Journal of Asia-Pacific Entomology 12 (2009) 247–252

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Journal of Asia-Pacific Entomology

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Reidentification of pheromone composition of sulfureana (Clemens) and evidence of geographic variation in male responses from two US states

Junwei Zhu a,⁎, Sridhar Polavarapu b, Kye-Chung Park c, Carolyn Garvey d, Daniel Mahr d, Satoshi Nojima e, Wendell Roelofs e, Tom Baker f a USDA-ARS-NPA, Agroecosystem Management Research Unit, Lincoln, NE 68583, USA b Marucci Center for Blueberry and Cranberry Research and Extension, Chatsworth, NJ 08019, USA c Plant and Food Research, Canterbury Research Center, Gerald Street, P.O. Box 51, Lincoln 8152, New Zealand d Department of Entomology, University of Wisconsin, Madison, WI 53706, USA e Department of Entomology, Barton Laboratory, Cornell University-New York State Agricultural Experiment Station, Geneva, NY 14456, USA f Department of Entomology, Pennsylvania State University, University Park, PA 16802, USA article info abstract

Article history: GC-EAD analyses of pheromone gland extracts of calling female revealed at least 6 Received 10 March 2009 compounds that consistently elicited antennal responses from male antennae. In addition to the major Revised 21 April 2009 pheromone compound, (E)-11-tetradecenyl acetate (E11–14:OAc), which was previously reported, the other Accepted 26 April 2009 compounds were found to be (E)-9-dodecenyl acetate (E9–12:OAc), (Z)-9-dodecenyl acetate (Z9–12:OAc), (Z)-9-tetradecenyl acetate (Z9–14:OAc), (Z)-11-tetradecenyl acetate (Z11–14:OAc), and (E)-11-tetradecenol Keywords: (E11–14:OH). Tetradecyl acetate, hexadecyl acetate and hexadecenyl acetates were also present in the extracts, Sex pheromone Sparganothis sulfureana but elicited no EAG response from male antennae. Wind tunnel tests demonstrated that males from Cranberries responded equally well to a blend containing five pheromone components in relative to the pheromone glands Pheromone polymorphism of calling females. Different male-response profiles from field-trapping tests conducted in the states of (E)-11-tetradecenyl acetate Wisconsin and New Jersey were observed, respectively. Significantly higher numbers of male S. sulfureana were (Z)-11-tetradecenyl acetate caught in New Jersey in traps baited with the binary blend of E11–14:OAc (30 μg) with 1% of Z11–14:OAc, but (E)-9-dodecenyl acetate males from Wisconsin responded equally well to traps containing blends of E11–14:OAc with 0–10% of Z11–14: (E)-11-tetradecenol OAc. The addition of more than 10% of Z11–14:OAc to the primary pheromone compound reduced male (Z)-9-tetradecenyl acetate captures significantly in both states. Male catches were doubled by adding E9–12:OAc and E11–14:OH to the most attractive binary blend in both states. The trapping test with caged live virgin female showed that males in Wisconsin preferred females from the local population than those from New Jersey. The differences in male responses observed may indicate the existence of pheromone polymorphism in this species. © Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society, 2009 Published by Elsevier B.V. All rights reserved.

Introduction begins in early June with a flight period of 4–5 weeks, and the second generation emerges in the first week of August, and continues for 8– The cranberry, macrocarpon, is a native North American 9 weeks (Cockfield et al., 1994). The second generation during late crop that is commercially grown on thousands of hectares in the summer has been considered as the most destructive, when the larvae northeast, north central and northwest regions of the United States. feed on both fruits and foliage. The Sparganothis fruitworm, Sparganothis sulfureana (Clemens), is Control of S. sulfureana in cranberry marshes relies heavily on the one of the three most important pests of cranberries in Massachusetts, application of organophosphate and carbamate insecticides. However, Wisconsin, and New Jersey, which three states together account for with increasing concern about potential environmental risks such as almost 90% of the cranberry production in United States (Cross, 2003). surface water pollution, and the elimination of organophosphate In New Jersey, S. sulfureana is regarded as the most important insecticides due to the 1996 Food Quality Protection Act, makes it cranberry pest during the post-pollination period and is the primary necessary to develop alternative pest management tools. Pheromone target for most insecticide applications. S. sulfureana has two mating disruption is one of these alternatives that has been used against generations per season (Averill and Sylvia, 1998). The first generation this pest with partial pheromone compounds, such as microencapsu- lated (E)-11-tetradecenyl acetate (E11–14:OAc) (Polavarapu et al., ⁎ Corresponding author. Fax: +1 402 437 5260. 2001). However, formulated natural blend of sex pheromone is argued E-mail address: [email protected] (J. Zhu). to provide most robust mating disruption (Cardé and Minks, 1995).

