Primary Attraction of the Fir Engraver, Scolytus ventralis

In laboratory bioassays, Porapak Q-captured and steam-distilled volatiles from the bark of host trees, Abies grandis, particularly from root-rot-infected trees, attracted 50–70% of male and female fir engravers, Scolytus ventralis. Gas chromatographic–electroantennographic detection (GC-EAD) analyses of Porapak Q-captured bark volatiles revealed 19 EAD-active compounds of which 13 (mostly monoterpenes) were identified by GC–mass spectrometry (GC-MS). In separate field experiments, multiple-funnel traps baited with two blends of these 13 synthetic volatiles released at 280 and 340 mg/ 24 hr attracted 66 and 93% of the total S. ventralis captured, respectively. The clerid predator, Thanasimus undulatus, also responded strongly to the kairomonal volatiles. Additional experiments produced no evidence for aggregation pheromones in S. ventralis. These included laboratory bioassays and GC and GC-EAD analyses of Porapak Q-captured volatiles from male- and female-infested logs or trees undergoing mass attack in the field, GC analyses and/or bioassays of extracts from female accessory glands, extracted volatiles from emerged, attacking and juvenile hormone-treated beetles of both sexes, and videotape analysis of the behavior of attacking beetles on the bark surface. We argue against the hypothesis of pheromone-mediated secondary attraction in S. ventralis and conclude that the attack dynamics of this species can be explained solely by its sensitive primary attraction response to host volatiles.     

Secondary attraction inGnathotrichus retusus and cross-attraction ofG. sulcatus (Coleoptera: Scolytidae)

Laboratory bioassays indicated that frass produced byGnathotrichus retusus males and male gut extracts were attractive to both sexes of beetles. The frass became attractive to females within 2 days of the commencement of boring activity, and attractive frass was produced for at least lOdays. Attraction of frass rose sharply after pairing of males with females. However, this increase in attraction could be due to increased boring activity and pheromone production by males following establishment of the mutualistic fungus. In field trapping experiments, male-infested logs were always highly attractive, but in one experiment, female-infested logs and logs infested by both sexes were also attractive. Thus, females could be involved in secondary attraction. High cross-attraction ofG. sulcatus toG. retusus infested logs in field experiments, as well as a moderate but significant response byG. retusus to sulcatol in the laboratory, suggests that they share sulcatol as a common pheromone.Research supported by Canadian Forestry Service Science Subvention Grant, National Research Council of Canada Operating Grant No. A3881, National Science Foundation (U.S.A.) Operating Grant No. PCM74-13643-A02, and the Council of Forest Industries of British Columbia.     

Juglone reduces growth,nitrogenase activity,and root respiration of actinorhizal black alder seedlings

European black alder trees [Alnus glutinosa (L.) Gaertn.] fix nitrogen with nodular symbionts and are interplanted with valuable black walnut trees (Juglans nigra L.) to increase soil nitrogen fertility. However, on some soils interplanted alder can be killed by black walnut's allelochemical juglone. In order to better understand the effects of juglone directly on the growth, nitrogen fixation, and root respiration of black alder, we grew nodulated alders hydroponically in a nitrogen-free nutrient solution at juglone levels of 2 × 10^–6, 2 × 10^–5, and 0 molar (M). Results indicate that nitrogenase activity (acetylene reduction) of alders growing in 2 × 10^–3 M juglone was reduced relative to alders without added juglone after one day, and in 2 × 10^–6 M juglone after five days. Root respiration (CO₂ evolution) and the relative increase of plant fresh weight were reduced in the 2 × 10^–5 M juglone treatment. In a related experiment, black alder germinants were grown in Flanagan silt loam soil dosed with 10^–3, 10^–4, and 0 M juglone. The inhibitory effects of 10^–3 M juglone on radicle elongation ceased 22–37 days after juglone treatments were started, suggesting that this soil can readily detoxify juglone.     

Identification of Semiochemicals Associated with Jeffrey Pine Beetle, Dendroctonus jeffreyi

Analysis of frass extracts and aerations from adult beetles confirmed earlier studies identifying 1-heptanol as a principal pheromone constituent produced by female Dendroctonus jeffreyi Hopkins. The aerations also indicated that males produced 33% (+) and 67% (–)-frontalin, previously unreported for this species, and >99% (+)-exo-brevicomin. Both males and females were responsive to these compounds in electrophysiological studies. Laboratory bioassays and field trapping experiments indicated that males and females were attracted to a mixture of 1-heptanol plus the principal host resin component heptane, but females were less attracted than males. Inclusion of racemic exo-brevicomin at 0.1% concentration in the traps increased the response of females. Inclusion of racemic frontalin above a 1% concentration in the normally attractive lures blocked arrival of beetles to the traps. While the female-produced pheromone components and host synergists are very different between the sibling species D. jeffreyi and Dendroctonus ponderosae Hopkins, the principal male-produced compounds are the same, and the response of the female beetles to those compounds appears to be similar.     

