Recognition of foreign oviposition-marking pheromone in a multi-trophic context

Both phytophagous and parasitic insects deposit oviposition-marking pheromones (OMPs) following oviposition that function to inform conspecifics of a previously utilized host of reduced suitability. The blueberry maggot fly, Rhagoletis mendax Curran (Diptera: Tephritidae), deposits eggs individually into blueberries and then marks the fruit surface with an OMP which reduces acceptance of fruit for oviposition by conspecifics. Diachasma alloeum (Muesebeck) (Hymenoptera: Braconidae) is a parasitic wasp attacking larval R. mendax which also deposits an OMP, signaling conspecifics of a wasp-occupied host. Behavioral studies were conducted testing the hypothesis that the OMP of the parasitic wasp modifies the oviposition behavior of its host fly. In this study, we show that the OMP of D. alloeum is recognized by R. mendax, and female flies will reject wasp-marked fruit for oviposition. Thus, we present a rare demonstration of pheromonal recognition between animals occupying different taxonomic orders and trophic levels. This chemical eavesdropping may enhance the ability of the fly to avoid fruit unsuitable for larval development.     

Antennal responses of the western pine beetle, <Emphasis Type="Italic">Dendroctonus brevicomis</Emphasis> (Coleoptera: Curculionidae), to stem volatiles of its primary host, <Emphasis Type="Italic">Pinus ponderosa</Emphasis>, and nine sympatric nonhost angiosperms and conifers

Summary. Stem volatile extracts from ten trees that are sympatric with the western pine beetle, Dendroctonus brevicomis LeConte (Coleoptera: Curculionidae) were assayed by gas chromatographic-electroantennographic detection analysis (GC-EAD). The extracts were from the primary host, ponderosa pine, Pinus ponderosa Dougl. ex Laws. (Pinaceae); two nonhost angiosperms, California black oak, Quercus kelloggii Newb. (Fagaceae), and quaking aspen, Populus tremuloides Michx. (Salicaceae); and seven nonhost conifers, white fir, Abies concolor (Gord. & Glend.) Lindl. ex Hildebr. (Pinaceae), incense cedar, Calocedrus decurrens (Torr.) Florin (Cupressaceae), Sierra lodgepole pine, P. contorta murrayana Grev. & Balf. (Pinaceae), Jeffrey pine, P. jeffreyi Grev. & Balf. (Pinaceae), sugar pine, P. lambertiana Dougl. (Pinaceae), Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco (Pinaceae), and mountain hemlock, Tsuga mertensiana (Bong.) Carr. (Pinaceae). Sixty-four compounds were identified from the ten trees, 42 of which elicited antennal responses in D. brevicomis, usually in both sexes. In addition, several synthetic compounds, including a number of the antennally-active compounds from the extracted trees and some bark beetle pheromone components, elicited antennal responses in a manner similar to that observed with the extracts. Of the antennally-active compounds known to be present in trees sympatric with D. brevicomis, only geraniol was unique to its host. Four antennally-active compounds were found in the host and in other conifers; five compounds were found only in nonhost conifers; eight compounds were found in either or both of the nonhost angiosperms; eight compounds were found in either or both of the angiosperms and in nonhost conifers, but not in the host; and 19 were found in both the host and in angiosperms and/or nonhost conifers. Several bark beetle pheromone components were found in the stem volatile extracts. Conophthorin was identified from both nonhost angiosperms; exo-brevicomin was identified in A. concolor; verbenone was identified from a number of nonhost conifers; and chalcogran was identified from P. tremuloides. The number of nonhost volatile chemicals that D. brevicomis encounters and is capable of detecting, and the diversity of sources from which they emanate, highlight the complexity of the olfactory environment in which D. brevicomis forages. This provides a basis for further work related to chemically-mediated aspects of foraging in this insect and perhaps other coniferophagous bark beetles, and highlights the need to consider foraging context in the design and implementation of semiochemical-based management tactics for tree protection.     

