Laboratory and field tests with the synthetic sex pheromone of threeMatsucoccus (1) pine bast scales.

Matsunone, (2E,4E)-4.6,10,12-tetramethyl-2,4-tridecadien-7-one, is the primary sex pheromone ofMatsucoccus resinosae, M. thunbergianae, andM. matsumurae. The synthetic compound was tested in the laboratory and in field tests with all three species, and significantly more males responded to synthetic matsuone than to controls. The attraction of synthetic matsuone to males in field tests was not significantly different from that of virgin females at the same matsuone release rate. An analog of matsuone previously shown to be biologically active was also tested in the laboratory withM. resinosae andM. thunbergianae, and field tested withM. resinosae. The active analog required a dose approximately 100 times greater than matsuone to yield maximum laboratory bioassay response, and in field tests, attraction was not significantly different from controls at doses at which matsuone was significantly attractive.

Seasonal variability in response ofIps pini (Coleoptera: Scolytidae) to ipsdienol in New York.

In May,Ips pini in New York did not respond in the field to 50-98.5% (R)-(-)-ipsdienol (synthetic). In September, beetles responded strongly to 50-60% (R)-(-)-ipsdienol (synthetic). In May and June, New York beetles showed marked preference for their own males over Arizona males, which produce an average of 94.1% (R)-(-)-ipsdienol. This suggested that ipsdienol stereochemistry alone does not ensure activity and that an additional compound is necessary for attraction in May. In the second year of field tests, attraction to synthetic ipsdienol and male beetles was tested in the spring, summer, and fall. There was response only to males in the spring and mid-summer and to both males and synthetic ipsdienol in the late summer and fall, causing a significant treatment x sampling period (date) interaction. Laboratory-reared beetles were not significantly more attracted to ipsdienol than to a blank airstream in laboratory assays, while male volatiles were significantly more attractive than ipsdienol and the blank. These data demonstrate that there is one or more unknown semiochemicals necessary for pheromonal response and that the behavioral activity of synthetic ipsdienol varies seasonally.

Lanierone: A new pheromone component fromIps pini (Coleoptera: Scolytidae) in New York.

A new pheromone component, lanierone, (2-hydroxy-4,4,6-trimethyl-2,5-cyclohexadien-1-one) was isolated and identified from a Porapak Q collection of volatiles from maleIps pini from New York through GC fractionation, bioassay, and spectrometry. In both the laboratory and the field, synthetic lanierone, in a 1:100 ratio with synthetic ipsdienol, is as attractive as natural pheromone sources. Synthetic ipsdienol alone is not attractive in the laboratory and only weakly attractive in the field. Varying the ratio of lanierone to ipsdienol in the field from 10(-4)∶1 to 1∶1 in 10-fold increments resulted in an increased number of beetles trapped at the three lower ratios, but also in an increase in the proportion of males trapped. In the field, all combinations of lanierone to ipsdienol attracted proportionately fewer males than did pheromone-producing male beetles. GC and GC-MS analyses of Porapak Q-trapped volatiles revealed that lanierone is produced in an amount equal to about 0.2% of that of ipsdienol and is produced exclusively by males. The small amount of lanierone produced, together with a GC retention time similar to that of ipsdienol on a nonpolar column, probably confounded its detection in earlier studies.

Identification of the sex pheromone of threeMatsucoccus pine bast scales.

Matsucoccus resinosae in the United States,M. matsumurae in China, andM. thunbergianae in Korea use (2E, 4E)-4,6,10,12-tetramethyl-2,4-tridecadien-7-one (1) (matsuone) as the primary component of their sex attractant pheromones. The structure was postulated from mass and NMR spectra and confirmed by synthesis of analogs3, (2E,4E)-4,6,11,12-tetramethyl-2,4-tridecadien-7-one, and4, (2E,4Z)-4,6,11,12-tetramethyl-2,4-tridecadien-7-one. Both analogs were attractive to the males ofM. resinosae in laboratory bioassays and toM. matsumurae in laboratory and field tests, but the 4Z analog (4) was much less so than the 4E analog (3) and had inhibitory effects at high concentrations. Dodecanol, isolated from aeration and solvent extracts of femaleM. resinosae, evoked characteristic wing-raising by pedestrian males, but the role of this response was not determined.

Interspecific activity of semiochemicals among sibling species ofPissodes (Coleoptera: Curculionidae).

