Calling Behavior of the Cerambycid Beetle <Emphasis Type="Italic">Neoclytus acuminatus acuminatus</Emphasis> (F.)

Males of the cerambycid beetle Neoclytus acuminatus acuminatus (F.) assume a body posture, never displayed by females, that appears to be associated with release of an aggregation pheromone: they periodically stop walking and fully extend their front legs, elevating their head and thorax above the substrate. In this article, we demonstrate that this body posture, the “pushup stance,” coincides with release of pheromone and that it serves to elevate pheromone glands above the substrate. We also use a pheromone proxy system (sublimation of naphthalene) to demonstrate that the pushup stance increases rates of pheromone dissemination. The pushup stance provides a convenient indictor for studying the role of pheromones in reproductive behavior and facilitating collection of pheromone in the laboratory.     

Enantiomers of fuscumol acetate comprise the aggregation‐sex pheromone of the South American cerambycid beetle Psapharochrus maculatissimus,and likely pheromones of the cerambycids Eupromerella plaumanni and Hylettus seniculus

There is increasing evidence that pheromone chemistry within the large coleopteran family Cerambycidae is often highly conserved, with numerous related species sharing the same pheromone components. As a result, traps containing these components can attract multiple cerambycid species simultaneously. In the present study, we exploited this concept in the identification of the male‐produced aggregation‐sex pheromone of the South American species Psapharochrus maculatissimus (Bates) (Coleoptera: Cerambycidae, subfamily Lamiinae, tribe Acanthoderini). Initially, live adults of both sexes were caught using a trap baited with a lure containing a blend of known cerambycid pheromone components. Headspace volatiles were collected from live beetles and analyzed by coupled gas chromatography‐mass spectrometry. Males of P. maculatissimus sex‐specifically produced a 1:38 blend of (R)‐fuscumol acetate ([2R,5E]‐6,10‐dimethylundeca‐5,9‐dien‐2‐yl acetate) and (S)‐fuscumol acetate, which were both components of the pheromone lures to which they had been attracted. In more focused field trials, traps baited with the (S)‐enantiomer, or a blend approximating the natural 1:38 ratio of (R)‐ to (S)‐enantiomers, attracted adults of both sexes in approximately equal numbers. During bioassays, adults of the lamiine species Eupromerella plaumanni (Fuchs) (tribe Acanthoderini) and Hylettus seniculus (Germar) (Acanthocinini) also were attracted, but to different lures, with E. plaumanni being attracted to the racemic mixture of the two enantiomers of fuscumol acetate, whereas H. seniculus was attracted specifically to (R)‐fuscumol acetate. Our results suggest that differences between these sympatric species in the stereochemistry of fuscumol acetate impart species‐specificity to pheromone communication channels, similar to what has been found recently with lamiine species from other continents.     

天牛成虫信息素及嗅觉感受机制研究进展

评述了天牛成虫信息素及嗅觉感受机制的研究进展。天牛雌、雄成虫均可释放性信息素,迄今已对31种天牛的性信息素进行了研究,其中完成组分鉴定的有13种。天牛性信息素包括长距离、短距离和接触性信息素3种类型。天牛性信息素存在变异现象,同种天牛分布在不同地区,其性信息素组分之间存在差异。触角是天牛感受性信息素的主要器官,也是判别成虫通讯方式的形态指标,性信息素发达的种类其触角常具显著的性二型现象。天牛利用寄主信息素（如萜烯类、醇类和酯类）寻找寄主。性信息素和寄主信息素在林间复合使用可提高诱捕率。天牛信息素还包括异种信息素、忌避信息素和产卵干扰素,能够提高天牛寄主定位效率。天牛触角嗅觉感受的神经细胞（RNs）有3类,气味信息经神经细胞群传输至中枢神经系统,神经信号按标记路线或交叉纤维样式输导。天牛气味结合蛋白（OBP）方面的研究尚未见报道。     

Field trials of aggregation pheromones for the stink bugs Chlorochroa uhleri and Chlorochroa sayi (Hemiptera: Pentatomidae)