Field Attraction of Hoplia communis to 2-Phenylethanol, A Major Volatile Component from Host Flowers, Rosa spp.

The attractiveness of volatile compounds from the floral scent of Rosa, one of the most preferred plants for adult Hoplia communis, was evaluated under field conditions. The beetles were attracted to most compounds tested, but 2-phenylethanol exhibited the highest capture rate. Catches increased with increasing emissions of between 9.1 and 287.2 mg/day. Catches in white traps were significantly larger (17.4-fold) than those in green traps when both were baited with anethole, an already known attractant; however, the trap color was not significant when a more attractive lure, 2-phenylethanol, was used. The use of a single funnel trap baited with 2 g of 2-phenylethanol at a heavily infested nursery exhibited promising results for mass trapping. Approximately 90,000 beetles of both sexes, which nearly corresponds to the estimated maximum population per 1000 m², were captured within six days.     

Identification of Female Sex Pheromone of the Rice Leaf Bug, Trigonotylus caelestialium

The female sex pheromone of the rice leaf bug, Trigonotylus caelestialium, was analyzed by GC-EAD and GC-MS. Ten EAD active compounds—n-hexyl n-hexanoate, (E)-2-hexenyl n-hexanoate, n-hexyl (E)-2-hexenoate, n-octyl n-butyrate, n-octyl n-hexanoate, (E)-2-octenyl n-hexanoate, n-pentyl n-hexanoate, n-hexyl n-butyrate, (E)-2-hexenyl n-butyrate, and n-hexyl (E)-2-butenoate—were detected in the ratio of 1000:414–491:trace–5:5–11:55–71:50–63:trace–3:225:90:32 from female body extracts, and in the ratio of 1000:271–342:10–43:1–3:58–78:14–19:trace:178:36:26 from male body extracts. Field trapping tests with these synthetic compounds indicated that n-hexyl n-hexanoate, (E)-2-hexenyl n-hexanoate, and n-octyl n-butyrate are pheromone components, and mixtures in ratios of 1000:400–500:10–100 were more attractive to males. Doses ranging from 4.29 g to 14.3 g of the three-component mixture in the ratio of 1000:400:30 loaded into glass capillary tubes were most attractive to males.     

Effects of Enantiomeric Blend of Verbenone on Response ofIps paraconfusus to Naturally Produced Aggregation Pheromone in the Laboratory

The aggregation pheromone produced by maleIps paraconfusus (Coleoptera: Scolytidae) tunneling in Monterey pine,Pinus radiata, logs was trapped on Porapak Q. A concentration of an extract of trapped volatiles that attracted beetles was determined in the laboratory through a concentration–response analysis of walking behavior of males and females. The interruptant effects of four concentrations of verbenone on response to a constant concentration of this naturally produced aggregation pheromone were tested with males and females. Independent of its enantiomeric composition [99.5% (S)-(–), 93.1 % (S)-(–), and 98.3% (R)-(+)], verbenone significantly reduced the percentage of females (but not males) reaching the attractant source. However, when the time required for beetles of both sexes taken together to reach the attractant source was considered, verbenone of higher enantiomeric purity had a greater effect on beetle behavior. Solutions of 99.5% (S)-(–)- and 98.3% (R)-(+)-verbenone increased the time required for beetles to reach the attractive source when compared to 93.1% (S)-(–)-verbenone. When pooled across enantiomeric blends, increasing concentrations of verbenone resulted in slower responses in beetles that reached the attractant source within 2 min. Males and females did not respond to verbenone alone.     

Aggregation Pheromone of Pityogenes knechteliand Semiochemical-Based Interactions with Three Other Bark Beetles

