Ability of stress‐related volatiles to attract and induce attacks by Xylosandrus germanus and other ambrosia beetles

• 1 Xylosandrus germanus typically colonizes physiologically‐stressed deciduous hosts but it is increasingly being recognized as a key pest of ornamental nursery stock. We tested the attractiveness of common plant stress‐related volatiles to ambrosia beetles occupying the nursery agroecosystem, as well as their ability to induce attacks on selected trees. Experiments were conducted in Ohio, U.S.A.
• 2 Stress volatile attractiveness was first assessed by positioning traps baited with acetaldehyde, acetone, ethanol and methanol in ornamental nurseries. Cumulative trap counts confirmed that ethanol was the most attractive stress‐related volatile to X. germanus. Methanol‐baited traps were slightly attractive to X. germanus, whereas traps baited with acetaldehyde and acetone were not attractive to any ambrosia beetle.
• 3 A series of tree injection experiments were also conducted to determine the ability of these volatiles to induce attacks by ambrosia beetles under field conditions. Injection of ethanol into Magnolia virginiana induced the largest number of attacks, whereas injection of acetaldehyde induced more attacks than methanol or acetone. Xylosandrus germanus was the most predominant species emerging from M. virginiana injected with each of the stress‐related volatiles. No attacks by wood‐boring beetles were observed on water injected or uninjected control trees.
• 4 Solid‐phase microextraction–gas chromatography–mass spectrometry confirmed the emission of acetaldehyde, acetone, ethanol and methanol after their injection into M. virginiana.
• 5 Xylosandrus germanus has an efficient olfactory‐based mechanism for differentiating among host volatile cues. Injecting select trees with stress‐related volatiles, particularly ethanol, shows promise as a trap tree strategy for X. germanus and other ambrosia beetles.

     

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.     

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

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Freeze stress of deciduous trees induces attacks by opportunistic ambrosia beetles

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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.

     

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

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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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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.     

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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Ambrosia beetle response to ethanol concentration and host tree species

Ethanol emitted by stressed trees is an olfactory cue used by ambrosia beetles (Coleoptera; Curculionidae; Scolytinae) to locate susceptible hosts to colonize. In addition, ethanol enhances the growth of ambrosia beetle fungal symbionts, improving colonization. Whether host selection and colonization are affected also by the amount of ethanol produced by stressed trees and by tree species is unclear. To investigate these mechanisms, we induced attacks by ambrosia beetles in bolts of eight tree species by coring and filling them with either 5% or 90% ethanol solutions in water. For each ethanol concentration, bolts of the eight different tree species were replicated six times in a randomized complete block design. Entry holes were used as a proxy for host selection whereas gallery development stage was used as a proxy for colonization. Ethanol concentration differentially affected host selection of the three ambrosia beetles that were active during this study. Anisandrus dispar Fabricius preferentially attacked bolts with 90% ethanol concentration, Xylosandrus crassiusculus (Motschulsky) preferentially attacked bolts with 5% ethanol concentration, and Xyleborinus saxesenii (Ratzeburg) attacked bolts irrespective of ethanol concentration. Colonization of X. crassiusculus reflected the same pattern observed for entry holes. The effect of host tree species on host selection was most prominent for X. saxesenii, while X. crassiusculus established a higher number of developed galleries in Ostrya carpinifolia Scopoli bolts than on five of the other tested tree species. Our results suggest that ethanol concentration and host tree species may influence ecological niche partitioning among ambrosia beetle 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.     

Short-term development of ambrosia and bark beetle assemblages following a windstorm in French broadleaved temperate forests