In field trials, adult Chlorochroa uhleri (St?l) (Heteroptera: Pentatomidae) of both sexes were caught in significant numbers in cylindrical screen traps baited with gray rubber septum lures loaded with the main component of the male-produced pheromone, methyl (E)-6-2,3-dihydrofarnesoate. Addition of the two possible minor components of the pheromone, methyl (E)-5-2,6,10-trimethyl-5,9-undecadienoate and methyl (2E,6E)-farnesoate, did not affect attraction. Combining the pheromone with different concentrations of volatiles mimicking the odors of a known host plant, alfalfa (Medicago sativa L.), had no significant effect on attraction of adult bugs, whereas combining the pheromone with the pheromones of two sympatric stink bug species, Chlorochroa sayi (St?l) and Euschistus conspersus Uhler, decreased trap captures, suggesting interference between the pheromones. Small numbers of Chlorochroa ligata (Say) adults also were attracted, but numbers caught were too low to allow statistical comparisons between lure blends. In field trials with C. sayi, all three of the male-specific pheromone compounds [methyl geranate, methyl citronellate, and methyl (E) -6-2,3-dihydrofarnesoate] were required for optimal attraction. As with C. uhleri, adults of both sexes were attracted to pheromone lures in approximately equal numbers. Because of the decreased volatility (=release rate) of methyl (E)-6-2,3-dihydrofarnesoate in comparison with the other two, lower molecular weight pheromone components, lures needed to be loaded with a disproportionately high amount of methyl (E)-6-2,3-dihydrofarnesoate to obtain the best trap catch. There was no indication that the pheromone components of C. uhleri or E. conspersus interfered with the attractiveness of the C. sayi pheromone in lures containing a blend of all three pheromones.     

天牛性信息素、引诱植物和植物性引诱剂的研究与应用

由于目前所拥有的各种天牛防治方法都难以获得理想的防治效果,研发新型天牛控制技术势在必行。其中利用引诱剂是最具潜力的研究领域之一。对天牛的性信息素、引诱植物和植物性引诱剂3个方面的研究及应用作了综述。天牛雌、雄成虫均可释放性信息素进行两性间的通讯联系。天牛性信息素的分泌部位因天牛种类不同而存在着差异。雄、雌成虫触角上均有感受性信息素的感受器,但报道较多的是雄虫感受器。迄今已鉴定出13种天牛性信息素的化学结构。一般而言,天牛成虫的嗜食树种,对天牛具有较强的引诱作用,可作为诱饵树种使用。寄主植物的某些挥发性物质对天牛成虫也有独特的引诱效果。除此之外,对天牛性信息素、引诱植物和植物性引诱剂研究与应用中存在的困难也进行了讨论,并对它们的应用前景做了展望。     

Response of native and exotic longhorn beetles to common pheromone components provides partial support for the pheromone-free space hypothesis

Longhorn beetles are among the most important groups of invasive forest insects worldwide. In parallel, they represent one of the most well-studied insect groups in terms of chemical ecology. Longhorn beetle aggregation-sex pheromones are commonly used as trap lures for specific and generic surveillance programs at points of entry and may play a key role in determining the success or failure of exotic species establishment. An exotic species might be more likely to establish in a novel habitat if it relies on a pheromone channel that is different to that of native species active at the same time of year and day, allowing for unhindered mate location (i.e., pheromone-free space hypothesis). In this study, we first tested the attractiveness of single pheromone components (i.e., racemic 3-hydroxyhexan-2-one, racemic 3-hydroxyoctan-2-one, and syn-2,3-hexanediol), and their binary and tertiary combinations, to native and exotic longhorn beetle species in Canada and Italy. Second, we exploited trap catches to determine their seasonal flight activity. Third, we used pheromone-baited “timer traps” to determine longhorn beetle daily flight activity. The response to single pheromones and their combinations was mostly species specific but the combination of more than one pheromone component allowed catch of multiple species simultaneously in Italy. The response of the exotic species to pheromone components, coupled with results on seasonal and daily flight activity, provided partial support for the pheromone-free space hypothesis. This study aids in the understanding of longhorn beetle chemical ecology and confirms that pheromones can play a key role in longhorn beetle invasions.     