Pissodes strobi, P. approximatus, andP. nemorensis are sibling species of pine weevils that can hybridize in the laboratory but are presumed to be reproductively isolated in nature. Males of all three species produce the terpenoids grandisol and grandisal; these compounds serve as an aggregation pheromone forP. approximatus andP. nemorensis when deployed with odors from pine bolts. A series of field experiments examined the possibility of cross-attraction among the three species. Tests in New York and Florida found that parapatrically distributedP. approximatus andP. nemorensis were cross-attractive, but different photoperiodic conditioning was required for pheromone production in males of the two species. Long-day pheromone production (P. approximatus-type.) was inherited in interspecific hybrids. Other tests showed thatP. strobi males, or hybrid males from crosses ofP. strobi withP. approximatus, were not attractive to sympatricP. approximatus. When the response ofP. strobi was assessed to males of eitherP. strobi orP. approximatus confined on white pine leaders (the breeding site ofP. strobi), no evidence of cross-attraction or pheromone activity was found;P. strobi were caught in equal numbers onP. strobi-baited leaders,P. approximatus-baited leaders, and unbaited leaders. Tests of interspecific interactions found that maleP. strobi produce an allelochemical signal that interrupts the response ofP. approximatus to its natural or synthetic aggregation pheromone. This interspecific response is apparently adaptive for members of both species (classified as an allomone-kairomone or synomone) because it may ultimately serve to prevent interspecific matings that would lower the fitness of the parents.

Components of moribund American elm trees as attractants to elm bark beetles,Hylurgopinus rufipes and Scolytus multistriatus.

Hylurgopinus rufipes male and female beetles were attracted to American elms infected with Dutch elm disease, and to American elms killed by injection of cacodylic acid.H. rufipes was also attracted to solvent extracts of elm, or to Porapak Q-trapped volatiles from elm. The major components of attractive fractions of Porapak Q-trapped volatiles were isolated, identified, and tested in field bioassays. Several artificially compounded mixtures of sesquiterpenes were attractive toH. rufipes, although no bait tested was as attractive as diseased tree controls. Laboratory bioassays with H. rufipes were marginally successful. In laboratory bioassays, nine of 14 sequiterpenes identified from active fractions of Porapak extracts elicited significant response from Scolytus multistriatus male and female beetles: δ- and γ-cadinene, α-cubebene, γ-muurolene, and ß-elemene were most active. However, in field tests, none of the sesquiterpenes alone or in combination significantly attracted S. multistriatus, nor did they significantly enhance the attraction of S. multistriatus to female-produced pheromone components (4-methyl-3-heptanol [H] and α-multistriatin [M]). In other field tests, α-cubebene (C) significantly enhanced response of S. multistriatus to H plus M, but foliage, logs, or chips of healthy elm did not enhance trap catch to HMC.

Cross-attraction between two species of Matsucoccus : Extraction, bioassay, and isolation of the sex pheromone.

In laboratory bioassays, males of Matsucoccus resinosae from Pinus resinosa in New York and Matsucoccus n. sp. from Pinus thunbergiana in Korea were strongly attracted to crude hexane extracts of M. resinosae females, and M. resinosae males responded strongly to extracts of Matsucoccus n. sp. females. Males of the two species responded similarly to gas chromatographic fractions and subfractions of a pentane extract of M. resinosae females. Sex pheromones of these two species appear to be the same or very similar.

Pheromone receptor cell specificity in interpopulational hybrids ofIps pini (Coleoptera: Scolytidae).

Electrophysiological studies of pheromone receptor cells keyed to ispdienol were performed in laboratory-raised hybrids of the eastern and western populations of the pine engraver,Ips pini. As previously shown in the parental beetles, the receptor cells keyed to ipsdienol could be classified as two distinct types: one keyed to (+)- and one to (-)-ipsdienol. None of the 20 ipsdienol cells recorded from F1 hybrids were of an intermediate type. Recordings of the summated receptor responses (EAGs) showed no significant difference between parental beetles and hybrids. Similar results were obtained in reciprocal crosses, eastern females with western males and the reverse. Thus, there was no indication that sex-linked alleles determined the specificity of the ipsdienol receptor cell. The ratio between (+) and (-) cells was 14∶6 in the hybrids compared to 1∶12 in the western and 9∶12 in the eastern populations.

Trap trees for elm bark beetles : Augmentation with pheromone baits and chlorpyrifos.

European elm bark beetles,Scolytus multistriatus (Marsh.), were strongly attracted to American elms,Ulmus americana L., baited with theS. multistriatus attractant, multilure, or killed by injection of the arboricide, cacodylic acid; a combination of the two treatments was most attractive. Comparisons of beetle catches on sticky bands affixed to the trees with samples of bark from the same trees showed that the number of beetles landing on cacodylic acid-treated trees was approximately 40 times greater than the number boring into them. Spraying the bark with the insecticide chlorpyrifos had no direct effect on attraction. No live bark beetle brood was found in trees that had been treated with cacodylic acid or chlorpyrifos, but trees that were only baited or left untreated (check) were attacked, killed, and colonized. We suggest that the contribution of the cacodylic acid trap tree technique to Dutch elm disease control will be enhanced by baiting treated trees with multilure and spraying their lower boles with 0.5% chlorpyrifos. This treatment will eliminate diseased and unwanted elms as potential breeding material and kill large numbers of elm bark beetles that might otherwise innoculate healthy elms with the Dutch elm disease fungus.

Aggregation pheromone of the deodar weevll,Pissodes nemorensis (Coleoptera: Curculionidae): Isolation and activity of grandisol and grandisal.

