Female Sex Pheromone Polymorphism in Adzuki Bean Borer, Ostrinia scapulalis, Is Similar to That in European Corn Borer, O. nubilalis

Individual analysis of the female sex pheromone of the adzuki bean borer, Ostrinia scapulalis, has shown that the sex pheromone of this species comprised (E)-11-tetradecenyl acetate (E11–14:OAc) and (Z)-11-tetradecenyl acetate (Z11–14:OAc) at variable blend ratios. The pheromone blend could be tentatively categorized into three types with respect to the proportion of E11–14:OAc: E type (94–100%, median 99.2%), Z type (0–16%, median 3.0%), and intermediate type (I type, 48–85%, median 63.7%). In addition to the identity of components, the blend ratios in the three types were similar to those of the E strain, Z strain, and hybrid of the European corn borer, O. nubilalis, respectively. This finding suggests that two closely related but morphologically distinct species, O. scapulalis and O. nubilalis, share almost the same sex pheromone communication systems. The significance of this similarity in the two sibling species is discussed.     

Response of male eye-spotted bud moth,Spilonota ocellana (Lepidoptera: Tortricidae), to different pheromone blends in north America and The Netherlands

Response of male eye-spotted bud moth,Spilonota ocellana (Denis and Schiffermüller), to different ratios of synthetic sex pheromone components, (Z)-8-tetradecenyl acetate (Z8-14:OAc) and (Z)-8-tetradecenyl alcohol (Z8-14:OH), were compared in four North American locations and in one location in The Netherlands. In British Columbia, Nova Scotia, Michigan, and The Netherlands, a 99:1 blend ofZ8-14:OAc andZ8-14:OH captured significantly more maleS. ocellana thanZ8-14:OAc alone or binary blends containing 10–50%Z8-14:OH. In Ontario, where population sizes were low compared to the other four locations, trends in trap catches were similar, and there was no indication that maleS. ocellana responded differently to the tested pheromone blends. A 99:1 blend ofZ8-14:OAc andZ8-14:OH should be most effective in pheromone-based control programs ofS. ocellana in North America and in The Netherlands. Our results confirm earlier studies that a 99:1 blend ofZ8-14:OAc andZ8-14:OH captures significantly more maleS. ocellana thanZ8-14:OAc alone. However, our finding that a 99:1 blend ofZ8-14:OAc andZ8-14:OH is significantly more attractive than binary blends containing 10–50%Z8-14:OH differs from previous findings in Germany and Switzerland.Agriculture Canada Summerland Research Station Contribution No. 821.     

Sex Pheromones and Reproductive Isolation of Three Species in Genus <Emphasis Type="Italic">Adoxophyes</Emphasis>

We tested differences in female pheromone production and male response in three species of the genus Adoxophyes in Korea. Females of all three species produced mixtures of (Z)-9-tetradecenyl acetate (Z9–14:OAc) and (Z)-11-tetradecenyl acetate (Z11–14:OAc) as major components but in quite different ratios. The ratio of Z9–14:OAc and Z11–14:OAc in pheromone gland extracts was estimated to be ca. 100:200 for Adoxophyes honmai, 100:25 for Adoxophyes orana, and 100:4,000 for Adoxophyes sp. Field tests showed that males of each species were preferentially attracted to the two-component blends of Z9–14:OAc and Z11–14:OAc mimicking the blends found in pheromone gland extracts of conspecific females. The effects of minor components identified in gland extracts on trap catches varied with species. Addition of 10-methyldodecyl acetate (10me-12:OAc) or (E)-11-tetradecenyl acetate (E11–14:OAc) to the binary blend of Z9–14:OAc and Z11–14:OAc significantly increased captures of A. honmai males, whereas E11–14:OAc exhibited a strongly antagonistic effect on catches of Adoxophyes sp. males. Moreover, (Z)-9-tetradecen-1-ol (Z9–14:OH) or (Z)-11-tetradecen-1-ol (Z11–14:OH) added to the binary blends increased attraction of male A. orana but not A. honmai and Adoxophyes sp. males, suggesting that these minor components, in addition to the relative ratios of the two major components, play an important role in reproductive isolation between Adoxophyes species in the southern and midwestern Korea where these species occur sympatrically.     