Adults of the cerambycid beetle Megacyllene caryae use both olfactory and visual information to locate mates

Many species of beetles in the family Cerambycidae use volatile pheromones to facilitate the location of mates. Visual cues may also influence the location of mates, as the adults of many species of cerambycids are often brightly patterned and diurnal. Theory predicts that combining signals or cues of different modalities (e.g., chemical, visual) to transmit information will increase the likelihood of an organism responding to this information, compared to when the signal or cue is presented alone. Here, we test the hypothesis that attraction of adults of the cerambycid beetle Megacyllene caryae (Gahan) (Coleoptera: Cerambycidae, Clytini) to their pheromones will be increased when visual cues are present. Consistent with that hypothesis, the number of beetles caught by traps baited with pheromones was increased 3.4‐fold when a dead adult beetle of that species was attached to the trap, relative to those with just pheromone alone. Capture of M. caryae in our study was also influenced by the position of traps within forest stands, with traps at 100 m within stands catching 2.5× as many beetles as traps at the forest edge. These findings suggest that vision and visual cues play an important role in the location of mates by cerambycid beetles and warrant further research. Also, the inclusion of visual cues on traps may enhance the efficacy of trapping cerambycid beetles, such as the detection of species that are non‐native and potentially invasive, or when monitoring species that are native and of conservation concern.     

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.     

A comparison of trap type and height for capturing cerambycid beetles (Coleoptera)

Wood-boring beetles in the family Cerambycidae (Coleoptera) play important roles in many forest ecosystems. However, increasing numbers of invasive cerambycid species are transported to new countries by global commerce and threaten forest health in the United States and worldwide. Our goal was to identify effective detection tools for a broad array of cerambycid species by testing some known cerambycid attractants and a pheromone in different trap designs placed across a range of habitats. We compared numbers and species richness of cerambycid beetles captured with cross-vane panel traps and 12-unit Lindgren multiple-funnel traps, placed either at ground level (1.5 m high) or canopy level (approximately 3-10 m high), at eight sites classified as either residential, industrial, deciduous forest, or conifer forest. We captured 3,723 beetles representing 72 cerambycid species from 10 June to 15 July 2010. Species richness was highest for the subfamilies Cerambycinae and Lamiinae, which accounted for 33 and 46% of all species captured, respectively. Overall, the cross-vane panel traps captured approximately 1.5 times more beetles than funnel traps. Twenty-one species were captured exclusively in traps at one height, either in the canopy or at ground level. More species were captured in hardwood sites (59 species) where a greater diversity of host material was available than in conifer (34 species), residential (41 species), or industrial (49) sites. Low numbers of beetles (n < 5) were recorded for 28 of the beetle species. The number of species captured per week ranged from 49 species on 21 June to 37 species on 12 July. Cross-vane panel traps installed across a vertical gradient should maximize the number of cerambycid species captured.     

Considering species richness and rarity when selecting optimal survey traps: comparisons of semiochemical baited flight intercept traps for Cerambycidae in eastern North America

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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.

     

Male-produced aggregation pheromone of the cerambycid beetle Neoclytus mucronatus mucronatus

Adult male Neoclytus mucronatus mucronatus (F.) (Coleoptera: Cerambycidae: Cerambycinae) were observed to display behaviors identical to calling behaviors of the congener N. acuminatus acuminatus F., males of which produce an aggregation pheromone. Odors collected from male N. m. mucronatus contained one major male‐specific compound, identified as (R)‐3‐hydroxyhexan‐2‐one. Bioassays determined that both sexes were weakly attracted to racemic 3‐hydroxy‐2‐hexanone. Further field trials determined that enantiomerically enriched (R)‐3‐hydroxyhexan‐2‐one (94% ee) attracted more beetles of both sexes than did the racemic blend. This aggregation pheromone is produced by glands that discharge through pores lying within shallow cuticular depressions in the pronotum of male N. m. mucronatus.     

