Sex Pheromone Blend Research Articles

Enhancement of attraction and trap catches of the old‐house borer, Hylotrupes bajulus (Coleoptera: Cerambycidae), by combination of male sex pheromone and monoterpenes

Monoterpenoid compounds extracted from wood of Scots pine, Pinus sylvestris L, synergized the attraction of the old-house borer, Hylotrupes bajulus (L), to the male pheromone (3R)-3-hydroxy-2-hexanone ((3R)-ketol) + 1-butanol. Glasshouse experiments using ground traps baited with extracts derived from Scots pine wood or the monoterpenes (+)-alpha-pinene, (-)-verbenone, (-)-trans-pinocarveol and (+)-terpinen-4-ol attracted significantly more H bajulus females, but caught fewer of them, than the synthetic pheromone mixture alone. Traps baited with higher concentrations of the monoterpene mixture attracted fewer females than those baited with lower dosages, whilst very high concentrations of the mixture (6-10 vials) caught no insects. However, a combination of (3R)-ketol + 1-butanol or (+/-)-3-ketol + 1-butanol with monoterpenes resulted in the capture of significantly more females than either the sex pheromone or the monoterpene mixture alone. Traps baited with a blend of the male's sex pheromone or the monoterpenes attracted significantly more, but caught fewer, males than females. Here again, a combination of the above blends enhanced the attraction of males significantly when compared with the attractancy of either of the compounds/mixtures used alone.

INTROGRESSING PHEROMONE QTL BETWEEN SPECIES: TOWARDS AN EVOLUTIONARY UNDERSTANDING OF DIFFERENTIATION IN SEXUAL COMMUNICATION

As a first step toward understanding how noctuid moths evolve species-specific pheromone communication systems, we hybridized and back-crossed two closely related moth species, Heliothis virescens (Hv) and H. subflexa (Hs), which differ qualitatively and quantitatively in their multicomponent sex pheromone blends. We used amplified fragment length polymorphism (AFLP) marker-based mapping of backcross families to determine which of the 30 autosomes in these moths contained quantitative trait loci (QTL) controlling the percentages of specific chemical components in the pheromone blends. In two previous backcrosses to Hs, we found a strong depressive effect of Hv-chromosome 22 on the percentage of three acetate components in the pheromone gland. These acetates are present in Hs and absent in Hv. Here, we describe how we introgressed Hv-chromosome 22 into the genomic background of Hs. Selection for Hv-chromosome 22 started from backcross 3 (BC3) females. All females that had Hv-chromosome 22 and a low percentage of acetates (<3% of the total amount of pheromone components present) were backcrossed to Hs males. In BC5 to BC8, we determined whether Hv-chromosome 22 was present by a) running only the primer pairs that would yield the markers for that chromosome, and/or b) determining the relative percentages of acetates in the pheromone glands. Either or both genotype and phenotype were used as a criterion to continue to backcross these females to Hs males. In BC9, we confirmed the isolation of Hv-chromosome 22 in the Hs genomic background, and backcrossed the males to Hs females to eliminate the Hv-sex chromosome as well as mitochondrial DNA. The pheromone composition was determined in BC3, BC5, and BC11 females with and without Hv-chromosome 22. All backcross females with Hv-chromosome 22 contained significantly less acetates than females without this chromosome. In addition, BC3 females with Hv-chromosome 22 contained significantly more Z11-16:OH than BC3 females without Hv-chromosome 22. However, in BC5 and BC11 females, the correlation between Z11-16:OH and Hv-chromosome 22 was lost, suggesting that there are separate QTL for the acetates and for Z11-16:OH, and that the relative amount of the alcohol component is only affected in epistasis with other (minor) QTL. Now that we have succeeded in isolating the chromosome that has a major effect on acetate production, we can test in behavioral experiments whether the presence of acetates may have been a driving force for a shift in pheromone composition. Such tests are necessary to move towards an evolutionary understanding of the differentiation in sexual communication in Heliothis spp. moths.