Abstract: In most temperate deciduous forests, windstorm is the main source of dead wood. However, the effects of this natural disturbance on ambrosia and bark beetle communities are poorly known. In managed oak‐hornbeam forests storm‐damaged in France in 1999, we sampled ambrosia (and second bark beetles) by ethanol‐baited window‐flight traps in 2001. By comparing uncleared gaps, undisturbed closed‐canopy controls and seedling‐sapling stands, we investigated the short‐term effects of gap formation, gap size and surrounding landscape to provide a snapshot of scolytid response. Contrary to expectations, neither the abundance nor the richness of ambrosia beetle species was significantly higher in gaps than in undisturbed stands. Few responses in abundance at the species level and only a slight difference in assemblage composition were detected between gaps and closed‐canopy controls. Gaps were more dissimilar from seedling‐sapling stands, than from closed‐canopy controls. More scolytid individuals and species were caught in gaps than in seedling‐sapling stands. Mean local and cumulative richness peaked in mid‐size gaps. Only mid‐size gaps differed from closed‐canopy controls in terms of species composition. We identified generalist gap species (Xyleborus saxesenii, X. cryptographus), but also species significantly more abundant in mid‐size gaps (Platypus cylindrus, Xyloterus signatus). The faunistic peculiarity of mid‐size gaps seemed to be partly related to a bias in oak density among gap size classes. Few landscape effects were observed. Only the scolytids on the whole and X. dispar were slightly favoured by an increasing density in fellings at the 78 ha scale. We did not find any correlation between scolytid abundance and the surrounding closed‐forest percentage area. We confirmed that temperate, deciduous, managed stands did not come under threat by ambrosia and bark beetle pests after the 1999 windstorm. Nonetheless, our data stressed the current expansion in Western Europe of two invasive species, X. peregrinus and especially X. germanus, now the predominant scolytid in the three oak forests studied.     

Attraction of ambrosia and bark beetles to coast live oaks infected by Phytophthora ramorum

1 Sudden oak death is caused by the apparently introduced oomycete, Phytophthora ramorum. We investigated the role of bark and ambrosia beetles in disease progression in coast live oaks Quercus agrifolia. 2 In two Marin County, California sites, 80 trees were inoculated in July 2002 with P. ramorum and 40 were wounded without inoculation. Half of the trees in each group were sprayed with the insecticide permethrin [cyclopropanecarboxylic acid, 3‐(2,2‐dichloroethenyl)‐2,2‐dimethyl‐(3‐phenoxyphenyl) methyl ester] to prevent ambrosia and bark beetle attacks, and then were sprayed twice per year thereafter. After each treatment, sticky traps were placed on only the permethrin‐treated trees. Beetles were collected periodically in 2003. 3 Inoculated trees accounted for 95% of all beetles trapped. The ambrosia beetles Monarthrum scutellare and Xyleborinus saxeseni and the western oak bark beetle Pseudopityophthorus pubipennis were the most abundant of the seven species trapped. 4 Permethrin treatment delayed initiation of beetle attacks and significantly reduced the mean number of attacks per tree. Beetles did not attack any wounded or noncankered inoculated trees. 5 Trees with larger cankers trapped more beetles early in the disease. Once permethrin lost effectiveness, the number of beetle entrance tunnels was a more reliable predictor of subsequent trap catch than was canker size. 6 Beetles were initially attracted to P. ramorum cankers in response to kairomones generated in the host‐pathogen interaction. After beetles attacked the permethrin‐treated trees, aggregation pheromones most probably were the principal factor in beetle colonization behaviour.     

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.     

Colonization of disturbed Scots pine trees by bark‐ and wood‐boring beetles

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Real‐time PCR identification of the ambrosia beetles,Trypodendron domesticum (L.) and Trypodendron lineatum (Olivier) (Coleoptera: Scolytidae)

The European hardwood ambrosia beetle (Trypodendron domesticum) and the striped ambrosia beetle (Trypodendron lineatum) are wood‐boring pests that can cause serious damage to lumber, resulting in a need for management of these pests in logging and lumber industries. Natural populations of ambrosia beetles exist throughout the world, but movement of ambrosia beetles into new habitats, particularly via international trade, can result in the establishment of invasive species that have the potential to spread into new territory. Efforts to monitor ambrosia beetle populations are time‐consuming and could be greatly enhanced by the use of molecular methods, which would provide accurate and rapid identification of potentially problematic species. Here, we present new real‐time PCR assays for the detection and identification of T. domesticum and T. lineatum. The methods described herein can be used with a variety of sampling strategies to enable timely and well‐informed decision‐making in efforts to control these ambrosia beetles.     