The mode of evolution of aggregation pheromones in Drosophila species

Aggregation pheromones are used by fruit flies of the genus Drosophila to assemble on breeding substrates, where they feed, mate and oviposit communally. These pheromones consist of species-specific blends of chemicals. Here, using a phylogenetic framework, we examine how differences among species in these pheromone blends have evolved. Theoretical predictions, genetic evidence, and previous empirical analysis of bark beetle species, suggest that aggregation pheromones do not evolve gradually, but via major, saltational shifts in chemical composition. Using pheromone data for 28 species of Drosophila we show that, unlike with bark beetles, the distribution of chemical components among species is highly congruent with their phylogeny, with closely related species being more similar in their pheromone blends than are distantly related species. This pattern is also strong within the melanogaster species group, but less so within the virilis species group. Our analysis strongly suggests that the aggregation pheromones of Drosophila exhibit a gradual, not saltational, mode of evolution. We propose that these findings reflect the function of the pheromones in the ecology of Drosophila, which does not hinge on species specificity of aggregation pheromones as signals.     

Effect of varying monoterpene concentrations on the response of Ips pini (Coleoptera: Scolytidae) to its aggregation pheromone: implications for pest management and ecology of bark beetles

Abstract 1 Host plant terpenes can influence attraction of conifer bark beetles to their aggregation pheromones: both synergistic and inhibitory compounds have been reported. However, we know little about how varying concentrations of individual monoterpenes affect responses. 2 We tested a gradient of ratios of α‐pinene, the predominant monoterpene in host pines in the Great Lakes region of North America, to Ips pini's pheromone, racemic ipsdienol plus lanierone. 3 Ips pini demonstrated a parabolic response, in which low concentrations of α‐pinene had no effect on attraction to its pheromone, intermediate concentrations were synergistic and high concentrations were inhibitory. These results suggest optimal release rates for population monitoring and suppression programmes. 4 Inhibition of bark beetle attraction to pheromones may be an important component of conifer defences. At terpene to pheromone ratios emulating emissions from trees actively responding to a first attack, arrival of flying beetles was low. This may constitute an additional defensive role of terpenes, which are also toxic to bark beetles at high concentrations. 5 Reduced attraction to a low ratio of α‐pinene to pheromone, as occurs when colonization densities become high and the tree's resin is largely depleted, might reflect a mechanism for preventing excessive crowding. 6 Thanasimus dubius, the predominant predator of I. pini, was also attracted to ipsdienol plus lanierone, but its response differed from that of its prey. Attraction increased across all concentrations of α‐pinene. This indicates that separate lures are needed to sample both predators and bark beetles effectively. It also provides an opportunity for maximizing pest removal while reducing adverse effects on beneficial species. This disparity further illustrates the complexity confronting natural enemies that track chemical signals to locate herbivores.     

Synthesis and Field Evaluation of Methyl-branched Ketones,Sex Pheromone Components Produced by Lithosiinae Female Moths in the Family of Arctiidae

Female moths of Lyclene dharma dharma (Arctiidae, Lithosiinae) produce a novel sex pheromone composed of three methyl-branched ketones (I–III) in a ratio of 2:1:1. In order to confirm the structure of III (6,14-dimethyl-2-octadecanone), a mixture of its four stereoisomers was synthesized via chain elongation by two Wittig reactions, starting from 1,7-hexanediol. GC-MS data of the synthetic III were satisfactorily coincident with those of the natural component. In addition to the racemic mixtures of I (6-methyl-2-octadecanone) and II (14-methyl-2-octadecanone), previously synthesized, the activity of III was evaluated in the Iriomote Islands, and effective male attraction was observed for the 2:1:1 mixture of I–III. This result indicates that the females do not produce only one stereoisomer for each component or that the response of the males is not disturbed by the other stereoisomers of natural isomers produced by the females. The field test also revealed that the two-component lure of I and II captured as many males as the mixture of I–III, while lures baited with two components in other combinations and with only one component scarcely exhibited any male attraction ability.     