Gas chromatographic–electroantennographic detection and GC-mass spectrometric analyses of volatile extracts from male and female Pityogenes knechteli Swaine identified hexanol, (±)-ipsdienol, and (S)-(–)-ipsenol as male-produced candidate pheromone components. In a lodgepole pine, Pinus contorta var. latifolia Engelmann, forest in the southern interior of British Columbia, multiple-funnel traps baited with (±)-ipsdienol alone, (S)-(–)-ipsenol alone, or both caught 60%, 6%, and 23%, respectively, of all P. knechteli trapped; unbaited traps caught the remaining 11%. In another field trapping experiment, (S)-(+)-ipsdienol was as attractive as (±)-ipsdienol, and (R)-(–)-ipsdienol was behaviorally benign. (S)-(+)-Ipsdienol is thus concluded to be the principal aggregation pheromone component of P. knechteli. At low release rates, hexanol increased attraction of beetles to (±)-ipsdienol, or to (±)-ipsidienol plus (S)-(–)-ipsenol, but at high release rates hexanol decreased attraction, suggesting a role in preventing overpopulation in the host tree. On the basis of laboratory bioassays in which walking beetles were attracted to (S)-(–)-ipsenol, we hypothesize that (S)-(–)-ipsenol serves as a short-range attractant for P. knechteli. Three sympatric scolytids were also captured in field experiments as follows: the pine engraver, Ips pini (Say), to its pheromone (±)-ipsdienol; I. latidens LeConte to its pheromone (S)-(–)-ipsenol; and I. mexicanus (Hopkins), for which the pheromone is unknown, to (S)-(–)-ipsenol with (±)-ipsdienol. Although all four species attack lodgepole pine, we have never observed I. latidens or I. mexicanus attacking the same hosts at P. knechteli or I. pini. These results suggest that ipsenol and ipsdienol serve as synomones involved in promoting aggregation on the host tree, maintaining species-specific communication, and thus contributing to resource partitioning and reduced competition among the four species.     

Interruption of response ofDendroctonus brevicomis to attractive pheromone by release of pheromone at several rates and spacings

The number of western pine beetles,Dendroctonus brevicomis, trapped at the center of three 90 × 90-m plots was reduced during a 16-hr period after surrounding an attractive source ofexo-brevicomin, frontalin, and myrcene with 16, 48, or 168 sources of the same attractant. Compounds were released from the center of the plot at 1 mg/16 hr/compound, and from the surrounding sources at 4–280 mg/16 hr/compound. About half of these amounts was released between 1530 hr and 1930 hr, the period of peak beetle flight. No treatment differences were apparent in reduction of catch within the range of release rates and spacings tested. When compounds were released continuously from 168 stations for 17 days, catch at the center of a plot was generally lower than catch before or after this period, but fluctuated daily. More beetles were caught on traps hung on ponderosa pines within a plot and the number of these traps catching beetles was greater when compounds were released from all stations than from only the center station. Beetles caught on traps were attracted into the plots from the surrounding forest, but appeared to be dispersed within the plot when compounds were released from many stations.D. brevicomis attacked at least 91 trees in the plots, of which 25 were killed during two summers.Coleoptera: Scolytidae.Trade names are mentioned solely for information. No endorsement by the U.S. Department of Agriculture is implied.     

Dominicalure 1 and 2: Components of aggregation pheromone from male lesser grain borerRhyzopertha dominica (F.) (Coleoptera: Bostrichidae)

Volatiles from lesser grain borers,Rhyzopertha dominica (F.) (Coleoptera: Bostrichidae), were collected on Porapak Q, and those from the male beetles were shown to contain two compounds, that were attractive individually and in combination to both sexes. These compounds were identified as (S)-(+)-1-methylbutyl (E)-2-methyl-2-pentenoate and (S)-(+)-1-methylbutyl (E)-2,4-dimethyl-2-pentenoate by spectrometry and comparison with synthesized compounds. The two compounds have been assigned the trivial names dominicalure 1 and dominicalure 2, respectively. Synthesized samples of these compounds, individually and in combination, were effective in trapping both sexes in field studies.Presented in part at the Northeastern Regional Meeting of the American Chemical Society, June 25–28, 1978, Boston.Approved as TA-14927 by the director of the Texas Agricultural Experiment Station in cooperation with the USDA-FR-SEA.     

Gallery initiation byTomicus piniperda (Coleoptera: Scolytidae) on scots pine trees baited with host volatiles

In a trapping experiment, wood chips from freshly cut Scots pine trees attracted flying adults of the bark beetleTomicus piniperda (L.) Healthy Scots pine trees, which usually are not attacked byT. piniperda, were baited with chips from freshly cut trees. In other experiments trees were baited with a mixture of (–)--pinene, (+)-3-carene, and terpinolene; the individual monoterpenes; or with ethanol alone. All baited treatments were attacked byT. piniperda. Most of the attacks resulted in short egg galleries and in larval galleries which were only a few millimeters in length. In trees from which a new generation of beetles emerged, net reproduction was well below 1.0. Unbaited control trees remained unattacked or received a few isolated attacks.     