The bark weevilPissodes nemorensis, a pest of pines and exotic cedars in the southeastern United States, utilizes a male-produced aggregation pheromone. The presumed pheromone components, grandisol (cis-2-isopropenyl-1-methylcyclobutaneethanol) and its corresponding aldehyde, grandisal, were isolated from extracts of male volatiles and male hindguts. A field test in northern Florida showed that the combination of grandisol, grandisal, and slash pine (Pinus elliottii) bolts acted synergistically to attract large numbers of male and femaleP. nemorensis. These components deployed in various paired combinations were not as attractive as the tripartite mixture. There was no evidence that flying weevils were attracted to unbaited pine bolts. The aggregation pheromone forP. nemorensis appears to be similar to that of a parapatric sibling species,P. approximatus.

Aggregation pheromone components of two species ofPissodes weevils (Coleoptera: Curculionidae) Isolation, identification, and field activity.

Two related volatile compounds were identified from each of two species ofPissodes bark weevils and implicated as components of their aggregation pheromones. Grandisol (cis-2-isopropenyl-1-methylcyclobutaneethanol), and its corresponding aldehyde, grandisal, were isolated from males of bothP. strobi andP. approximatus and were found in the abdomens and hindguts of the respective species. In field tests synthetic grandisol and grandisal together with odors from cut pine acted synergistically in attracting both sexes ofP. approximatus. This response was similar to that elicited by maleP. approximatus feeding on cut pine. Males and females of natural populations ofP. strobi were more responsive to caged males feeding on leaders of white pine than they were to leaders alone. The combination of grandisol, grandisal, and leaders was less attractive than males on leaders, but more attractive than leaders alone. From isolation of pheromone components at different times of the year, it was determined that males of both species produced grandisol and grandisal only at times when cohort females were reproductively mature.

Effects of multilure components on twig-crotch feeding by European elm bark beetles.

The three components of the European elm bark beetle pheromone dispensed from polyethylene vials attached to the boles of healthy juvenile elms affected the rates of beetles landing and twig feeding on the baited trees. Maximum attraction to the tree occurred when all three pheromone components were presented together in a ratio of 1∶1∶8 for 4-methyl-3-heptanol (H), α-multistriatin (M), and α-cubebene (C). M released either alone or in excess of its natural ratio with H and C induced twig-crotch feeding. H presented alone had no effect on attraction or twig-crotch feeding, but in combination with M it induced landing on and boring into the tree bole. We concluded that the ratio of M and H being released influenced incoming beetles either to land on and colonize the bole or to feed in twig crotches. M in excess of H, known to occur when most females are mated, terminates colonization and deflects incoming beetles to crowns of elms.

Electroantennogram responses ofScolytus multistriatus (Coleoptera: Scolytidae) to its pheromone components and to associated compounds.

Electroantennograms were recorded fromScolytus multistriatus in response to 4-methyl-3-heptanol, the four geometric isomers of multistriatin, and cubeb oil. Charateristic dose-response curves for response amplitude and the time required for the voltage trace to return to 1/2 baseline (recovery rate) were established. Recovery rates were significantly more rapid following stimulation with 4-methyl-3-heptanol or cubeb oil than with the multistriatin isomers. At most intensities, α-multistriatin, the isomer that evokes behavioral response, gave significantly larger EAGs with significantly longer recovery rates than the other isomers. Results of differential adaptation experiments suggested that 4-methyl-3-heptanol interacted with the processes involving multistriatin and cubeb oil activity. However, cross-activity of acceptors for these compounds seems unlikely; single sensillum recordings are needed to ascertain the response spectra for individual receptor neurons.

Response ofScolytus multistriatus (Coleoptera: Scolytidae) to α- and δ-multistriatin in Switzerland.

Laboratory bioassays and field tests demonstrated that a Swiss population ofS. multistriatus responded much more strongly to α- than to δ-miltistriatin in combination with 4-methyl-3-heptanol and α-cubebene. High concentrations of brevicomin appeared to replace α-multistriatin in evoking a response byScolytus species, but this effect can be explained by the fact that the brevicomin was contiminated with small amounts of α-multistriatin. Frontalin, another bicyclic ketal, showed no biological activity. Field tests indicated thatS. pygmaeus aggregates to the same attractant mixture asS. multistriatus. S. scolytus also responded preferentially to this mixture, but the relative amounts of α-multistriatin to 4-methyl-3-heptanol do not appear to be as important as forS. multistriatus.

Sex pheromones: abolition of specificity in hybrid bark beetles.

Specificity of sex pheromones maintains breeding isolation among three closely related species of spruce-infesting Ips. Hybrids produced in the laboratory were intermediate to the parent species in both attractiveness and response. Pheromones and pheromone receptor types in the hybrids are probably mixtures of those of the parent species.

"Sex-ratio" condition: unusual mechanisms in bark beetles.

Although males and females occur in similar frequency in the broods of most scolytid beetles, certain individuals of several species produce offspring highly aberrant in the ratio of sexes-all or mostly female. In some instances this condition reflects gynogenesis (parthenogenesis stimulated by mating); in others, a maternally transmitted factor.