Identification of sex pheromone of bristly cutworm,Lacinipolia renigera (Stephens)

The sex pheromone of the bristly cutworm moth,Lacinipolia renigera was identified as a blend of (Z)-9-tetradecenyl acetate (itZ9–14): Ac and (Z, E)-9,12-tetradecadienyl acetate (ZE-9,12–14: Ac). Extracts of female glands were analyzed by capillary gas chromatography on three columns of different polarities. In each analysis, peaks with retention times identical to Z9–14:Ac andZE–9, 12–14: Ac were observed. GC-MS analysis of gland extracts supported the identification of these two compounds. Volatiles emitted from female sex pheromone glands during 10-min collection periods contained 7.8 ±2.01 ng ofZ9- 14: Ac. On average the blend contained 3.8 ± 1.43%ZE-9,12–14: Ac. Blends ranging from 1% to 10%ZE- 9,12–14: Ac in Z9-14: Ac (1 mg) were effective in capturing males in the field. The number of males captured in traps baited with a 3 % blend ofZE- 9,12-14: Ac in Z9-14: Ac was not significantly different than the number caught in traps containing two virgin females.     

Individual variation in sex pheromone of smaller tea tortrix moth,Adoxophyes sp. (Lepidoptera: Tortricidae)

Female smaller tea tortrix mothsAdoxophyes sp. (Lepidoptera: Tortricidae), which initiated calling at 1, 2, or 3 days old, respectively, were analyzed individually for (Z)-11-tetradecenyl acetate (Z11-14:OAc) and (Z)-9-tetradecenyl acetate (Z9–14: OAc) in the pheromone gland via GLC. Among different age groups, broad and similar distributions were found for pheromone quantity (¯X=58.6±52.9 ng/female; range 1.3–219.8 ng/female). The ratio of the two pheromone components averaged 6535 but ranged from 8416 to 4060. The significance of the pheromone blend variation to the attraction of males was tested in a field experiment. The ratio of males trapped by the most attractive blend versus the least attractive one was 2.2.     

Calling behavior and pheromone titer in the smaller tea tortrix moth,Adoxophyes sp. (Lepidoptera: Tortricidae)

The calling behavior and pheromone titer in the female smaller tea tortrix moth,Adoxophyes sp., were investigated under a 1410-hr light-dark photoperiod. Quantitative gas chromatographic analysis of ovipositor extract for (Z)-11-tetradecenyl acetate (Z11–14Ac) and (Z)-9-tetradecenyl acetate (Z9–14Ac), the major pheromone components of this species, obtained on the third day postemergence, indicated that extractable amounts of sex pheromone were present throughout the period of observation. Maximal pheromone titer and calling activity was reached at 8 and 10 hr after onset of scotophase, respectively. The ratio ofZ11–14Ac toZ9–14Ac through the 24-hr period varied significantly. The significance of the sex pheromone component ratio variation on the attraction of males was tested in a field experiment. The ratio of males trapped by the most attractive blend versus the least attractive one was 2.16.     

Sex pheromone components isolated from china corn borer,Ostrinia furnacalis guenée (lepidoptera: Pyralidae), (E)- and (Z)-12-tetradecenyl acetates

The sex pheromone components from the corn borer spreading widely in China,Ostrinia furnacalis Guenée, have been identified as (E)-and (Z)-12-tetradecenyl acetates (E andZ12–14 Ac). The ratio ofE isomer toZ isomer was 53 47. Traps containing 1 × 10^–7–1 × 10^–5 g of these compounds captured more males than did live females or their tip extract (3–6 female equivalents). Tetradecyl acetate (14 Ac) was also identified in the tip extract. Its quantity was about 1.8 times the sum of the other two isomers. However, including this compound in its natural ratio in pheromone traps resulted in a decrease in trap catches (P< 0.05).     