Expression and evolution of Δ 9 and Δ 11 desaturase genes in the moth Spodoptera littoralis

Desaturation of fatty acids is a key reaction in the biosynthesis of moth sex pheromones. The main component of Spodoptera littoralis sex pheromone blend is produced by the action of Δ 11 and Δ 9 desaturases. In this article, we report on the cloning of four desaturase-like genes in this species: one from the fat body (Sls-FL1) and three (Sls-FL2, Sls-FL3 and Sls-FL4) from the pheromone gland. By means of a computational/phylogenetic method, as well as functional assays, the desaturase gene products have been characterized. The fat body gene expressed a Δ 9 desaturase that produced ( Z)-9-hexadecenoic and ( Z)-9-octadecenoic acids in a (1:4.5) ratio, whereas the pheromone gland Sls-FL2 expressed a Δ 9 desaturase that produced ( Z)-9-hexadecenoic and ( Z)-9-octadecenoic acids in a (1.5:1) ratio. Although both Δ 9 desaturases produced ( Z)-9-tetradecenoic acid from myristic acid, transformed yeast grown in the presence of a mixture of myristic and ( E)-11-tetradecenoic acids produced ( Z, E)-9,11-tetradecadienoic acid, but not ( Z)-9-tetradecenoic acid. The Sls-FL3 gene expressed a protein that produced a mixture of ( E)-11-tetradecenoic, ( Z)-11-tetradecenoic, ( Z)-11-hexadecenoic and ( Z)-11-octadecenoic acids in a 5:4:60:31 ratio. Despite having all the characteristics of a desaturase gene, no function could be found for Sls-FL4.

Enhancement of Attraction to Sex Pheromones of Spodoptera exigua by Volatile Compounds Produced by Host Plants

We measured the effects of exposure to volatile compounds produced by host plants on the rate of capture of male Spodoptera exigua using synthetic sex pheromones. Exposure to volatile compounds stimulated strong electroantennographic responses of male S. exigua. The behavioral responses of male moths to combinations of sex pheromone and volatile compounds were tested in wind tunnel experiments. When lures were baited with synthetic sex pheromone plus benzaldehyde, phenylacetaldehyde, (Z)-3-hexenyl acetate, or linalool, respectively, the landing rate of S. exigua males was increased by 101.4%, 79.6%, 60.6%, and 34.3%, respectively, compared to sex pheromone alone. In field tests, traps baited with either pheromone + (E)-2-hexenal, pheromone + phenylacetaldehyde, pheromone + (Z)-3-hexenyl acetate, or pheromone + (Z)-3-hexenol enhanced moth catches by 38.8%, 34.6%, 24.6%, and 20.8%, respectively compared to traps baited with pheromone alone. In a second field experiment, more S. exigua males were trapped with a combination of a synthetic sex pheromone blend and several individual host plant volatiles compared to synthetic sex pheromone alone. These results suggest that some host plant volatiles enhance the orientation response of S. exigua male moths to sex pheromone sources.

Behavioral Activity of Stereoisomers and a New Component of the Contact Sex Pheromone of Female German Cockroach, Blattella germanica

(3S,11S)-3,11-Dimethylnonacosan-2-one is a major component of the courtship stimulating, contact sex pheromone of the female German cockroach. Although the four synthetic stereoisomers of this compound have been tested in behavioral assays, their relative activity remains unresolved. Using isolated male antennae dosed with synthetic test compounds to assay male behavior, we found that at high doses all four stereoisomers elicited responses from 100% of the males. However, at physiologically relevant doses similar to those found on the female antenna, the (3S,11S)-isomer was the least effective of the four stereoisomers at eliciting courtship responses in males. This is the first example of a natural stereoisomer having less bioactivity than related stereoisomers that do not occur naturally. Another component of the sex pheromone blend, 3,11-dimethylheptacosan-2-one, was previously purified from the female's epicuticule and behaviorally assayed, but its activity was not confirmed through synthesis. We now confirm that synthetic (3S,11S)-3,11-dimethylheptacosan-2-one elicits behavioral responses, but less so than its C29 homolog.

Sex Pheromones and Reproductive Behavior of Spodoptera litura (Fabricius) Moths Reared from Larvae Treated with Four Insecticides

When Spodoptera litura (Fabricius) larvae were treated with the insecticides deltamethrin, endosulfan, malathion, and carbaryl at dosages causing 30% mortality (LD30), female moths reared from survivors showed similar patterns in the frequency of calling behavior. However, females reared from larvae dosed with deltamethrin had significantly higher titers of sex pheromone than those treated with endosulfan, malathion, or carbaryl, or control individuals. The ratio of the sex pheromone blend, Z9,E12-14:OAc to Z9,E11-14:OAc in females from the deltamethrin treatment group differed significantly from that of female moths in other treatments. In a wind-tunnel bioassay, males reared from larvae treated with deltamethrin, malathion, or carbaryl were less likely to display behaviors resulting in the location of a sex pheromone source than males reared from larvae treated with endosulfan or a solvent control (acetone). Mating success was significantly decreased in pairs containing either males or females reared from larvae that survived treatment with deltamethrin.