Opposing Effects of Host Monoterpenes on Responses by Two Sympatric Species of Bark Beetles to Their Aggregation Pheromones

The effects of host tree monoterpenes on primary and secondary attraction of the bark beetles, Ips pini and Ips grandicollis, and their associated phloeophagous insects were investigated. Monoterpenes alone were not attractive to I. pini. However, monoterpenes mediated the attraction of I. pini to its aggregation pheromones. With the exception of 3–carene, the effect of monoterpenes on I. pini's response to its pheromone, ipsdienol plus lanierone, was inhibitory. In contrast, (–)--pinene both attracted I. grandicollis and enhanced the attraction of I. grandicollis to its pheromone, ipsenol. No monoterpene inhibited the response of I. grandicollis to its pheromone. The inhibitory effect of host monoterpenes on I. pini response to its aggregation pheromone differs from previous work, in which monoterpenes either synergized responses or had no effect. In addition to possible geographic differences, the concentrations used in our study simulated trees that had begun to respond to attack, whereas previous studies deployed lower concentrations, which simulated constitutive phloem from unattacked trees. These results support the view that trees that undergo induced responses to bark beetles sometimes inhibit attraction of additional beetles, despite the beetles' production of aggregation pheromones. Neither species displayed cross-attraction to the pheromone of the other. The red turpentine beetle, Dendroctonus valens, showed weak and consistent attraction to (+)--pinene and in some cases to (–)--pinene. Attraction to (–)--pinene was usually enhanced by Ips spp. pheromones. The absence of strong attraction to (+)--pinene and partial attraction to (–)--pinene suggest that the effects of different stereoisomers of -pinene on D. valens vary throughout its geographical range. Hylastes porculus was also attracted to some monoterpenes, particularly (–)--pinene. An additional 10 species of phloeophagous insects were caught in response to monoterpenes and/or pheromones, including the pine root weevils, Hylobius pales and Pachylobius picivorus, and the longhorned beetle, Monochamus carolinensis.     

An estimate of the effectiveness of pheromone-baited traps for the suppression ofScolytus multistriatus (Coleoptera: Scolytidae)

We attempted to suppress a population ofScolytus multistriatus (Marsham), the principal vector of Dutch elm disease, by trapping flying beetles on sticky traps baited with synthetic pheromone. The estimated catch on 421 traps distributed throughout a 1-km² plot in Detroit, Michigan, was nearly 1 million beetles. Because an estimated 5 million beetles emerged in the plot during the study period, we conclude that this preliminary trapping study had no appreciable effect on suppressing the population. Studies that employ improved materials and techniques are continuing.     

Influence of aggregation inhibitors (verbenone and ipsidenol) on landing and attack behavior ofDendroctonus brevicomis (Coleoptera: Scolytidae)

The influence of the aggregation inhibitors verbenone and ipsdienol on the response of western pine beetle,Dendroctonus brevicomis, to attractive host trees was investigated. Paired ponderosa pine trees (Pinus ponderosa) were baited with aggregation semiochemicals to stimulate mass attack. One tree in each pair received an inhibitor treatment consisting of five sets of two verbenone and two ipsdienol dispensers spaced 1 m apart vertically along the tree bole. Beetle landing was monitored with sticky traps on the tree bole, and attack density was assessed from bark samples removed four or seven days after baiting. The inhibitor treatment resulted in a significant reduction of both the numbers of beetles landing on trees and the density of attacking beetles compared to control trees (without inhibitors). The ratios of beetle landing density to attacking density were not different between inhibitor-treated and control trees, nor were the vertical distributions of beetles landing or attacking, suggesting that beetle behavior was primarily influenced at a longer range, prior to landing on the tree. Although the application of verbenone and ipsdienol did not preventD. brevicomis from attacking baited trees, our results suggest that when applied to unattacked (and unbaited) trees, their effectiveness at reducing the attack pressure might allow trees having a certain amount of resistance to survive attack by pioneer beetles.     

Evaluation of two attempts to trap defined populations ofScolytus multistriatus

Two experiments were conducted to investigate the use of a pheromone-based trap-out technique for suppressing populations ofScolytus multistriatus. In the first experiment, elm bolts containing the immature stages ofS. multistriatus were placed in an isolated community that contained elm trees, but which was essentially devoid of a resident beetle population. The infested bolts produced a total of 46,485 adult beetles of which 20% were recovered on traps baited with synthetic pheromone. In the second experiment beetles were released in a desert valley containing no resident elms or beetles. Only 1% of 20,000 released beetles were recaptured on traps erected on vertical cardboard cylinders and on elm logs. These rates of recapture are related to the attraction of beetles to naturally occurring brood sources versus pheromone-baited traps.Coleoptera: Scolytidae.     

