trees treated with herbicides in northeastern United States. In addition, the effects of these compounds were determined independently with pheromone-baited multiple-funnel traps. None of the interruptants reduced numbers of I. pini or I. grandicollis either arriving on trap trees or caught in pheromone-baited traps. However, verbenone increased catches of I. grandicollis in traps baited with its pheromone, ipsenol. The mix of green leaf volatiles reduced catches of a native ambrosia beetle, Gnathotrichus materiarius (Fitch), whereas verbenone reduced trap catches of an exotic ambrosia beetle, Xylosandrus germanus (Blandford). Catches of X. germanus in traps adjacent to trap trees were enhanced with conophthorin.     

Attraction of the cerambycid beetles Tetropium gabrieli,T. castaneum and T. fuscum to pheromones and host tree volatiles

1. Tetropium gabrieli is native to the European Alps where it breeds in European larch and may be an important pest during drought periods by killing trees. It has spread to larch plantations in several European countries including Sweden.
2. In trapping studies conducted in Sweden, we tested whether T. gabrieli was attracted to E-fuscumol, E,Z-fuscumol, E,Z-fuscumol acetate and the host volatile combination of α-pinene plus ethanol. We also compared its response with those of the native congeners, T. fuscum and T. castaneum.
3. All three Tetropium species were caught in significantly higher numbers in traps baited with E-fuscumol plus host volatiles than in traps baited with host volatiles alone or in unbaited traps.
4. There was no difference in catch of any Tetropium species between traps baited with E-fuscumol plus host volatiles versus traps baited with E,Z-fuscumol plus host volatiles, but the latter treatment caught more T. fuscum and T. castaneum than did E,Z-fuscumol acetate plus host volatiles.
5. The addition of E-fuscumol to host volatile-baited traps significantly reduced mean catch of T. gabrieli, significantly increased mean catch of T. castaneum, and had no significant effect on catch of T. fuscum. Mean catch of all three species was significantly greater in traps baited with E-fuscumol alone than in traps baited with host volatiles alone.
6. All three species were caught from May to August. The median date of captures was about 2 weeks earlier for T. fuscum compared with the other two species.

     

Landscape‐scale genetic differentiation of a mycangial fungus associated with the ambrosia beetle,Xylosandrus germanus (Blandford) (Curculionidae:Scolytinae) in Japan

In this study, we examined the genetic structures of the ambrosia fungus isolated from mycangia of the scolytine beetle, Xylosandrus germanus to understand their co‐evolutionary relationships. We analyzed datasets of three ambrosia fungus loci (18S rDNA, 28S rDNA, and the β‐tubulin gene) and a X. germanus locus dataset (cytochrome c oxidase subunit 1 (COI) mitochondrial DNA). The ambrosia fungi were separated into three cultural morphptypes, and their haplotypes were distinguished by phylogenetic analysis on the basis of the three loci. The COI phylogenetic analysis revealed three distinct genetic lineages (clades A, B, and C) within X. germanus, each of which corresponded to specific ambrosia fungus cultural morphptypes. The fungal symbiont phylogeny was not concordant with that of the beetle. Our results suggest that X. germanus may be unable to exchange its mycangial fungi, but extraordinary horizontal transmission of symbiotic fungi between the beetle's lineages occurred at least once during the evolutionary history of this symbiosis.     

Efficacy of traps, lures, and repellents for Xylosandrus compactus (Coleoptera: Curculionidae) and other ambrosia beetles on Coffea arabica plantations and Acacia koa nurseries in Hawaii

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.     

Volatiles from the symbiotic fungus Raffaelea lauricola are synergistic with Manuka lures for increased capture of the Redbay ambrosia beetle Xyleborus glabratus

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