Multiple acyl-CoA desaturase-encoding transcripts in pheromone glands of Helicoverpa assulta, the oriental tobacco budworm

Seven desaturase cDNAs were isolated from pheromone glands of Helicoverpa assulta, a moth producing a sex pheromone blend with high Z9-16:Ald and low Z11-16:Ald, opposite to what is found in other heliothine moths such as Helicoverpa zea. Six of the seven sequences map onto recently defined lepidopteran desaturase sequence lineages and the other is orthologous to a desaturase sequence previously reported only in H. zea. The levels of desaturase-encoding transcripts in pheromone glands were determined and the three most abundant ones were functionally expressed in a desaturase-deficient mutant strain of Saccharomyces cerevisiae. The HassNPVE transcript, shown to encode a Δ9 desaturase producing more Z9-18:Acid than Z9-16:Acid, was the most abundant, followed by the HassKPSE transcript, shown to encode a Δ9 desaturase producing more Z9-16:Acid than Z9-18:Acid, and by the HassLPAQ transcript, shown to encode a Δ11 desaturase producing only Z11-16:Acid. Thus, the relative amounts of transcripts encoding two Δ9 desaturases and a single Δ11 desaturase in H. assulta pheromone glands were consistent with the relative amounts of unsaturated fatty acid precursors required to produce the major and minor sex pheromone components of this species. Desaturase transcript levels in pheromone glands were also found to be as high during scotophase as during light phase, when pheromone production ceases. The other four transcripts were present at extremely low levels in H. assulta pheromone glands and the functional roles of their encoded desaturase-homologous proteins could not be determined.

Sex Pheromone Composition and Monitoring of the Oriental Fruit Moth, Grapholita molesta (Lepidoptera: Tortricidae) in Naju Pear Orchards

The sex pheromone blend of the oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae) from pear orchards of Naju, Korea, has been identified. Four candidate sex pheromone components in abdominal tip extracts of G. molesta female were detected by gas chromatography (GC). GC analysis indicated that it included (Z)-8-dodecenyl acetate (Z8-12: Ac), ( E)-8-dodecenyl acetate ( E8-12: Ac), (Z)-8-dodecenol (Z8-12: OH), and dodecanol (12: OH) in a ratio of 100:6.8:19.1:5.4. In the field trappings, Z8-12: Ac alone was not attractive to G. molesta male, and more males were caught by traps baited with Z8-12: Ac containing 4, 5 or 6 % E isomer than by traps baited with other Z: E mixtures. Addition of Z8-12: OH to the two acetates increased significantly catch of G. molesta males. Although a significant amount of Z8-12: OH was detected in the female abdominal tip extracts, addition of Z8-12: OH at the 1 % level of the total acetates was sufficient to enhance the trap catch. Traps baited with 95: 5: 1 blend of Z8-12: Ac, E8-12: Ac, and Z8-12: OH were successfully used to monitor G. molesta males flights. Analysis of seasonal trap catches over three years summarizes that the moth has four main generation peaks over at least seven months per year in Naju.

Electroantennogram responses of Apanteles obliquae (Hym., Braconidae) to various infochemicals

Abstract: The electroantennogram recording technique (EAG) was used to study the olfactory sensitivity of Apanteles obliquae (Hym., Braconidae), a gregarious larval endoparasitoid of Spilosoma obliqua (Walker) (Lep., Arctiidae), to 25 general plant volatiles belonging to alcohol, aldehyde and terpenoid groups and also to volatiles from the host and plant–host complex. The EAG data indicated different olfactory sensitivity between the sexes, not only to individual plant volatiles but also to the volatiles from host and plant–host complex. Females were found to be more responsive than males. However the synthetic sex pheromone blend of the host insect elicited similar EAG responses in both sexes. The EAG data of the present study is correlated with the reported behaviour observed in other parasitoids.