Identification and Field Evaluation of Components of Female Sex Pheromone of Millet Stem Borer, Coniesta ignefusalis

Five active compounds were detected during analyses of ovipositor washings and effluvia from virgin female Coniesta ignefusalis moths by gas chromatography (GC) linked to electroantennographic (EAG) recording from a male moth. These were identified as (Z)-7-dodecen-1-ol (Z7–12:OH), (Z)-5-decen-1-ol (Z5–10:OH), (Z)-7-dodecenal (Z7–12:Ald), (Z)-7-dodecenyl acetate (Z7–12:Ac), and (Z)-9-tetradecen-1-ol (Z9–14:OH) by comparison of their GC retention times, mass spectra, and EAG activities with those of synthetic standards. Laboratory tests of dispensers for these compounds showed that release rates from polyethylene vials increased to relatively uniform values after three to four days, but release from septa was very rapid and nonuniform and decreased to low levels after two to three days. Trapping tests in Niger showed that the major component, Z7–12:OH, and two of the minor components, Z5–10:OH and Z7–12:Ald, were essential for attraction of male C. ignefusalis moths. The most attractive blend contained these three components in a 100:5:3.3 ratio in a polyethylene vial, which emitted the components in similar proportions to those produced by the female C. ignefusalis moth. Water traps baited with this blend containing 1 mg of Z7–12:OH caught more male C. ignefusalis moths than traps baited with newly emerged female moths. Addition of up to 10% of the corresponding E isomers of the pheromone components had no effect on catches, but addition of the other two minor components detected, Z7–12:Ac and/or Z9–14:OH, to the attractive blend at naturally occurring levels caused significant reductions in trap catch.     

Behavioral responses of elm bark beetles to baited and unbaited elms killed by cacodylic acid

Diseased elms, treated with various doses of cacodylic acid in northwest England, became attractive to elm bark beetles (Coleoptera: Scolytidae). This attraction seemed to be independent of pheromone baits. However attractive the trees became, they were unsuitable to the beetles as breeding sites since significantly more beetles visited the trees than were stimulated to penetrate and attempt to breed. It seems as if colonization of trap trees by the bark saprophytePhomopsis oblonga following cacodylic acid treatment made the trees unsuitable to beetles for breeding.     

Effects of verbenone andtrans-verbenol on the response ofDendroctonus brevicomis to natural and synthetic attractant in the field

Verbenone andtrans-verbenol were investigated as candidate interruptants for use as tree protectants. Verbenone andtrans-verbenol, pheromones released byDendroctonus brevicomis during host colonization, reduced the trap catch ofD. brevicomis near sources of the attractant composed ofexo-brevicomm, frontalin, and myrcene. Catch reduction at some trap positions was greater at a high release rate than at a low release rate oftrans-verbenol alone and of the combination of verbenone plustrans-verbenol. Verbenone also reduced catches at traps baited with attractive bolts from trees under attack byD. brevicomis. Attempts to use verbenone to protect living trees fromD. brevicomis attack were inconclusive.Coleoptera: Scolytidae.This research was supported in part by the Forest Service, U.S. Department of Agriculture, and in part by grants from the National Science Foundation/Environmental Protection Agency (grant NSF-6B-34718/BMS75-04223), and in part by grants from the Rockefeller Foundation to the University of California. Mention of a proprietary or commercial product does not constitute recommendation or endorsement of the product by the U.S. Department of Agriculture and does not imply its approval to the exclusion of other products that also may may be suitable.     

Identification of major component of male-produced aggregation pheromone of larger grain borer,Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae)

Male beetles of the larger grain borer,Prostephanus truncatus Horn (Coleoptera: Bostrichidae) were shown to produce material causing an electroantennographic (EAG) response from both male and female beetles. Volatiles from mixed sex cultures were collected on Porapak Q and fractionated by liquid chromatography (LC) with gradient elution. The total volatiles and the fractions were analyzed by gas chromatography (GC) linked to EAG, and two compounds eluting in the same LC fraction elicited EAG responses from male and female beetles. These were assumed to be components of the male-produced aggregation pheromone. Amounts of these compounds obtained were very low and less than 10^–3 times the amounts of the aggregation pheromone components produced by the related species,Rhyzopertha dominica, under similar conditions. The most abundant of these EAG-active compounds was identified as 1-methyiethyl (2E)-2-methyl-2-pentenoate by comparison of its GC retention times and mass spectrum with those of synthetic analogs. The synthetic compound elicited electrophysiological and behavioral responses in both male and female beetles significantly greater than those to structural analogs. This compound is attractive toP. truncatus beetles in the field, and has been given the trivial name of trunc-call 1.     

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.Deceased.