Walnut twig beetle landing rates differ between host and nonhost hardwood trees under the influence of aggregation pheromone in a northern California riparian forest

1. Host selection behaviour of the walnut twig beetle (WTB) among hardwood trees was investigated in a riparian forest in northern California by monitoring the landing rate of the beetle with sticky traps on branches baited with 3-methyl-2-buten-1-ol, the male-produced aggregation pheromone.
2. The assay was conducted over 7 days (22 May to 29 May 2017) and compared landing rates on branches of six nonhost species paired with northern California black walnut, Juglans hindsii (the host).
3. A total of 2242/1192 WTB were collected on branches of host/nonhost pairs, and more WTB landed on J. hindsii than on nonhosts in 42 of 58 instances. Female landing rate generally exceeded male landing rate, which underscores the influence of the male-produced synthetic pheromone in this system.
4. Landing rates of WTB males, females, and the combined sexes on boxelder, Acer negundo, and valley oak, Quercus lobata, did not differ significantly from the landing rates on J. hindsii, suggesting that these two nonhost riparian hardwoods do not repel WTB (in the context of the aggregation pheromone).
5. Significantly fewer WTB landed on Oregon ash, Fraxinus latifolia, river red gum, Eucalyptus camaldulensis, Fremont cottonwood, Populus fremontii, and red willow, Salix laevigata, than on J. hindsii, which suggests that these four nonhosts may repel one or both sexes of WTB in the context of the aggregation pheromone. Future analysis of the volatiles from these four hardwood species may lead to the discovery of semiochemical repellents for WTB.

     

Additive and synergistic integration of multimodal cues of both hosts and non-hosts during host selection by woodboring insects

Host selection by herbivorous insects is primarily thought to depend on attraction to olfactory cues emitted from the host species. However, the discrimination of these hosts from non-host species may also arise from the adaptive detection and avoidance of non-host cues, including visual characteristics. Many generalist, conifer-colonising beetles, for example, use characteristic volatiles to identify and discriminate against non-host angiosperm trees such as aspens and birches while flying. These trees also differ in bark reflectance characteristics, which could also mediate host/non-host discrimination by interacting with semiochemicals. We tested this hypothesis by evaluating the responses of eight species of polyphagous woodboring beetles to traps which simulated the visual appearance of coniferous hosts (black) and angiosperm non-hosts (white), and which were baited with host or non-host volatiles. As predicted, three species of large woodboring beetle and a woodboring wasp all avoided white, non-host-simulating traps that were baited with attractive kairomones, and preferred black, host-simulating traps. Conversely, three ambrosia beetle species demonstrated weaker visual preferences, possibly because they preferentially colonise fallen hosts that would transmit less accurate visual information. However, the ambrosia beetle Trypodendron lineatum did show a greater preference for host-coloured traps when these released host-associated kairomones in addition to their pheromone, and also avoided white non-host traps, but only when these released non-host volatiles. To our knowledge, this is the first evidence that multiple non-host cues could synergistically mediate the adaptive discrimination of hosts and non-hosts. Our results suggest that successful host location by generalists arises from the complex integration of cues in multiple sensory modes, and that foraging herbivores evaluate both hosts and non-hosts during their search.     

Attraction to monoterpenes and beetle‐produced compounds by syntopic Ips and Dendroctonus bark beetles and their predators