Specialized olfactory receptor neurons mediating intra- and interspecific chemical communication in leafminer moths Eriocrania spp. (Lepidoptera: Eriocraniidae).

We performed a physiological and morphological characterization of sensilla auricillica in male Eriocrania semipurpurella moths. Each auricillic sensillum contained three olfactory receptor neurons. Responding neurons (87 of 139) could be grouped into five physiological types. Type 1 responded to (R,Z)-6-nonen-2-ol and type 2 to its enantiomer (S,Z)-6-nonen-2-ol, both of which are pheromone components of E. semipurpurella. Type 3 responded to both (R)-heptan-2-ol and (R,Z)-4-hepten-2-ol, which are pheromone components of the sympatric species E. cicatricella. Types 4 and 5 responded to the ketones (Z)-6-nonen-2-one and/or nonan-2-one, which are found in the pheromone glands of female E. semipurpurella. Field-trapping showed that type 3 receptor neurons mediate strongly antagonistic effects of (R)-heptan-2-ol and (R,Z)-4-hepten-2-ol on E. semipurpurella, while nonan-2-one should possibly be included as a synergist in the sex pheromone blend of this species. The attraction of E. cicatricella and E. sparrmannella to compounds mixed with the pheromone blend of E. semipurpurella shows that the pheromone components of E. semipurpurella have little or no antagonistic effects on these species. The morphology and physiology of eriocraniid pheromone sensilla are very similar to those found in the order Trichoptera (caddisflies), suggesting a homology between pheromone detection systems in the two sister orders Lepidoptera and Trichoptera.

Resistance to a mating disruptant composed of (Z)-11-tetradecenyl acetate in the smaller tea tortrix, Adoxophyes honmai (Yasuda) (Lepidoptera: Tortricidae).

At tea fields in Shimada in Shizuoka Prefecture, a mating disruptant composed of (Z)-11-tetradecenyl acetate (trade name: Hamaki-con) had been used for controlling Adoxophyes honmai (Yasuda) annually since 1983. In 1986, four years after use of the disruptant was initiated, the disruption percentage of pheromone trap catches was 96%, and a high control effect was observed. However, from 1996 to 1998, 14 to 16 years after treatment was started, the percentages became less than 50%. When the disruptant was applied in other tea fields where it had not been previously used, the disruption percentages were over 99%. These results strongly suggest the possibility that a selection pressure caused by continuous use of the disruptant should have caused the low disruption percentage in Shimada. We propose that the phenomenon be termed “resistance” to a mating disruptant composed of (Z)-11-tetradecenyl acetate in A. honmai. When the sex pheromone blend of A. honmai, 63 : 31 : 4 : 2 mixture of (Z)-9-tetradecenyl acetate, (Z)-11-tetradecenyl acetate, (E)-11-tetradecenyl acetate and 10-methyldodecyl acetate (4-component blend), was applied as a disruptant for the “resistance” population, the disruption percentage was 99% and the larval density in the next generation decreased. These results suggest that the 4-component blend is useful as a control tool for the “resistance” population.

Heritable variation of sex pheromone composition and the potential for evolution of resistance to pheromone-based control of the Indian meal moth, Plodia interpunctella.

The short-term evolutionary effect of pheromone-based mating disruption on the mating ability of the Indian meal moth, Plodia interpunctella, was investigated. Three independent selection lines were established, and the mating ability of moths in plastic tents treated with high doses of pheromone and in control tents was compared for two consecutive generations. In addition, the heritability of the sex pheromone blend, measured as the ratio of two major pheromone components (Z,E)-9,12-tetradecadienyl acetate and (Z,E)-9,12-tetradecadienol, was estimated. Based on a mother-daughter regression analysis including 21 families, the heritability of the pheromone blend was 0.65 +/- 0.14, indicating a potential for evolutionary change of the character. However, no increase in mating ability of females in pheromone-treated tents or alteration of the pheromone blend was observed in any selection line when compared with control lines, indicating no or weak selection on the pheromone blend as well as other traits influencing mating ability of this species under the created mating disruption conditions. Factors contributing to the lack of selection effects are discussed.