Sex pheromone of aPlanotortrix excessana sibling species and reinvestigation of related species

The sex pheromone of aPlanotortrix excessana sibling species was investigated. Females were found to produce eight potential pheromone components: dodecyl acetate, tetradecyl acetate (14OAc). (Z)-5-tetradecenyl acetate (Z5-14OAc), (Z)-7-tetradecenyl acetate (Z7-14OAc), (Z)-9-tetradecenyl acetate, hexadecyl acetate, (Z)-7-hexadecenyl acetate, and (Z)-9-hexadecenyl acetate. When these compounds were bioassayed using field-trapping and wind-tunnel techniques, only 14OAc,Z5-14OAc, andZ7-14OAc were found to be behaviorally active. The sex pheromone glands of females of other species including,Planotortrix MBS,Planotortrix M,P. notophaea, Ctenopseustis servana, and aC. obliquana sibling species, were also found to containZ5-14OAc orZ7-14OAc, singly or in combination. In the case ofPlanotortrix M, the addition ofZ7-14OAc to the previously identified sex pheromone blend ofZ5-14OAc and 14OAc was found to increase trap captures of male moths of this species. Thus in these New Zealand species (and in some Australian species),Z5-14OAc andZ7-14OAc appear to be utilized in combination in pheromonal communication just as (Z)-11-tetradecenyl acetate and (E)-11-tetradecenyl acetate are used by many species of Holarctic Tortricidae in the tribe Archipini.Lepidoptera: Tortricidae: Tortricinae.     

Sex pheromone components of the blueberry leaftier mothCroesia curvalana kearfott (Lepidoptera: Tortricidae)

The GC-MS and GC-EAD analyses of sex-gland components and sex-gland-released volatiles have identified (E)- and (Z)-11-tetradecenal (9010) and (E)- and (Z)-11-tetradecenyl acetate (8515) as components of the sex pheromone chemical communication system ofCroesia curvalana (Kearfott). The aldehyde-to-acetate ratio of 91 was found in the gland-released components. This blend at a 0.1–1 g/day release rate from PVC lures was shown to be as effective as virgin females in the trapping of maleC. curvalana.     

Comparative analysis of sex-pheromone-response antagonists in three races of European corn borer

Behavioral responses of males from three New York races of European corn borer,Ostrinia nubilalis (Hübner), to various suspected sexpheromone-response antagonists were investigated with a flight tunnel protocol. Males from a bivoltine (Z) race and from a univoltine (Z) race utilizing a natural pheromone blend of 982 (Z)-(E)-11-tetradecenyl acetates and males from a bivoltine (E) race utilizing a 199 (Z)-(E)-11-tetradecenyl acetate ratio were flown to their natural blends and to blends containing 1 % of one of the following additional compounds: Z9–12OAc,E9–12OAc,Z9–14OAc, andE9–14OAc. In each race, the added components lowered significantly the number of individuals completing the behavioral sequence. The only exception was bivoltineE males flown to a blend withE9–12OAc added. The number completing the sequence in this case was statistically not significantly lower than the number that completed the sequence to the standard blend. In all three races, theZ9–14OAc produced the most dramatic reduction in completed flights, and in the univoltineZ race, this added component was significantly more effective in reducing completed flights than any other added component. The response to the Z9–14OAc is understandable given recently published findings of an electrophysiological study of single sensilla in the European corn borer.     

Identification of female sex pheromone components of rice looper,Plusia festucae (L.), (Lepidoptera: Noctuidae)

The rice looper,Plusia festucae, is a defoliator of the rice plant. Chromatographic behavior, chemical reactions, and GC-MS analyses of the female sex pheromone revealed that the main component was (Z)-5-dodecenyl acetate (Z5–12: OAc, component I). The GC-MS analysis also indicated that the pheromone gland extract included another three monounsaturated components, (Z)-5-dodecen-l-ol (Z5–12: OH, component II), (Z)-7-tetradecenyl acetate (Z7–14: OAc, component III), and (Z)-7-tetradecen-l-ol (Z7–14: OH, component IV) in the following ratio: I:II:III:IV=100:6:15:1. In a paddy field, the mixture of synthetic I, II, and III in a ratio of 100:6:15 showed stronger attractancy than the virgin female, while the role of IV was unknown.     