Wind tunnel attraction of grapevine moth females, Lobesia Botrana, to natural and artificial grape odour

Summary. Host plant volatiles which attract insect herbivores for egg-laying are of principal interest with respect to insect ecology and evolution. Direct applications concern population monitoring and control through behavioural manipulation. Identification of behaviourally active plant secondary metabolites is essential also for plant breeding for insect resistance. Grapevine moth females Lobesia botrana are attracted by upwind flight to green grape berries Vitis vinifera. The headspace of grape berries was collected on air filters. A solvent extract of these filters, released from a sprayer, attracted females in the wind tunnel. The results demonstrate that volatile cues mediate attraction of grapevine moth females to grape berries, and that headspace collections capture the essence of this odour signal. The air filter extracts were examined by gas chromatography coupled with electroantennographic detection, and the compounds eliciting a consistent antennal response in L. botrana females were identified by mass spectrometry. The headspace collection apparatus was calibrated for collection efficiency of the active compounds. Their recovery rate ranged from 35 % for methyl salicylate to 83 % for (E,E)-α-farnesene. A synthetic ten-component blend was then formulated. The blend consisted of compounds eliciting an antennal response, formulated in a blend ratio corrected for differences in collection efficiency. Subsequent wind tunnel tests showed that female attraction to this synthetic ten-component blend was not significantly different from attraction to grape berries, or to headspace collections of the same berries. At a release rate of 35 ng/h of the most abundant compound (E)-β-caryophyllene, 20 % of the test females approached the source of sprayed headspace collections and the ten-component synthetic blend, respectively. In comparison, 100 g of green berries, releasing the main compound (E)-β-caryophyllene at a rate of ca. 4.7 ng/h, attracted 10 % of the females by upwind flight followed by source contact.     

Leaf surface compound fromBrassica oleracea (Cruciferae) induces oviposition byPieris brassicae (Lepidoptera: Pieridae)

Summary Chemicals present on the surface of cabbage (Brassica oleracea L.) leaves were extracted by dipping these leaves for 3 s in dichloromethane followed by a 3 s dip in methanol. When offered in dual choice bioassays using green paper cards as a substrate, the methanol extract stimulated oviposition activity byPieris brassicae L. (Lepidoptera: Pieridae) females. The oviposition stimulant was isolated using medium pressure liquid chromatography, reversed-phase HPLC, ion-pair HPLC and ion exchange chromatography. Using¹H-NMR spectroscopy, the stimulant could be identified as glucobrassicin (3-indolyl-methyl-glucosinolate). When pure glucobrassicin was offered at a dose identical to that in the crude methanol extract, butterflies did not discriminate between these two substrates in a dual choice test. It is argued that a high sensitivity for indole glucosinolates as host recognition factors may confer an adaptive value for these specialist crucifer feeders. The nutritional significance of their precursor tryptophan and the non-volatile nature of the aglycones formed upon enzymic hydrolysis in damaged tissues are proposed as properties of indole glucosinolates that contribute to this possible adaptive advantage.     

GC-EAD responses to semiochemicals by eight beetles in the subcortical community associated with Monterey pine trees in coastal California: similarities and disparities across three trophic levels

