Spatial Displacement of a Lure Component Can Reduce Catches of Two Nontarget Species During Spring Monitoring of Southern Pine Beetle.

Local outbreak risk for the southern pine beetle, Dendroctonus frontalis Zimmermann (Coleoptera: Curculionidae), is forecast with a trapping survey conducted every spring throughout the southeastern United States. Traps baited with pine odors and components of the D. frontalis aggregation pheromone are used to obtain abundance estimates of both this species and its clerid predator Thanasimus dubius (F.) (Coleoptera: Cleridae); these data are entered into a predictive model that estimates outbreak risk. An attractant synergist for D. frontalis, endo-brevicomin, has recently been included in the survey lure, but it can have the unintended effect of attracting nontarget species Hylesinus pruinosus Eichhoff (Coleoptera: Curculionidae: Scolytinae) and Enoclerus nigripes (Say) (Coleoptera: Cleridae) which, due to their sometimes large numbers and general similarity in appearance to the target species, could complicate sorting and counting of trap catches. Analysis of bycatch data from a previously-published, 31-mo trapping study in Mississippi indicated that displacement of the endo-brevicomin releaser 6 m from the trap largely eliminated catches of the nontarget species H. pruinosus and E. nigripes while not reducing catches of the target species D. frontalis and T. dubius. Our analysis demonstrates that interspecific differences in spatial responses to attractive semiochemicals can be used to improve insect trap selectivity. Both nontarget beetle species were captured in highest numbers during late winter/early spring, coinciding with the D. frontalis survey.

Sex Pheromone of the Baldcypress Leafroller (Lepidoptera: Tortricidae).

The baldcypress leafroller, Archips goyerana Kruse (Lepidoptera: Tortricidae), is a specialist on Taxodium distichum (L.) Richard and has caused serious defoliation in swamps of southeastern Louisiana, accelerating decline of baldcypress forests concurrently suffering from nutrient depletion, prolonged flooding, and saltwater intrusion. We investigated the composition of the sex pheromone of this species. Coupled gas chromatography-electroantennographic detection (GC-EAD) analyses indicated that male antennae were sensitive to four compounds [(Z)-11-tetradecenyl acetate (Z11-14:OAc), (E)-11-tetradecenyl acetate (E11-14:OAc), (Z)-9-tetradecenyl acetate (Z9-14:OAc), and (Z)-11-tetradecen-1-ol (Z11-14:OH)] present in female abdominal tip extracts in an approximately 100:1.5:0.6:10 ratio. In trapping trials performed in a cypress-tupelo swamp in southeastern Louisiana, moths were attracted to blends of these four components presented in approximately the female-produced ratios. Elimination of Z11-14:OH had no impact on moth response, whereas elimination of any of the three acetates strongly reduced or eliminated attraction. A blend in which the E11:Z11 ratio of 14:OAc was 5:100 was much less attractive than the same blend with the female produced ratio of 1.5:100. A. goyerana is closely related to the sympatric species Archips argyrospilus (Walker) with which it was previously synonymous. Our data revealed differences between the pheromone composition of A. goyerana and that reported for A. argyrospilus, which could account for the apparent absence of cross-attraction between these species. We conclude that a lure containing a 100:1.5:0.6 ratio of Z11-14:OAc, E11-14:OAc, and Z9-14:OAc has the potential to be used in traps to detect and measure A. goyerana populations and thereby monitor an important biotic factor contributing to the loss of coastal baldcypress forests.

Potential for a minor pine bark beetle pest, Dendroctonus terebrans (Coleoptera: Curculionidae: Scolytinae), to mediate host location by a major pine killer, Dendroctonus frontalis.

The southern pine beetle, Dendroctonus frontalis Zimmermann is an important mortality agent of Pinus in the eastern United States of America where it commonly shares hosts with the black turpentine beetle, Dendroctonus terebrans (Olivier), which infrequently kills trees. Unlike D. frontalis, which must kill its hosts to become established in the bark and reproduce, D. terebrans can occupy living hosts as a parasite. Olfactory mechanisms whereby D. frontalis initially locates hosts have not been demonstrated, whereas D. terebrans responds strongly to host odors. Because D. terebrans produces frontalin, the primary aggregation pheromone component for D. frontalis, and commonly arrives on hosts prior to D. frontalis, it has been hypothesized that D. terebrans pheromone components can mediate D. frontalis location of suitable, living trees. We assessed this possibility with studies of the semiochemical interactions between D. frontalis and D. terebrans. Coupled gas chromatography-electroantennographic detection analyses indicated that D. terebrans produces nine different olfactory stimulants for D. frontalis, nearly all of them known semiochemicals for D. frontalis. A trapping experiment designed to address the potentially confounding influence of lure contamination confirmed that the D. terebrans pheromone component exo-brevicomin enhances attraction of D. frontalis and thus could be an attractive kairomone. In ambulatory bioassays, male D. frontalis were strongly attracted to odors of frass of solitary female and paired D. terebrans, indicating their attraction to the naturally occurring semiochemicals of D. terebrans. Cues from D. terebrans may influence host and mate-finding success of D. frontalis and, thereby, the latter's virulence.