Sex pheromones and reproductive isolation in four european small ermine moths

Reproductive isolation among four sympatric small ermine moths (Yponomeuta) is analyzed in terms of niches in the sexual communication channel. Potential pheromone components were identified from pheromone gland secretions ofY. evonymellus,Y. cagnagellus,Y. padelius, andY. vigintipunctatus by gas chromatography with flame ionization and electroantennographic detection and tested for behavioral activity in the field. The species were found to share (Z)-11-tetradecenyl acetate (Z11-14: OAc) in combination with varying proportions of theE isomer as primary sex pheromone components.Y. cagnagellus differs from the rest of the species by having only a small amount (1.5%) ofE isomer relative toZ isomer in its pheromone. The closely relatedY. padelius has a three-component pheromone including large amounts (ca. 400%) of (Z)-11-hexadecenyl acetate (Z11–16:OAc) in addition toZ11–14: OAc (100%) andE11–14: OAc (34%).Z11–16: OAc appears to reduce trap catches ofY. evonymellus andY. vigintipunctatus when added to the pheromone. Although these species are the two most distantly related European small ermine moths, they seem to share the same sex pheromone, i.e.,Z11–14OAc (+20%E) and (Z)-11-tetradecenol. Our interpretation is that this might have been the sex pheromone of the ancestor of today'sYponomeuta species.1 Lepidoptera: Yponomeutidae Latr.     

Changes in pheromone titer of oblique-banded leafroller,Choristoneura rosaceana,virgin females as a function of time of day,age, and temperature

Under a 16:8 hr light-dark photoperiod and 20C constant temperature, the titers of (Z)-11-tetradecenyl acetate (Z11–14:Ac), (E)-11-tetradecenyl acetate (E11–14:Ac) and (Z)-11-tetradecenol (Z11–14:OH) produced by different-agedChoristoneura rosaceana virgin females varied significantly during the scotophase, with the maximum titer occurring before the onset of calling in day-0 and day-3 females, while in day-5 females the titer remained constant throughout the calling period. There was a significant decrease in the titer of all pheromone components with age, explaining the lesser attractiveness of day-5 females relative to day-0 and day-3 females observed in the field. Under a cold thermocycle simulating condition during the second flight period in the fall, the titers of all pheromone components did not vary with time of day. There was a significant decrease in the amount ofZ11–14:Ac with age but no changes occurred in the minor components. Furthermore, for any given age tested, the amount of each component produced during the period of maximal calling activity remained relatively similar at the two temperature regimes. However, as with the expression of calling behavior, pheromone production was initiated earlier at cooler than at warmer temperatures. At both temperature regimes, female age and time of day influenced the ratio of each pheromone component. These results are discussed in relation to the hypothesis that by calling earlier, less attractive older females may increase their probability of mating.     

Identification of sex pheromone component of spruce budmothZeiraphera canadensis

The analyses of virgin female sex pheromone gland extracts by gas chromatography (GC), GC-electroantennographic detection (GC-EAD) and GC-mass spectrometry (GC-MS) followed by field-trapping experiments, have identified (E)-9-tetradecenyl acetate (E9–14:Ac) as the primary sex pheromone component of the spruce budmoth,Zeiraphera canadensis. Dosages of 1.0–100.0 g ofE9–14:Ac impregnated in rubber septa provide effective trap baits.Lepidoptera: Tortricidae: Eucosminae.     