1226-8615/$ – see front matter © Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society, 2009 Published by Elsevier B.V. All rights reserved. doi:10.1016/j.aspen.2009.04.005 248 J. Zhu et al. / Journal of Asia-Pacific Entomology 12 (2009) 247–252

Field screening tests with several aliphatic alcohols and acetates antenna was used for EAG recordings. A capillary glass Ag–AgCl have identified (E)-11-tetradecenyl alcohol (E11–14:OH) (Roelofs and recording electrode filled with electrode saline (0.1 M KCl) was placed Comeau, 1970) and E11–14:OAc (Comeau and Roelofs, 1973; Mayer in contact with the cut tip of the antenna. Another electrode filled and McLaughlin, 1991) as sex attractants for S. sulfureana. Recently it with the same saline was connected with the base of the antenna and was found by two of us (Roelofs and Nojima, unpublished) that E11– served as the reference electrode. The EAG setup with a high-input DC 14:OAc was present in female pheromone gland extracts. Additional with automatic baseline drift compensation that was used in this field-trapping tests indicated that the addition of two other com- study was from Syntech (Hilversum, The Netherlands). A GC-EAD pounds, (Z)-9-tetradecenyl acetate (Z9–14:OAc) and (E)-9-dodecenyl program (version 2.3) developed by Syntech was used to record and acetate (Z9–12:OAc), increased male captures by 2- to 10-fold at apple analyze the amplified EAD and FID signals on a Pentium III PC. orchards in New York State (Hill and Roelofs, unpublished). However, GC-MS analyses were performed by using a Hewlett Packard 5890 there are no reports to date that determine sex pheromone constituents Series II gas chromatograph interfaced to a Hewlett Packard 5972 of S. sulfureana based on sex pheromone gland extraction or effluvia Mass Selective Detector (MSD), equipped with the same columns collections. Identification of the full natural blend of sex pheromone of described in GC-EAD analyses. Concentrated pheromone gland S. sulfureana will provide further insights into pheromone-mediated extracts were injected for analyses. The GC operation conditions sexual communication, and may further optimize mating disruption were same as those described above. Mass spectra were recorded from technology for this cranberry pest. 30 to 550 a.m.u. after electronic impact ionization at 70 eV. The chemical In the present study, we report the results from gas chromato- identification of pheromone compounds was performed by comparing graphic-electroantennogram (GC-EAD) analyses of pheromone gland the retention indices and mass spectra with those of authentic extracts from calling female S. sulfureana that reveal six pheromone standards in a mass spectral library (Wiley 138K, John Wiley & Sons, candidate compounds eliciting antennal responses from male anten- Inc., New York, USA). nae. Further, we report pheromone structure identification and male All synthetic saturated and mono-unsaturated aliphatic acetates, behavioral responses to pheromone blends containing minor pher- aldehydes and alcohols (C12–C16) tested in the present study were omone components in laboratory wind tunnel studies and in purchased from the PheroBank (The Netherlands) and the purity was cranberry fields. Different profiles of male responses in different N99% according to analyses by GC-MS. The two groups of conjugated field test sites have also been observed, which may suggest acetates (7,9-dodecadienyl acetates and 9,11-tetradecadienyl acetates) pheromone polymorphism in this species. were generously provided by Dr. Christer Löfstedt (Pheromone Group, Lund University, Sweden) with purities described in Zhu et al. (1996). Methods and materials Behavioral