• 1 Bark beetles are significant mortality agents of conifers. Four beetle species, the pine engraver Ips pini, the six‐spined pine engraver Ips calligraphus sub. ponderosae, the southern pine beetle Dendroctonus frontalis, and the western pine beetle Dendroctonus brevicomis, cohabitate pines in Arizona.
• 2 A pheromone trapping study in ponderosa forests of Arizona determined the attraction of beetles to conspecific and heterospecific pheromone components in the presence and absence of host volatiles, and tested whether predators differ in their attraction to combinations of pheromone components and tree monoterpenes.
• 3 All four bark beetle species differed in their responses to heterospecific lures and monoterpenes. Ips calligraphus was the only species that increased in trap catches when heterospecific lures were added. Heterospecific lures did not inhibit the attraction of either Dendroctonus or Ips species. The replacement of myrcene with α‐pinene increased the attraction of Dendroctonus, whereas the addition of α‐pinene had mixed results for Ips. The prominent predators Temnochila chlorodia and Enoclerus lecontei were more attracted to the I. pini lure than the D. brevicomis lure, and the combination of the two lures with α‐pinene was most attractive to both predator species.
• 4 Cross attraction and limited inhibition of bark beetles to heterospecific pheromones suggest that some of these species might use heterospecific compounds to increase successful location and colonization of trees. Predator responses to treatments suggest that tree volatiles are used to locate potential prey and predators are more responsive to Ips than to Dendroctonus pheromone components in Arizona.

     

Primary attraction and kairomonal host discrimination in three species of Dendroctonus (Coleoptera: Scolytidae)

Abstract 1 Synthetic blends of bole and foliage volatiles of four sympatric species of conifers were released from pheromone‐baited multiple‐funnel traps to determine if three species of tree‐killing bark beetles (Coleoptera: Scolytidae): (i) exhibited primary attraction to volatiles of their hosts and (ii) discriminated among volatiles of four sympatric species of host and nonhost conifers. 2 Bole and foliage volatiles from Douglas‐fir, Pseudotsuga menziesii (Mirb.) Franco, increased the attraction of coastal and interior Douglas‐fir beetles, Dendroctonus pseudotsugae Hopkins, to pheromone‐baited traps. Primary attraction to bole volatiles was observed in interior D. pseudotsugae. Beetles were significantly less attracted to the pheromone bait when it was combined with volatiles of lodgepole pine, Pinus contorta var. latifolia Engelm. or interior fir, Abies lasiocarpa × bifolia. 3 The monoterpene myrcene synergized attraction of mountain pine beetles, Dendroctonus ponderosae Hopkins, to their aggregation pheromones, but there was no evidence of primary attraction to host volatiles or discrimination among volatiles from the four conifers. 4 There was significant primary attraction of the spruce beetle, Dendroctonus rufipennis Kirby, to bole and foliage volatiles of interior spruce, Picea engelmannii × glauca, but beetles did not discriminate among volatiles of four sympatric conifers when they were combined with pheromone baits. 5 Our results indicate that host volatiles act as kairomones to aid pioneer Douglas‐fir beetles and spruce beetles in host location by primary attraction, and that their role as synergists to aggregation pheromones is significant. For the mountain pine beetle, we conclude that random landing and close range acceptance or rejection of potential hosts would occur in the absence of aggregation pheromones emanating from a tree under attack.     

Longhorn beetles and predatory clerid beetles attracted to a blend of longhorn beetle pheromone compounds in a Central European oak forest (Coleoptera: Cerambycidae,Cleridae)

1. A blend of longhorn beetle pheromones was tested as a generic attractant in a Central European oak forest. Overall, 20 cerambycid species totalling 1250 specimens were captured using two trap types.
2. More adults of Phymatodes testaceus and Leiopus nebulosus nebulosus were attracted to pheromone-baited traps compared to solvent controls. Significant numbers of four other species were caught by panel traps but not funnel traps. For the cerambycine Pyrrhidium sanguineum, significantly more beetles were caught in treatment traps than controls. For the cerambycine Anaglyptus mysticus, lepturines Cortodera humeralis and Rhagium sycophanta, the numbers of beetles caught in treatment or control traps were similar, indicating no attraction to the lure blend. Adults of a predatory clerid beetle Clerus mutillarius were caught in significantly larger numbers by both baited trap types in comparison to controls, totalling 1514 specimens.
3. Antennae of both sexes of C. mutillarius showed responses to 3-hydroxyhexan-2-one and 2-methylbutan-1-ol, and males also responded to syn-2,3-hexanediol.
4. Strong attraction of C. mutillarius suggests eavesdropping on the pheromones of cerambycids and that such attractant baits, without traps, might be used to manipulate the local population density of predators in a push-pull biological control program.