Sex Pheromone of Apple Blotch Leafminer, Phyllonorycter crataegella, and Its Effect on P. mespilella Pheromone Communication

(Z)-10,(Z)-12-Tetradecadienyl acetate (Z10,Z12–14:OAc) and (E)-10,(E)-12-tetradecadienyl acetate (E10,E12–14:OAc) are sex pheromone components of the apple blotch leafminer (ABLM), Phyllonorycter crataegella. Compounds extracted from female pheromone glands were identified by coupled gas chromatographic–electroantennographic detection (GC-EAD) analyses, retention index calculations of EAD-active compounds, and by comparative GC-EAD analyses of female ABLM-produced and authentic (synthetic) compounds. In field experiments in apple Malus domestica orchards in Connecticut, Z10,Z12–14:OAc alone attracted ABLM males. Addition of E10,E12–14:OAc to Z10,Z12–14:OAc at 0.1:10 or 1:10 ratios enhanced attractiveness of the lure. Geometrical isomers Z10,E12- or E10,Z12–14:OAc at equivalent ratios were behaviorally benign and slightly inhibitory, respectively. In field experiments in British Columbia, Z10,Z12–14:OAc plus E10,E12–14:OAc did not attract Phyllonorycter moths, supporting the contention that ABLM is not present in the fruit growing regions of British Columbia. Z10,Z12–14:OAc added to P. mespilella pheromone, (E)-4,(E)-10-dodecadienyl acetate, strongly inhibited response by P. mespilella males. Recognition of the ABLM pheromone blend by allopatric P. mespilella males suggests a phylogenetic relationship and previous sympatry of these two Phyllonorycter spp. If pheromonal attraction of ABLM males were reciprocally inhibited by P. mespilella pheromone, a generic Phyllonorycter pheromone blend could be tested for pheromone-based mating disruption of the apple leaf-mining Phyllonorycter guild in North America.     

Response of male potato stem borer moths,Hydraecia micacea (Esper) to conspecific females and synthetic pheromone blends in the laboratory and field

Behavior of males ofHydraecia micacea (Esper) responding to virgin females and to synthetic pheromone blends were investigated in a laboratory wind tunnel. The synthetic blend consisted of saturated 14Ac (68.9%),Z9–14Ac (3.4%),E11–14Ac (14.6%), andZ11–14Ac (13.1%). Virgin females were significantly better lures than the four-component synthetic blend for most behaviors. By deleting components individually from the four-component blend,Z9–14Ac,Z11–14Ac, and saturated 14Ac were found to be necessary for communication butE11–14Ac was found to have no effect on typical pheromone-mediated reproductive behaviors. Close-range studies suggested that chemicals of low volatility, released from moths, were important in eliciting copulation attempts. Field studies reinforced laboratory findings regarding the effectiveness of different lures and indicated thatHeliothis traps were the most effective for monitoring.     

Sex pheromone ofAdoxophyes orana: Additional components and variability in ratio of (Z)-9- and (Z)-11-tetradecenyl acetate

Twelve products related to the sex pheromone main components (Z)-9- and (Z)-11-tetradecenyl acetate (Z9–14Ac andZ11–14Ac, respectively), were identified in female pheromone gland extracts of the laboratory-reared summerfruit tortrix moth,Adoxophyes orana F.v R. These are the geometric isomers and the alcohols of the main components, (Z)-9-dodecenyl acetate, (Z)-11-hexadecenyl acetate, and saturated acetates of 12–22 carbons. The ratio ofZ9–14Ac toZ11–14Ac in individuals varied from 3.51 to 111 with an average of 6.2; their total added up to 462 ng/female with an average of 182 ng for 2- to 7-day-old individuals. No qualitative or quantitative differences were observed between laboratory and field insects.Z9–14Ac,Z11–14Ac and the corresponding alcohols were also found in female effluvia. Addition of either of the two alcohols to a blend of the two acetates augmented trap catch in the field. The same was true for (Z)-9,(E)-12-tetradecadienyl acetate which was not detected in gland extracts.     