Southern pine beetle, Dendroctonus frontalis, antennal and behavioral responses to nonhost leaf and bark volatiles.

A growing body of evidence suggests that bark beetles detect and avoid release points of volatile compounds associated with nonhost species, and thus such nonhost volatiles may have potential utility in the management of bark beetles. We used a coupled gas chromatograph-electroantennographic detector (GC-EAD) to assay the olfactory sensitivity of the southern pine beetle, Dendroctonus frontalis Zimmermann, to volatiles from leaves and bark of eight species of nonhost angiosperm trees that are common in the range of D. frontalis. Tree species sampled were red maple (Acer rubrum L.), mockernut hickory [Carya alba (L.) Nutt. ex Ell.], sweetgum (Liquidambar styraciflua L.), black tupelo (Nyssa sylvatica Marsh.), black cherry (Prunus serotina Ehrh.), southern red oak (Quercus falcata Michx.), blackjack oak [Quercus marilandica (L.) Muenchh.], and water oak (Quercus nigra L.). Beetle antennae responded to a total of 28 identifiable compounds in these samples. The relative olfactory responsiveness to 14 of these, as well as to nonanoic acid and four additional volatiles reported to be associated with nonhost angiosperms, was assessed in GC-EAD analyses of synthetic dilutions spanning six orders of magnitude. The largest response voltage amplitudes were obtained with trans-conophthorin, nonanoic acid, terpinen-4-ol, phenylethyl alcohol, and eucalyptol, whereas the lowest response thresholds were to nonanoic acid, nonanal, linalool, (E)-2-hexen-1-ol, and phenylethyl alcohol. Funnel traps baited with various combinations of eleven antennally-active angiosperm volatiles along with a standard attractant captured significantly fewer male and female D. frontalis than traps baited with the standard attractant alone. Our data suggest that a diversity of semiochemicals may be involved in host species discrimination by D. frontalis, and several may have utility in their management.

Alternative Formulations of Trap Lures for Operational Detection, Population Monitoring, and Outbreak Forecasting of Southern Pine Beetle in the United States.

The southern pine beetle, Dendroctonus frontalis Zimmermann (Coleoptera: Curculionidae: Scolytinae) is a major destructive pest of Pinus L. In the southeastern United States, numbers of this species and a major predator, Thanasimus dubius (Fabricius) (Coleoptera: Cleridae), captured during an annual springtime trapping survey are used to make forecasts of the likelihood and severity of an outbreak during the following summer. We investigated responses by both species to six lure formulations to evaluate their suitability for the survey and allow integration of historical data sets produced with differing lure compositions. Trapping trials were performed at four locations across three states (Louisiana, Mississippi, and Alabama) during spring, and at these and one additional location (North Carolina) in fall 2016. All lures included the pheromone component frontalin. Southern pine beetle preferred lures that additionally included the pheromone component endo-brevicomin and turpentine as a source of host odors (rather than a 7:3 mixture of monoterpenes alpha- and beta-pinene). Thanasimus dubius displayed little discrimination among lure compositions. Lure preferences by southern pine beetle did not differ significantly among locations in spring but were influenced by season. Gas chromatography (GC)-electroantennographic detection analyses with southern pine beetle and GC-mass spectrometry identified numerous known and potential semiochemicals that distinguished volatiles released by the tested host odor devices. The lure combination that included endo-brevicomin and alpha/beta-pinene is recommended for the trapping survey because of its high sensitivity for southern pine beetle and potential for greater data integrity resulting from its reproducible composition.

Southern Pine Beetle (Coleoptera: Curculionidae: Scolytinae) Pheromone Component trans-Verbenol: Enantiomeric Specificity and Potential as a Lure Adjuvant.

The southern pine beetle, Dendroctonus frontalis Zimmermann (Coleoptera: Curculionidae: Scolytinae) utilizes a multi-component aggregation pheromone to mediate mass-attacks and thereby colonize otherwise unsusceptible trees. Females produce the attractant frontalin and a synergist, trans-verbenol. We investigated trans-verbenol to determine whether enantiomeric composition, airborne concentration, and possibly other factors might affect its biological activity. Newly-emerged females from Mississippi populations produced 54-87% of the (-)-enantiomer; females initiating galleries in logs produced lower amounts and a wider range of enantiomeric ratios [12-92% (-)]. Coupled gas chromatography-electroantennographic detection (GC-EAD) studies did not suggest large differences in the concentration threshold of olfaction for the two enantiomers. We examined the effect of adding trans-verbenol to traps located outside infested areas and baited with components of the aggregation attractant. Male attraction was similarly increased by lures with 3, 81, or 98% of the (-)-enantiomer of trans-verbenol, whereas females preferred 81 over 3%. When release rate of 81% (-)-trans-verbenol in traps was varied across three orders of magnitude (0.3, 3, and 30 mg/d), the data suggested a positive dose-response trend. A high release (i.e., 2-5 g/d) device of host-odor alpha-pinene had a much stronger enhancing effect on trap catches than a trans-verbenol device (~30 mg/d), and trans-verbenol did not further enhance attraction when alpha-pinene was present. Our results suggest that the weak attraction-enhancing activity of trans-verbenol reported previously cannot be improved by adjusting the enantiomeric composition or release rate of lures, and furthermore there are no anticipated benefits of adding trans-verbenol to the D. frontalis monitoring lure.