     

Comparison of naturally and synthetically baited spruce beetle trapping systems in the central Rocky Mountains

We compared naturally baited trapping systems to synthetically baited funnel traps and fallen trap trees for suppressing preoutbreak spruce beetle, Dendroctonus rufipennis Kirby, populations. Lures for the traps were fresh spruce (Picea spp.) bolts or bark sections, augmented by adding female spruce beetles to create secondary attraction. In 2003, we compared a naturally baited system ("bolt trap") with fallen trap trees and with synthetically baited funnel traps. Trap performance was evaluated by comparing total beetle captures and spillover of attacks into nearby host trees. Overall, the trap systems did not significantly differ in spruce beetle captures, although bolt traps caught 6 to 7 times more beetles than funnel traps during the first 4 wk of testing. Funnel traps with synthetic lures had significantly more spillover than either trap trees or bolt traps. The study was repeated in 2004 with modifications including an enhanced blend synthetic lure. Again, trap captures were generally similar among naturally and synthetically baited traps, but naturally baited traps had significantly less spillover. Although relatively labor-intensive, the bolt trap could be used to suppress preoutbreak beetle populations, especially when spillover is undesirable. Our work provides additional avenues for management of spruce beetles and suggests that currently used synthetic lures can be improved.     

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.

     

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.     

Responses of stored-product Anobiidae to pheromone lures and plant-derived volatiles

Experiments were conducted in different locations to investigate responses of adult Stegobium paniceum and Lasioderma serricorne (Col., Anobiidae) to different commercially available or prototype fabrications of their female‐produced sex pheromones. The results showed that the number of S. paniceum captured in traps baited with the commercially available lures was significantly higher than those captured in traps baited with the prototype stegobinone lures. The three commercially available brands of serricornin lures investigated were equally effective in capturing L. serricorne. In a related study, we conducted a 9‐week trapping experiment to determine if responses of L. serricorne to serricornin can be enhanced by the presence of host plant odours. Traps were baited with serricornin alone, serricornin plus dried red chilli (Capsicum frutescens L.) or red chilli alone. The results showed that the number of beetles captured in traps baited with a combination of serricornin and chilli volatiles were significantly higher than in traps baited with pheromone or chilli volatiles alone, indicating that potential exist for improved monitoring or mass trapping of L. serricorne by combining pheromone with plant‐derived volatiles present in Capsicum spp.     

β-Pinene + ethanol attracts more red turpentine beetles than carene+ ethanol,with or without traces of frontalin,at prescribed burn or thinned sites

1. Red turpentine beetle, Dendroctonus valens LeConte (Coleoptera: Curculionidae: Scolytinae), previously responded more strongly to (−)-β-pinene + ethanol than (+)-3-carene + ethanol lures at sites burned the prior year by wildfire in Oregon and northeastern California, whereas at a thinned-unburned Arizona site (+)-3-carene + ethanol was the stronger attractant. This discrepancy was further examined to tease apart whether D. valens attraction varies by region or previous forest disturbance types.
2. Here, (−)-β-pinene + ethanol and (+)-3-carene + ethanol lures were tested in pine stands at two Oregon sites disturbed the previous year by a prescribed burn or thinning only. Both lures were tested also with or without trace amounts of the pheromone frontalin, as its presence enhanced attractions in China but had not been tested in North America.
3. At both sites, regardless of prior forest disturbance, (−)-β-pinene + ethanol lures attracted the most beetles. Lures releasing trace frontalin attracted more beetles than their corresponding lures without it at both sites, except in one case.
4. Overall, previous year disturbances from disparate management treatments had minimal influence on lure attraction to D. valens. For detection, monitoring or management (−)-β-pinene + ethanol + frontalin in trace amounts attracts the most beetles of lures tested to date in Pacific Northwest pine forests.