Differences in sex pheromone communication systems of closely related species:Spodoptera latifascia (walker) andS. descoinsi lalannecassou & silvain (Lepidoptera: Noctuidae)

S. latifascia andS. descoinsi are closely related species that occur sympatrically over limited areas in French Guiana. We examined allopatric populations,S. latifascia originating from Barbados andS. descoinsi from French Guiana. Studies on nocturnal activity cycles showed temporal partitioning of female calling behavior, male sexual activity, and mating behavior.S. descoinsi were sexually active in the first half of the scotophase whereasS. latifascia were sexually active in the second half. Seven compounds (Z9–14: Ac,Z9,E12–14: Ac,Z11–16: Ac,E9,E12–14: Ac,Z9–14: Ald,Z9,E11–14: Ac andZ11–14: Ac) were identified in females of bothS. latifascia andS. descoinsi extracts.Z9–14: Ac was a main pheromone component for the two species. The major difference between the pheromones ofS. latifascia andS. descoinsi was the proportion ofZ9,E12–14: Ac in the extracts: 7% forS. latifascia and 42% forS. descoinsi. The proportion ofZ9,E12–14: Ac relative to the sum ofZ9–14: Ac andZ9,E12–14: Ac in individual gland extracts was 4±1% (mean ± standard deviation) forS. latifascia and 44.8±6% forS. descoinsi. Electrophysiological studies showed no major differences between species in the morphology and physiology of the pheromone receptors of males. Receptors were identified forZ9–14: Ac andZ9,E12–14: Ac, but no receptor was found for the other compounds. In the wind tunnel, synthetic blends withZ9–14: Ac andZ9,E12–14: Ac gave the same behavioral responses as conspecific female extracts for the males of the two species. Some cross-attraction was observed with synthetic blends and female extracts. Nethertheless, previous field trapping experiments in French Guiana were species-specific and suggested differences in the attractivity of males. In the laboratory,S. latifascia andS. descoinsi could hybridize in both reciprocal crosses. FemaleS. descoinsi × maleS. latifascia mating rate was significantly lower than for the reciprocal cross, and 26.7% of femaleS. descoinsi could not separate from maleS. latifascia after mating. These copulatory problems may involve genital incompatibilities between males and females. Several barriers against interbreeding betweenS. latifascia andS. descoinsi seem to combine including differences in nocturnal activity cycles, pheromone differences, and genital barriers. The study of sympatric populations will be necessary to define the role of sex pheromones in the reproductive isolation ofS. latifascia andS. descoinsi.     

Mint root borer,Fumibotys fumalis (Guenee): Identification and field tests of female sex pheromone gland components

Compounds identified in sex pheromone gland extracts of femaleFumibotys fumalis (Guenee) consisted of (E,E)-10,12-tetradecadienyi acetate, (Z)-11-tetradecenyl acetate, (E)-11-tetradecenyl acetate, and (Z)-9-tetradecenyl acetate in a ratio of 100:18: 8:4, respectively. The compounds were identified by electroantennographic, gas Chromatographic, mass spectrometric, and chemical derivatization procedures. In mint fields synthetic components in gray elastomeric septa at ratios found in the sex pheromone gland and at doses of 3 or 10 mg of the diene produced trap catch comparable to traps baited with three females.Lepidoptera: Pyralidae     

Sex pheromone of a conifer-feeding budworm,Choristoneura retiniana,Walsingham

Sex pheromone components collected from femaleChristoneura retiniana included 11-tetradecenyl acetates and alcohols. The major component wasE-11-tetradecenyl acetate (E11–14Ac) with a lesser amount ofZ isomer necessary to induce male response. A 928 ratio ofE,Z11–14 Ac appeared optimal. The alcohol component was present at about 10% of the total pheromone mixture, and traps baited with acetates plus alcohol surpassed unmated females in their degree of attractiveness. Chemical analysis indicated a 9:1 ratio of theE — Z isomers of 11-tetradecenyl alcohol (11–14ol) pheromone components, although bait formulations containing a predominance of either theE orZ isomers were equally successful in field bioassays. Based on male response to traps,E- andZ11–14Ac (928E — Z) are essential pheromone components for long-range sex attraction. The 11–14 ol enhanced attraction when added at 10% of the total pheromone blend.This paper reports the results of research only. Mention of a commercial product does not constitute a recommendation by the U.S. Department of Agriculture.