Verbenone reduces landing of the redbay ambrosia beetle,vector of the laurel wilt pathogen,on live standing redbay trees

1. Laurel wilt is a disease that has caused extensive mortality of redbay Persea borbonia in the southeastern U.S.A. The redbay ambrosia beetle Xyleborus glabratus is the vector of the causal agent of laurel wilt, the fungus Raffaelea lauricola.
2. We tested two potential repellents to the redbay ambrosia beetle, verbenone and methyl salicylate (MeSA) in an 8‐month large‐scale experiment conducted in three locations in Florida. In each location, redbay trees were treated with a single or double application of SPLAT (Specialized Pheromone and Lure Application Technology; ISCA Technologies, Riverside, California) verbenone, as well as SPLAT with a 1:2 mix of MeSA and verbenone.
3. The MeSA + verbenone mixes did not reduce beetle captures compared with the control treatment, whereas SPLAT verbenone alone significantly reduced the number of beetles captured on sticky traps placed on redbay trees in the three locations. The reduction of beetle capture was similar regardless of one or two treatments of SPLAT verbenone. The reduction of tree death with the SPLAT verbenone treatment was not statistically significant.
4. The results of the present study suggest that trunk application of verbenone can reduce landing rates of the redbay ambrosia beetle on live redbay trees and shows promise for use in an integrated pest management strategy against laurel wilt.

     

Spatio‐temporal dynamics of an Ips acuminatus outbreak and implications for management

• 1 Understanding spatio‐temporal processes of bark beetle infestations is crucial for predicting beetle behaviour and aiding management decisions aiming to prevent or mitigate tree mortality. We recorded the spatial and temporal distribution of killed trees during the 5‐year period of an Ips acuminatus outbreak.
• 2 Killed trees were always grouped in well‐defined patches (infestation spots). In years of high population density, infestation spots were large and aggregated, whereas, in years of low density, infestation spots were small and weakly aggregated or randomly distributed within the study area.
• 3 Most trees were killed in the spring by beetles that had hibernated but, in some years, trees were also killed in the summer by new‐generation beetles originating from spring attacks. Spring‐killed trees always formed new infestation spots at new locations (i.e. spot proliferation). By contrast, summer‐killed trees always occurred at the edge of active spots established in the spring, thus resulting in spot growth.
• 4 With regard to management strategies, the results obtained in the present study suggest that areas located in close proximity to infestations of the previous year should be prioritized for risk assessment. Because large spots account for most of the observed tree mortality, the cut‐and‐remove method should be focused on these spots as soon as crown discoloration appears in the summer. If applied timely, this strategy will remove the new‐generation beetles originating from the spring attacks before they emerge and also reduce the risk of spot growth.

     

Comparative efficacy of plant-derived essential oils for managing ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) and their corresponding mass spectral characterization

Ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) pose a significant challenge to producers of ornamental nursery stock. Conventional insecticides are commonly used for management purposes, but plant-derived essential oils also may discourage ambrosia beetles from initiating attacks. To identify promising commercially available products, field-based efficacy trials were conducted in Ohio in 2009 and 2010 with the following products: Armorex (Soil Technologies), Cinnacure (Proguard, Inc.), EcoTrol (EcoSMART Technologies, Inc.), and Veggie Pharm (Pharm Solutions, Inc.). Potted Magnolia virginiana L. were first injected with 75 ml of 5% ethanol to ensure ambrosia beetle pressure on experimental trees. Mixtures of each product (10% in water) and a water control were applied until runoff and attacks occurring under field conditions were quantified at 1, 4, 7, and 14 d after treatment (DAT). Ambrosia beetle attacks generally increased over time but at differing rates depending on the particular treatment. In 2009, Armorex and Veggie Pharm were associated with the lowest cumulative attacks 14 DAT. In 2010, Armorex and Cinnacure were associated with the fewest attacks 14 DAT. Solid phase microextraction-gas chromatography-mass spectrometry was used to characterize the volatile compounds associated with each product. Allyl isothiocyanate, a compound with known repellent and insecticidal properties, was unique and predominant in Armorex. These experiments identified commercially available botanicals containing plant essential oils with activity against ambrosia beetles, along with demonstrating the usefulness of ethanol-injection to ensure ambrosia beetle pressure under field conditions. Characterizing the constituents of efficacious botanically based products could also lead to the development of improved botanical insecticides.     

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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