assays in wind tunnel Triangular pieces of filter paper (Whatman white filter paper, Male and female moths of S. sulfureana used for the present 0.5 cm2) were used as dispensers for all behavioral assays in the wind study were emerged from pupae shipped from Marucci Center for tunnel. The filter paper lure was attached to an pin, which was Blueberry and Cranberry Research and Extension, Rutgers University, provided with a rubber septum support base and set on a wire stand Chatsworth, New Jersey. The male pupae were maintained at 20 °C, approximately 40 cm above the wind tunnel floor. The filter paper was 55% relative humidity in a climate-control chamber with a 14:10 L:D impregnated with pheromone solution 2 min before testing, and cycle, whereas female pupae were set in a separate chamber at 23 °C replaced every 20 min. All synthetic lures tested in the wind tunnel under a reversed photoperiod. Emerged adults were held in paper contained approximately 100 ng (in a volume of ∼10 μl) of the major cartons (3.75 L), and supplied with a 5% sucrose solution until used in the experiments. Table 1 Preparation of gland extracts Comparisons of retention time between the synthetic standards and pheromone candidates from gland extracts of New Jersey female Sparganothis sulfureana on two different columns in GC-MS analyses. Female S. sulfureana were observed to have a period of maximum calling between 1 to 2 h into the scotophase. Pheromone glands of 2– Compounds Carbowax DB-225 3 day old females were dissected at their peak calling time, and Synthetics Extracts Synthetics Extracts extracted in hexane (B&J high grade solvent) for 30 min. Batch E9–12:OAc 12.12 12.12 11.94 11.93 Z9–12:OAc 12.24 12.24 12.10 12.09 extracts were pooled from 10–30 glands in 30 μl of solvent with 10 ng Z9–14:Ald 12.54 – 12.64 – of internal standard (pentadecane). These batched extracts were then E11–14:Ald 12.58 12.57 12.70 – analyzed with gas chromatograph-mass spectrometry (GC-MS). The Z7,E9–12:OAc 13.63 – 13.22 13.21 solvent extracts without the addition of the internal standard were E7,Z9–12:OAc 13.75 – 13.31 – – ⁎ – saved for use in behavioral wind tunnel bioassays. Z7,Z9 12:OAc 13.83 13.85 13.37 E7,E9–12:OAc 13.95 – 13.37 13.36 14:OAc 13.84 13.85 13.72 13.72 Chemical analysis E11–14:OAc 14.30 14.25–14.36 14.05 14.06 Z9–14:OAc 14.26 14.25–14.36 13.99 14.00 A Hewlett Packard 5890 Series II gas chromatography was equipped Z11–14:OAc 14.44 14.45 14.15 14.16 Z7–14:OH 14.87 – 13.63 13.64 either with a DB-225 column or a carbowax column (30 m×0.25 mm E11–14:OH 15.07 15.08 13.63 13.64 fi fl i.d., J & W Scienti c, Folsom, CA). An ef uent split at 1:1 ratio allowed Z11–14:OH 15.21 15.21 13.80 13.80 simultaneous flame ionization detector (FID) and male electroanten- Z9,E11–14:OAc 15.69 15.68 15.19 – nogram (EAD) responses to female pheromone gland extracts. The E9,Z11–14:OAc 15.80 – 15.26 15.25 – – extracts were injected in a splitless mode. The injector temperature Z9,Z11 14:OAc 15.89 15.86 15.33 E9,E11–14:OAc 16.00 – 15.33 – was set at 250 °C, and the split valve was opened 1 min after injection. 16:OAc 15.88 15.86 15.46 15.47 The initial column temperature was 80 °C for 1 min and then linearly Z9–16:OAc 16.17 16.14 15.81 15.79 increased to 230 °C at a rate of 15 °C/min. The outlet for the EAG was Z11–16:OAc 16.29 16.28 15.91 15.90 fi fl continuously supplied with a puri ed air stream owing over the ⁎ Retention time in italics of the two adjacent compounds means that these two antennal preparation at a speed of 0.5 m/s. A dissected male compounds co-elute from the analyzing column used for analyses. J. Zhu et al. / Journal of Asia-Pacific Entomology 12 (2009) 247–252 249