Transient post–mating inhibition of behavioural and central nervous responses to sex pheromone in an insect

Mating is costly for both male and female insects and should therefore only occur if it is likely to be successful. Within one scotophase, which is the dark period of the light cycle, male moths can only produce one single spermatophore, which is transferred to the female during mating. Remating within the same scotophase would thus be unsuccessful. We tested the hypothesis that newly mated males of the moth Agrotis ipsilon have developed an energy-saving strategy based on the transient inhibition of their sexual behaviour, thus avoiding unsuccessful remating. Agrotis ipsilon males do not copulate more than once during the same scotophase. Moreover, newly mated males do not respond behaviourally to the female sex pheromone although electroantennograms showed that their peripheral olfactory system is fully functional. However, intracellular recordings of antennal lobe neurons showed that the sensitivity for the synthetic sex pheromone blend decreased as compared with that of unmated males. Both the sexual behaviour and the sensitivity of the antennal lobe neurons were restored when tested during the next scotophase. Our results show a fast, transient neuronal plasticity that 'switches off' the olfactory system, which could prevent males from mating unsuccessfully.

Sex Pheromone Composition and Male Trapping of The Oriental Fruit Moth, Grapholita molesta (Lepidoptera: Tortricidae) in Korea

Sex pheromone composition was analyzed for the oriental fruit moth ( Grapholita molesta), and field trapping studies were also conducted to develop monitoring system with its sex pheromone in Korea. The calling and mating behavior of G. molesta were seen maximally 2 to 1 hr before lights-off and these behavior ceased after lights-off. Analysis of abdominal tip extracts from virgin females of G. molesta revealed 4 compounds, Z8-dodecenyl acetate ( Z8-12: Ac), E8-dodecenyl acetate ( E8–12: Ac), Z8-dodecenol ( Z8–12: OH) and dodecanol (12: OH). In the field trappings, the best capturing activity for G. molesta was observed with the 95:5:1 blend of Z8–12: Ac, E8–12: Ac and Z8–12: OH but 12: OH showed no synergic effect. The most effective amount of sex pheromone blend was 0.5 mg for a lure.

Isolation, identification, synthesis, and bioefficacy of female Diacrisia obliqua (Arctiidae) sex pheromone blend. An Indian agricultural pest

Diacrisia obliqua is a polyphagous pest especially on oil seed crops. Adult female sex pheromone blend consists of five pheromone components, which include (3Z,6Z)-cis-9,10-epoxyl,3,6-henicosatriene and (3Z,6Z)-cis-9,10-epoxy3,6-henicosadiene. Synthesis of these enantiomers was achieved through alkylative epoxide rearrangement and stereoselective Wittig olefination reactions as key steps. Bioefficacy experiments both at laboratory and minifield were very positive.

Detection of prey by a spider that aggressively mimics pheromone blends

Adult female bolas spiders have a unique hunting tactic that combines aggressive chemical mimicry of the sex pheromone blends of their prey moths with a specialized weapon (the bolas) and behaviors to capture attracted male moths. This report shows that female bolas spiders can release the attractive allomone before they make the bolas and that females detect moth wing vibrations from attracted prey. In response to this detection, females initiate the construction of a bolas. This ability to sample for prey presence may allow this predator to adapt its hunting activity to the temporal and spatial availability of its prey and, thereby, may reduce the constraints associated with extreme prey specialization.

A chiral sex pheromone system in the pea midge, Contarinia pisi.

The sex pheromone of the pea midge consists of 2-acetoxytridecane, (2S,11S)-diacetoxytridecane and (2S,12S)-diacetoxytridecane. The responses of male pea midges to the corresponding stereoisomers of (2S,11S)-diacetoxytridecane and (2S,12S)-diacetoxytridecane were tested in field trapping experiments and by electroantennographic recordings. When added at 20% of the pheromone component to the sex pheromone blend, the (2S,11R)- and (2R,11S)-stereoisomers of (2S,11S)-diacetoxytridecane, were shown to have a strong inhibitory effect on male attraction in the field. At the same dose, (2R,11R)-diacetoxytridecane, (2R,12R)-diacetoxytridecane, and meso-2,12-diacetoxytridecane, did not have a significant effect on male behavior. It was also shown that substitution of either (2S,11S)-diacetoxytridecane or (2S,12S)-diacetoxytridecane with the related stereoisomers reduced trap catches to the level of blank traps. The electroantennographic recordings showed similar dose-response curves for the pheromone components and the stereoisomers shown to have an inhibitory effect. It seems likely that male antennae have receptors for bothpheromone components and for inhibitory stereoisomers. Scanning electron microscopy and transmission electron microscopy of the antennae revealed three types of sensilla involved in chemoreception: sensilla circumfila, sensilla trichodea, and sensilla coeloconica. The sensilla circumfila and trichodea are both innervated by two sensory cells, whereas the sensilla coeloconica are innervated by four to five cells.