The mode of pheromone evolution: evidence from bark beetles

Sex and aggregation pheromones consist of species-specific blends of chemicals. The way in which different species' blends have evolved has been the subject of some debate. Theoretical predictions suggest that differences between species have arisen not through the accruing of small changes, but through major shifts in chemical composition. Using data on the aggregation pheromones of 34 species of bark beetle from two genera, Dendroctonus and Ips, we investigated how the distributions of the chemical components of their pheromone blends mirror their phylogenetic relationships. We tested whether there were consistent patterns that could be used to help elucidate the mode of pheromone evolution. Although there were obvious differences in pheromone blends between the two genera, the differences between species within each genus followed a less clear phylogenetic pattern. In both genera, closely related species are just as different as more distantly related species. Within Dendroctonus, particularly, most chemical components were distributed randomly across the phylogeny. Indeed, for some chemicals, closely related species may actually be more different than would be expected from a random distribution of chemical components. This argues strongly against the idea of minor shifts in pheromone evolution. Instead, we suggest that, within certain phylogenetic constraints, pheromone evolution in bark beetles is characterized by large saltational shifts, resulting in sibling species being substantially phenotypically (i.e. pheromonally) different from one another, thus agreeing with theoretical predictions.     

Role of ipsdienol, ipsenol, and cis-verbenol in chemical ecology of Ips avulsus, Ips calligraphus, and Ips grandicollis (Coleoptera: Curculionidae: Scolytinae)

Stressed or damaged pine (Pinus sp.) trees in the southeastern United States are often colonized simultaneously by three southern Ips species (Coleoptera: Curculionidae: Scolytinae): small southern pine engraver, Ips avulsus (Eichhoff); sixspined ips, Ips calligraphus (Germar); and eastern fivespined ips, Ips grandicollis (Eichhoff). All three species mediate colonization of host material with volatile pheromones. All of the southern Ips produce cis-verbenol, and either ipsdienol or ipsenol, and electrophysiological studies have demonstrated that all three southern Ips are able to detect all three compounds. This study examined the role of ipsdienol, ipsenol, and cis-verbenol in the chemical ecology of the southern Ips in Georgia and Louisiana. The most attractive blends of pheromones, with the fewest number of components, were ipsdienol plus ipsenol for I. avulsus, cis-verbenol plus ipsdienol for I. calligraphus, and either cis-verbenol plus ipsenol or ipsdienol plus ipsenol for I. grandicollis. Cross-attraction of I. grandicollis to the pheromone blend most attractive to I. avulsus was observed. Although the presence of heterospecific pheromone reduced the catches of all three species (i.e., the tertiary blend captured fewer beetles than the most attractive binary blends) in both states (significantly in two cases), high numbers of all three species were still captured in traps baited with all three compounds. These results suggest that the pheromones cis-verbenol, ipsdienol, and ipsenol can be combined for monitoring all three species of the southern Ips simultaneously.     

Species overlap, speciation and the evolution of aggregation pheromones in bark beetles

Theoretical predictions suggest that species‐specific signals used in the attraction of mates should evolve to reduce the risk of mismating and hybridization. These predictions lead to the hypothesis that the signals of spatially overlapping (i.e. sympatric or syntopic) species should differ more substantially than those of non‐overlapping species. Earlier studies have tested this prediction for auditory and visual signals. Here we test the hypothesis using olfactory signals, specifically the aggregation pheromones of species from two genera of bark beetles, Dendroctonus and Ips. We found no direct evidence from within these genera regarding the fact that the chemical blends that make up these pheromones differ more substantially in species that overlap in their geographical ranges and/or host‐tree use than in allopatric taxa. However, when comparing between genera, the pheromones of overlapping species appear to be more similar than non‐overlapping species. We hypothesize that the species of host tree utilized by the beetles may have some influence on their pheromone blends. Additionally, within genera, species that overlap in host use tend to be more closely related than species that use different hosts. These results may provide indirect evidence for an effect of species overlap on the evolution of bark beetle pheromones.