Fig. 1. GC-EAD analyses of male antennae of Sparganothis sulfureana to female pheromone gland extracts (2 female equivalents) on a DB-225 column (GC temperature program rate at 15 °C/min). Insert shows male antennae responses to synthetic pheromone standards on the same column, but with a 10 °C/min programmed rate. Compound1:E9–12:OAc, 2: Z9–12:OAc, 3: E11–14:OH, 4: Z9–14:OAc, 5: E11–14:OAc, 6: Z11–14:OAc. pheromone component (E11–14:OAc), and other compounds were #1780-J07) were used as dispensers. Approximately 30 μg of synthetic added in relative amounts based on the GC-MS analyses of calling pheromone blend compounds (in 50 μl of hexane) was impregnated female gland extracts (see details in Table 2). Hexane (10 μl) was used into each rubber septum. Wing traps (Great Lakes IPM, Vestaburg, as control. ) were used for tests in both states. Traps were placed in the A 1.5×1.5×2 m Plexiglas wind tunnel was used for the bioassays cranberry beds at ca. 20 cm above the canopy. Treatments were tested with New Jersey male moths. The temperature inside the wind tunnel in randomized complete-block design with blocks placed in separate was maintained at 22±2 °C during the experiment, and the wind cranberry beds. Within a replicate (block) the traps were set at least velocity was 0.5 m/s, humidity was 40–50%, and the light intensity 10 m apart in Wisconsin fields and 22–25 m in New Jersey. Traps were was around 5 lux. Males of S. sulfureana aged 2–3 days were checked every 3–4 days, and trap positions within a series were re- transferred to glass tubes (8×2.5 cm) and placed in the wind tunnel randomized after checking, to minimize the effects of habitat room during the last 2 h of the photophase. Male moths were tested heterogeneities. In the first field-trapping test, effects of different 0–2 h after the onset of the scotophase. Individual males were ratios of E11–14:OAc and Z11–14:OAc on male attraction were released from the glass tube at 1.5 m downwind from the tested conducted in June, 2001. The second set of experiments was then pheromone source. Each male was allowed 1 min to initiate flight after carried out later during July to determine the attractiveness of blends being introduced into the pheromone plume. Once flight was with addition of secondary compounds that were identified from the initiated, males were observed for 2 min or until they stopped female pheromone gland extracts. The third trapping test with caged responding. A number of behavioral characteristics were scored, including take-off, oriented flight in the plume, halfway to the source, and finally landing on the source. Due to the limited number of male Table 2 Amounts of pheromone-gland constituents of New Jersey female Sparganothis moths available for the wind tunnel test, only selected blends were sulfureana and their relative ratios to the major pheromone component, E11–14:OAc. tested. In those tests, we attempted to test all treatments during the same day's wind tunnel sessions. Compounds Titers (ng/FE) Relative ratios E9–12:OAc 0.012±0.003 0.053±0.03 – Field-trapping tests Z9 12:OAc 0.001±0.003 0.004±0.001 Z9–14:OAc 0.26±0.05 1.15±0.06 E11–14:OAc 22.71±0.03 100.00 Trapping tests to evaluate the attractiveness of different combina- Z11–14:OAc 0.35±0.05 1.54±0.10 tions of the pheromone component candidates were carried out in E11–14:OH 1.27±0.25 5.59±1.77 fi cranberry elds in New Jersey and Wisconsin. Synthetic blends were The results of this quantification were based on analyses of more than 10 batches of prepared in hexane, and red rubber septa (Arthur H. Thomas Co. extracts with a total of pheromone glands dissected from 194 calling females. 250 J. Zhu et al. / Journal of Asia-Pacific Entomology 12 (2009) 247–252 live virgin female moths from Wisconsin (field collected pupae) and Table 3 New Jersey (laboratory pupae from a colony that was refreshed every Number of male Sparganothis sulfureana caught in traps baited with different ratios of E11- and Z11–14:OAc in Wisconsin and New Jersey cranberry fields (2001). year with field collected adults) was conducted in cranberry marshes in Wisconsin. Three female moths from each state were placed inside % of pheromone candidates Mean number of moths caught per trap a metal mesh cage, and hung from the top of a wing trap. The (30 μg dosage) (±std. error) experiment was run from the end of August to mid September, 2001. E11–14:OAc Z11–14:OAc Wisconsin (6/15–7/10) New Jersey (6/21–6/29) Female moths were refreshed every week. Trap catches were checked 100 0 10±2 ab 39±4 b every 4th day, and trap positions were randomized within a series of 100 1 10±2 ab 52±4 a 97 3 11± 4 a 17 ± 3 c five replicates. 90 10 6±1 ab 4±1 d 70 30 3±1 b 3±1 d Results Means followed by different letters indicate that there are significant differences among treatments (ANOVA followed by Fisher's PLSD test, Pb0.05, n=6). GC-EAG analyses and chemical identification