SEX PHEROMONE BLEND OF THE PANDORA MOTH (LEPIDOPTERA: SATURNIIDAE), AN OUTBREAK PEST IN PINE FORESTS (PINACEAE)

AbstractThe female-produced sex pheromone blend of the pandora moth,Coloradia pandora pandoraBlake, a pest responsible for periodic defoliation of pine forests in the western United States, has been identified. The pheromone consisted of a mixture ofE10,Z12-hexadecadienal,E10,Z12-hexadecadienyl acetate, andE10,E12-hexadecadienyl acetate. All three components were essential for optimal attraction.E10,Z12-Hexadecadienol, tentatively identified in trace amounts in pheromone gland extracts, elicited strong responses from male antennae in coupled gas chromatography – electroantennogram analyses but had no apparent effect as a pheromone component at physiologically relevant levels. Hexadecanal, octadecanal, and hexadecyl acetate also were identified in extracts of female pheromone glands but elicited no responses from male antennae, nor did they appear to be components of the attractant blend. In field trials, each trap baited with the optimized three-component blend ofE10,Z12-hexadecadienal,E10,Z12-hexadecadienyl acetate, andE10,E12-hexadecadienyl acetate captured an average of more than 40 male moths per night.

Pheromone Blend Variation and Cross-Attraction Among Populations of the Black Cutworm Moth (Lepidoptera: Noctuidae)

Abstract Sex pheromone quantity, blend ratio, and cross-attraction were studied in four populations of the black cutworm, Agrotis ipsilon (Hufnagel). Analysis of pheromone gland extracts showed that the three pheromone components of A. ipsilon, (Z)-7-dodecenyl acetate (Z7–12:Ac), (Z)-9-tetradecenyl acetate (Z9–14:Ac) and (Z)-11-hexadecenyl acetate (Z11–16:Ac) were present in all four populations. On average, Z7–12:Ac was the most abundant component in the populations from Kentucky and Kansas, whereas in the Egyptian and French populations Z11–16:Ac was the most abundant component. All populations had a similar quantity of Z7–12:Ac. The population from Kentucky had significantly more Z9–14:Ac than the population from Kansas, and the other two populations had intermediate quantities of this component (P < 0.05). The Egyptian population had significantly more Z11–16:Ac than the French population, which had significantly more than the populations from Kentucky and Kansas (P < 0.05). Comparing Nearctic (Kentuc...

Geographical variation in female sex pheromones of the rice leaffolder moth, Cnaphalocrocis medinalis: identification of pheromone components in Japan

AbstractSex pheromone components of the Japanese rice leaffolder moth, Cnaphalocrocis medinalis Guenée (Lepidoptera: Pyralidae) were identified from ovipositor extracts of virgin females as (Z)‐11‐octadecenal, (Z)‐13‐octadecenal, (Z)‐11‐octadecen‐1‐ol and (Z)‐13‐octadecen‐1‐ol at a ratio of 11:100:24:36 by GC‐EAD, GC, GC‐MS. The total amount was estimated to be ca.0.9 ng/female. Field bioassays in Kagoshima, Japan, showed that the two aldehydes are essential for male attraction and the alcohols may have a synergistic effect on the aldehydes. A rubber septum containing 0.9 mg of the four components at the natural ratio was shown to be an effective lure for monitoring this pest in Japan. The above four components are quite different from the sex pheromone components reported previously for the same species of either Philippine or Indian origin; components were shown to be (Z)‐11‐hexadecenyl acetate and (Z)‐13‐octadecenyl acetate at a ratio of 98:2 in the Philippine blend and 1:10 in the Indian blend. Furthermore, in the field tests in Japan, neither the Philippine blend nor the Indian blend showed any attractive activity, while the Japanese blend attracted significant numbers of male moths. These results suggest that there are remarkable geographical variations in the sex pheromone composition of this species or there are several distinct species using different sex pheromone blends.