Characterization of an aggregation pheromone in Hylesinus pruinosus (Coleoptera: Curculionidae: Scolytinae).

We conducted laboratory and field bioassays to characterize the pheromone system of an ash bark beetle, Hylesinus pruinosus Eichhoff (Coleoptera: Curculionidae: Scolytinae). Solitary females in newly initiated galleries in ash logs produced (+)-exo-brevicomin, whereas male beetles paired with females produced (+)-endo-brevicomin, lesser quantities of (+)-exo-brevicomin, and a third compound that could not be identified. Beetles produced these compounds also after exposure to juvenile hormone III, and they were the sole volatile chemicals isolated from beetles or aerations of infested logs that elicited electrophysiological responses from antennae of either sex. In the field, both sexes were strongly attracted to traps baited solely with either racemic or pure (+)-endo-brevicomin. Racemic exo-brevicomin was much less attractive to both sexes than racemic endo-brevicomin, and it did not increase attraction of endo-brevicomin when released in combination. Host odors (volatiles from mechanically damaged ash branches) failed to attract beetles or increase attractiveness of racemic exo-brevicomin. Our evidence suggests that male-produced (+)-endo-brevicomin is the major component of an aggregation pheromone for H. pruinosus, with (+)-exo-brevicomin and the unidentified male compound playing an indeterminate role in the chemical ecology of this species. Our data thus show an instance in which the major aggregation pheromone component of a bark beetle is produced by the secondarily arriving sex, a rare occurrence in bark beetles but one which has been reported previously for the Hylesini.

Evidence that (+)-endo-brevicomin is a male-produced component of the Southern pine beetle aggregation pheromone.

Previous research indicated that the aggregation pheromone of the southern pine beetle, Dendroctonus frontalis, is produced only by females, the sex that initiates attacks. We provide evidence indicating that secondarily arriving males augment mass aggregation by releasing the attractive synergist (+)-endo-brevicomin. Healthy pines artificially infested with both sexes of D. frontalis were significantly more attractive to conspecifics than trees infested solely with females. Coupled gas chromatography-electroantennographic detection (GC-EAD) analyses of volatiles isolated from male beetles revealed substantially greater olfactory sensitivity by D. frontalis to endo-brevicomin than to any other component. The threshold of detection of both sexes for (+)-endo-brevicomin was four orders of magnitude lower than for its antipode and at least one order of magnitude lower than for either enantiomer of frontalin, the major female-produced aggregation pheromone component. Pairing with a female in a gallery stimulated individual male beetles to produce hundreds of nanograms of (+)-endo-brevicomin. (+)-endo-Brevicomin was detected in a small percentage of female D. frontalis, whereas (-)-endo-brevicomin was never detected in either sex. In field trapping bioassays, we confirmed that (+)-endo-brevicomin is a potent synergist for attractive combinations of frontalin and pine turpentine. However, (+)-endo-brevicomin failed to attract D. frontalis either when presented alone or in combination with turpentine. We postulate that mass colonization of host trees by D. frontalis is mediated by distinct semiochemicals from both sexes rather than females alone. Our discovery of a key aggregation pheromone component in such an apparently well-studied species implies that the pheromone models of other bark beetles could benefit from systematic reexamination using newer technologies. Additionally, baits fortified with (+)-endo-brevicomin may enhance pest management strategies that exploit attractants for D. frontalis.

Electrophysiological and olfactometer responses of two histerid predators to three pine bark beetle pheromones.

We measured electrophysiological responses in the antennae of two predaceous hister beetles, Platysoma parallelum and Plegaderus transversus, exposed to racemic mixtures of primary aggregation pheromones of scolytid bark beetle prey, ipsenol, ipsdienol, and frontalin. No significant differences were found for either histerid species between male and female antennal responses to any of the three pheromones. Measurement of antennal threshold responses indicated that Pla. parallelum has increasing antennal sensitivity to ipsdienol, ipsenol, and frontalin. In contrast, Ple. transversus exhibited similar detection thresholds to all three pheromones. Pla. parallelum antennae exhibited different response amplitudes to the three pheromones at quantities above the detection threshold, while Ple. transversus had similar responses to each. Behavioral responses to the same three pheromones were evaluated for both histerid species using pedestrian olfactometer bioassays. Both species were attracted to frontalin and ipsenol, but not ipsdienol. Pla. parallelum was significantly more attracted to frontalin than ipsenol, while Ple. transversus showed no significant preference for either compound. Our results suggest that histerids that prey upon pine bark beetles may have different host or host habitat preferences, which could reduce interspecific competition.