By comparing retention times with those of synthetic pheromone Male behavioral responses in the wind tunnel compounds commonly found in tortricid moths (including saturated, mono-unsaturated and conjugated acetates, aldehydes and alcohols Responses of male S. sulfureana from New Jersey to a five-component from carbon chain length of C12 to C16), we tentatively identified a synthetic blend containing E11–14:OAc, Z11–14:OAc, E9–12:OAc, Z9– total of 19 compounds from pheromone gland extracts in GC-MS 14:OAc and E11–14:OH were equivalent to female extracts (Fig. 2). analyses (Table 1). Among them, E9–12:OAc, Z9–12:OAc, 14:OAc, Z9– However, the overall rates of successful landing were only 30%. The 14:OAc, E11–14:OAc, Z11–14:OAc, E11–14:OH, Z11–14:OH, 16:OAc, major pheromone component, E11–14:OAc, alone was not attractive to Z9–16:OAc and Z11–16:OAc, were the compounds found in pher- males in these tests, and did not even evoke oriented flight in the plume omone extracts analyzed from both columns. GC-EAD analyses in these tests. Sequential addition of several candidate minor pher- showed that 6 compounds elicited antennal responses more con- omone components, starting with Z11–14:OAc, then E9–12:OAc, Z9–12: sistently than others (Fig. 1). Further GC-MS analyses of the extract OAc and Z9–14:OAc, to the major pheromone component did not from the same batch analyzed in GC-EAD confirmed their chemical increase male responses. However, a 5-component blend that included structure identifications based on their retention times and char- E11–14:OH, increased male response significantly to the levels not acteristic fragments of mass spectra compared to those from authentic differing from those occurring in response to female extract (Fig. 2). No standards. Two alcohols (Z7–14:OH and E11–14:OH), which were males were observed to respond to the control (either stayed inside the inseparable on the DB-225 column, were resolved by analysis on a releasing cage, or randomly flew towards the wind tunnel ceiling). Carbowax column to confirm the identification of E11–14:OH (Table 1). Although the retention time of one peak matched that of Field-trapping tests synthetic Z11–14:OH, it was excluded due to the ambiguity of the mass spectrum. We also excluded conjugated dodecadienyl and In the first field-trapping experiment, different ratios of the binary tetradecadienyl acetates as candidate pheromone compounds because blend of E11–14:OAc and Z11–14:OAc were tested in both New Jersey they were not consistently present from analyses on columns with and Wisconsin. In New Jersey, traps baited with E11–14:OAc and 1% of different polarities. Further GC-EAD analyses of a synthetic standard Z11–14:OAc caught the highest number of male S. sulfureana followed mixture confirmed the identification of above six EAD-active by the blend containing the primary pheromone component, E11–14: pheromone compounds from conspecific male antennae (Fig. 1 OAc, alone (Table 3). Further increases in the amount of Z11–14:OAc insert). The titers and their relative ratios of those pheromone above 1% in the binary blend lowered trap catches significantly. compounds are summarized in Table 2. However, in Wisconsin, no significant differences in male captures