Summary.  Antennae of six sympatric bark and ambrosia beetles (Scolytidae), Dendroctonus valens LeConte, Gnathotrichus retusus (LeConte), Hylastes tenuis Eichhoff, Ips mexicanus (Hopkins), Ips plastographus maritimus Lanier, and Pseudohylesinus sericeus (Mannerheim), and two scolytid predators, Enoclerus sphegeus (F.) (Cleridae) and Lascontonus tuberculatus Kraus (Colydiidae), were analyzed by gas chromatographic-electroantennographic detection (GC-EAD) for their responses to synthetic Ips spp. pheromone components, and host and nonhost volatiles. The beetles emerged from cut logs of pitch canker-infected Monterey pine trees, Pinus radiata D. Don. There were significant disparities in EAD response patterns to the hemiterpene and monoterpene alcohol pheromone components that are typically produced by Ips spp. Antennae of I. p. maritimus responded strongly to ( ± )-ipsdienol, ( ± )-ipsenol, amitinol, and lanierone; antennae of I. mexicanus responded strongly to (1S,2S)-(–)-cis-verbenol, with weaker responses to ( ± )-ipsdienol, ( ± )-ipsenol, and amitinol; antennae of H. tenuis responded to (1S, 2R)-(–)-trans-ver-benol, with less pronounced responses to (–)-cis-verbenol and 2-methyl-3-buten-2-ol; and antennae of D. valens, G. retusus, and P. sericeus generally responded to all Ips spp. pheromone components except 2-methyl-3-buten-2-ol (D. valens and G. retusus) and E-myrcenol (G. retusus and P. sericeus). Ips mexicanus responded only to the (–)-enantiomers of ipsenol and ipsdienol, whereas I. p. maritimus responded to (–)-ipsenol, but to both the (+)- and (–)-enantiomers of ipsdienol. The antennae of the two predaceous insects (E. sphegeus and L. tuberculatus) responded to a range of the Ips spp. pheromone components. Host monoterpenes elicited no antennal responses from E. sphegeus, G. retusus, H. tenuis, and I. mexicanus, but several monoterpenes elicited various levels of responses from D. valens and I. p. maritimus antennae. Interestingly, antennae of female D. valens responded to (–), but not (+)-limonene. α- and β-Pinene elicited weak responses from L. tuberculatus antennae. EAD responses to selected nonhost volatiles were almost identical among the six scolytid species, with trans-conophthorin eliciting the strongest response in most cases, followed by three C₆- alcohols and two C₈-alcohols. The antennal responses by most of these species to linalool or geranylacetone were very weak; (E)-2-hexenal, (Z)-3-hexenyl acetate, and benzyl alcohol elicited almost no response. The response pattern of P. sericeus to nonhost volatiles differed slightly from the rest of the scolytids: a strong response to linalool, weaker response to the C₈-alcohols. The two predaceous Coleoptera generally had weak, but detectable, responses to nonhost volatiles, except for a relatively strong response to trans-conophthorin by L. tuberculatus. No notable differences in EAD responses were observed between males and females of the two Ips spp. Our results provide an electrophysiological baseline for future efforts to identify attractive and repellent semiochemicals (aggregation pheromones, host kairomones, or nonhost interruptants) for this guild of scolytids and their key predators that are associated with moribund and pitch canker- infected P. radiata.     

Behavior and ecology of Old World Luperini beetles of the genusAulacophora (Coleoptera: Chrysomelidae)

Summary The attraction of Old World leaf beetles in the genusAulacophora to kairomones and parakairomones which are effective lures for New WorldDiabrotica andAcalymma were investigated. Beetles captured on sticky traps baited with single and multicomponent lures were no different from the control traps for two species ofAulacophora. Yellow colored traps and squash blossoms are attractive toAulacophora beetles which detect sub-microgram quantities of cucurbitacins on silica gel. Leaf feeding behavior and flight activity data are correlated with varietal preference of threeAulacophora species. The common response byDiabrotica andAulacophora to cucurbitacins reinforces the two groups' coevolutionary association with the Cucurbitaceae. The apparent lack of a common response toCucurbita blossom volatiles suggests recent evolutionary pathways are substantially different for these two groups of beetles.     

Sequestration of cucurbitacin analogs by New and Old World chrysomelid leaf beetles in the tribe Luperini

Summary Two South American polyphagous leaf beetles,Diabrotica speciosa andCerotoma arcuata, selectively accumulated the bitter tasting compound 23,24-dihydrocucurbitacin D in their body after ingesting root tissues of cucurbit plants. Similarly, three Asian Cucurbitaceae-feeding specialists in the genusAulacophora were found to sequester the same compound. Cucurbitacin analogs were shown to deter feeding by a bird predator, indicating an allomonal role for these compounds in cucurbitacin-associated chrysomelid leaf beetles both of New and Old Worlds. The strong affinity to cucurbitacins, selective sequestration of the analogs and consequent protection from predators suggested an ecological adaptation mechanism developed in common among these two geographically isolated subtribes in the Luperini.