     

Determination of Agriotes obscurus (Coleoptera: Elateridae) sex pheromone attraction range using target male behavioural responses

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Limonene: attractant kairomone for white pine cone beetles (Coleoptera: Scolytidae) in an eastern white pine seed orchard in western North Carolina

I report on the attraction of the white pine cone beetle, Conophthoru.s coniperda (Schwarz) (Coleoptera: Scolytidae), to traps baited with the host monoterpene limonene in western North Carolina. Both (+)- and (-)-limonene attracted male and female cone beetles to Japanese beetle traps in an eastern white pine, Pinus strobus L., seed orchard near Murphy, NC. Catches of cone beetles were directly proportional to the release rate of (-)-limonene; (+)-limonene was not tested for dose response. Attraction of cone beetles to the pheromone (+/-)-trans-pityol was increased significantly by both enantiomers of limonene. In all experiments, catches of C. coniperda were strongly male biased with no treatment effect on sex ratio. (- )-Limonene had no effect on trap catches of the predator Enoclerus nigripes (Say) to pityol, whereas (+)-limonene interrupted the attraction of E. nigripes to traps baited with pityol. Of six monoterpenes commonly found in white pine cones, only (-)-alpha-pinene elicited attraction of E. nigripes to Japanese beetle traps.     

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.

     

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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Comparative evaluation of pheromone activity for the European spruce bark beetle <Emphasis Type="Italic">Ips typographus</Emphasis> (L.) (Coleoptera,Scolytidae) and methods of their application

Comparative assessment of attractiveness of different pheromone products is presented and the possibility of using pheromone-baited sticky traps for identification of the pest foci and monitoring of its abundance is considered. The test variants showed high selectivity in the territory of the Tsarskoye Selo open-air museum. Pheromone with the addition of a-pinene and limonene was more effective when using barrier traps. During the season, 2636 beetles were captured in standard traps, 2431 beetles, in standard traps with the addition of a-pinene, and 2469 beetles, in standard traps with the same components and limonene. Comparison of the barrier and sticky traps showed the same tendency in the pest population dynamics revealed by the two types of traps (correlation coefficient 0.8862). The use of additional components increased the duration of pheromone activity. Thus, improvement of the component composition of pheromones and the types of traps is required for using them as tools to monitor the European spruce bark beetle abundance and assess the risk of damage to trees.     

Behavioural response of the invasive Halyomorpha halys (Hemiptera: Pentatomidae) to host plant stimuli augmented with semiochemicals in the field

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Mate finding in the forest cockchafer, Melolontha hippocastani, mediated by volatiles from plants and females

The response of the forest cockchafer, Melolontha hippocastani F. (Coleoptera, Scarabaeidae), towards volatiles emitted by different host plants and conspecifics was tested in field experiments during the flight period at dusk. Funnel traps containing artificially damaged leaves from the host plants Carpinus betulus L. and Quercus rubra L., as well as from the non‐host plant Prunus serotina Ehrh. caught significantly more beetles than empty control traps. On the other hand, traps baited with undamaged leaves from Q. rubra did not catch significantly more beetles than empty controls. Leaves from C. betulus damaged by beetle feeding did not attract more beetles than artificially damaged leaves. By use of gas chromatography coupled with electroantennographic detection (GC‐EAD) electrophysiological responses of males and females were shown for 18 typical plant volatiles. A synthetic mixture of selected typical green plant volatiles was also highly attractive in the field. A total of 9982 beetles was caught during the field experiments, among them only 33 females. This suggests that attraction to damaged foliage during flight period at dusk is male‐specific. Field experiments testing the attractiveness of female M. hippocastani towards conspecific males by employing caged beetles and beetle extracts indicated that males of M. hippocastani use a female‐derived sex pheromone for mate location. On wired cages containing either unmated feeding females, or unmated females without access to foliage, or feeding males in combination with extracts from unmated females, significantly more males landed during the flight period than on comparable control cages containing feeding males or male extracts. A possible scenario of mate location in M. hippocastani involving feeding‐induced plant volatiles and a female‐derived sex pheromone is discussed.     

Early detection of Agrilus planipennis: investigations into the attractive range of the sex pheromone (3Z)-lactone

The emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is a highly destructive primary pest of ash (Fraxinus sp., Oleaceae) trees outside of its native range. Ash is an important component of many ecosystems and its loss would be detrimental to both the economy and the environment. The present study aimed to improve our understanding of the effectiveness of green sticky prism traps baited with host kairomone and insect pheromone lures for A. planipennis and to collect data for modelling the range of attraction of the pheromone (3Z)-dodecen-12-olide [(3Z)-lactone]. Traps were deployed over a single flight season in urban locations of Ontario, Canada, with low densities of EAB. Traps were placed in pairs of trees separated by not more than 25 m. All traps contained the host kairomone, (3Z)-hexenol, with the remaining half in each pairing additionally baited with (3Z)-lactone pheromone. Both lure types were highly effective in capturing EAB, with >90% detection rates overall. However, traps baited with the lactone pheromone and host volatile lures doubled trap captures of EAB over distances of at least 25 m from the nearest traps baited with only host volatiles. Although the baseline detection rate of traps containing (3Z)-hexenol alone is not significantly reduced compared with traps containing (3Z)-lactone, the overall trap effectiveness is significantly increased when (3Z)-lactone is present. The implications for the use of (3Z)-lactone at 3 mg per septum dose in an early warning trapping system are discussed. Trap layout methods and risk-based analysis models can now be further refined by including these data about the attractive range of lures and their behaviour in different plot environments.     

The ambrosia beetle Anisandrus maiche (Stark) is repelled by conophthorin and verbenone and attracted to ethanol in a dose-dependent manner

1. Anisandrus maiche (Stark) (Coleoptera: Curculionidae: Scolytinae) is a non-native ambrosia beetle recently detected in northwestern Indiana. There is a critical need for advanced methods to detect and manage this potentially destructive beetle. Identifying semiochemicals that function as attractants or repellents can inform management practices to protect high-value plantings of hardwood trees.
2. We evaluated the extent to which (S)-(−)-verbenone, (E)-(±)-conophthorin and ethanol influence trap capture of A. maiche using two trap types and heights. We also investigated the effect of ethanol release rate on trap capture.
3. Traps baited with ethanol alone captured the most beetles, while traps baited with (S)-(−)-verbenone or (E)-(±)-conophthorin alone captured few A. maiche, and each compound decreased capture when paired with ethanol lures.
4. There was no difference in mean capture of A. maiche between trap types across treatments, and the height of ethanol-baited traps did not influence the capture rate suggesting this species is widely distributed throughout the canopy.
5. Our results suggest that (S)-(−)-verbenone and (E)-(±)-conophthorin are effective repellents for A. maiche and a release rate of at least 3 g/day of ethanol enhances trap to capture.

     

Sex pheromone of Nearctic Agriotes mancus and its similarity to that of three Palearctic Agriotes invasive in North America

1. The wheat wireworm, Agriotes mancus (Coleoptera: Elateridae), is a predominant elaterid pest species in the Nearctic region, with a life history and morphology similar to those of Agriotes obscurus, Agriotes lineatus and Agriotes sputator, three Palearctic pest elaterids invasive in North America. Here, we report the identification and field testing of the sex pheromone of A. mancus.
2. We collected headspace volatiles from female beetles on Porapak Q, and analysed aliquots of Porapak extract by gas chromatography with electroantennographic detection (GC-EAD) and by GC-mass spectrometry. In GC-EAD recordings, two esters—geranyl butanoate and geranyl hexanoate—elicited antennal responses from A. mancus males. In field experiments, trap lures containing both geranyl butanoate and geranyl hexanoate afforded large captures of A. mancus males, which were—on average—approximately 30-fold higher than captures in traps baited with a single ester.
3. Traps baited with geranyl butanoate as a single-component lure captured a significant number of Palearctic A. sputator, indicating the establishment of A. sputator in its invaded Nearctic range.
4. With the A. mancus sex pheromone now known, it can be included in the development of pheromone-based programmes to monitor and manage native and invasive Agriotes pests in North America.