Fig. 2. Behavioral responses of male Sparganothis sulfureana in a subtractive assay in a wind tunnel. Data points with the same letters do not differ significantly (PN0.05, Ryan's test). The relative ratios of the tested pheromone blends are based on the GC-MS results shown in Table 2. J. Zhu et al. / Journal of Asia-Pacific Entomology 12 (2009) 247–252 251

Table 4 Number of male moths caught in traps baited with different ratios of pheromone candidates of Sparganothis sulfureana in Wisconsin and New Jersey cranberry fields (July of 2001).

% of pheromone candidate components Mean number of moths caught/ (30 μg total dosage) trap (±std. error) E11–14:OAc Z11–14:OAc E9–12:OAc Z9–12:OAc Z9–14:OAc E11–14:OH Wisconsin New Jersey 100 1 6±1 b 16±3 b 100 1 0.05 4±1 b 20±5 ab 100 1 1 10±2 ab 15±4b 100 1 66±1b23±3a 100 1 0.05 1 12±2 ab 15±4 b 100 1 0.004 1 8±1 b 15±3 b 100 1 0.05 0.004 1 6 15±3 a 29±6 a

Means followed by different letters indicate that there are significant differences among treatments (ANOVA followed by Fisher's PLSD test, Pb0.05, n=7). were observed with traps baited with blends containing E11–14:OAc E9–12:OAc, Z9–12:OAc, Z9–14:OAc, and E11–14:OH found from and from 0 to 10% of Z11–14:OAc. In both states, the addition of 30% of pheromone gland extracts, were identified as potential pheromone Z11–14:OAc to the primary pheromone compound significantly components. Among these constituents, Z9–14:OAc and E11–14:OH reduced male catches. appeared to contribute to attraction of males in Wisconsin and New Further field tests showed that in New Jersey, traps baited with a 6- Jersey, respectively. Field studies in New Jersey also indicated the compound blend or the binary blends of E11–14:OAc plus 1% Z11–14: importance of E9–12:OAc in sex pheromone communication. OAc with either E9–12:OAc or E11–14:OH added to it caught the Although field-trapping tests in both states showed that the 6- highest number of males (Table 4). No differences in trap catches were compound blend attracted significantly more males than the binary found among the binary blend with other secondary compounds. blend, exactly which of these additional minor gland constituents Although the 6-compound blend was also the most attractive blend to designated as being sex pheromone components remains to be males in Wisconsin, the male response pattern observed in the field determined. differed from that in New Jersey. No increases in male catches were Many of the pheromone-gland constituents reported from female observed in Wisconsin when the binary blend was added with either S. sulfureana in the present study were also identified as being either E11–14:OH or E9–12:OH. However, the same combinations tested in pheromone components or sex attractants in other species within the New Jersey resulted in significant increases in male captures. genus Sparganothis (Roelofs and Brown, 1982). E11–14:OAc was The third trapping test using caged live virgin female moths from reported to be attractive to the males of S. albicaudana and S. tunicana Wisconsin and New Jersey conducted in Wisconsin cranberry fields (Roelofs and Comeau, 1971; Daterman et al., 1977). Females of S. clearly showed that the male moths from Wisconsin were preferen- directana, S. pilleriana and Sparganothis sp. were shown to produce a tially attracted to traps containing Wisconsin females (Fig. 3). Females binary pheromone blend containing E11–14:OAc and Z11–14:OAc from New Jersey attracted significantly fewer Wisconsin males. No (Bjostad et al., 1980a,b; Guerin et al., 1986). E11–14:OH is a major males were caught in control traps baited with hexane only pheromone component of the tufted apple bud moth, Platynota throughout our field tests. idaeusalis, and of the spotted fireworm, parallela (Hill et al., 1974; Polavarapu and Lonergan, 1998). This compound is also a Discussion pheromone component of the woodbine leafroller, Sparganothis sp. (Daterman et al., 1977; Bjostad et al., 1980a; Polavarapu and Lonergan, Among the six EAG-active compounds identified from female 1998). gland extracts of S. sulfureana,bothE11–14:OAc and Z11–14:OAc (in a Although we found that retention times of several other 1% blend with E11–14:OAc) exhibited significant effects on male compounds from female gland extracts matched those of mono- attraction in field tests in New Jersey, and must now be considered unsaturated acetates, alcohols and aldehydes with carbon chain pheromone components. In the present study, four other compounds, lengths with C12–C14, their chemical structures were not confirmed due to either mismatched mass spectra of synthetic standards or absence in gas chromatograph analyses on two different columns. The presence of traces of two groups of conjugated acetates (7,9–12:OAc and 9,11–14:OAc) could not be confirmed by mass spectrometric data due to only partial matches observed. Although 14:OAc, 16:OAc, Z9– 16:OAc and Z11–16:OAc were tentatively identified, they were not consistently found in each analyses, nor elicited any EAD responses, therefore, they were not included in the subsequent behavioral tests. Various ratios of the binary blend of E11–14:OAc and Z11–14:OAc have been considered as the most common pheromone components in many species within the (Roelofs and Brown, 1982). Results from our field tests demonstrated that a blend with 100:1 of E11–14:OAc and Z11–14:OAc was very attractive to male S. sulfureana in both New Jersey and Wisconsin. Subsequent field tests using the most active binary blend with the addition of other compounds revealed apparent differences in blends that increased attraction of males in the two states. The addition of either E11–14:OH or E9–12: OAc to the binary blend appeared to be most important for increasing Fig. 3. Trapping male Sparganothis sulfureana in Wisconsin cranberry fields with caged male catches in New Jersey, whereas Z9–14:OAc appeared to be an virgin females collected from New Jersey and Wisconsin (5 replicates, Aug.–Sept., important minor constituent in Wisconsin. Furthermore, in a field test 2001). Error bars indicate standard error. Columns labeled with different letters fi indicate significant differences (Pb0.05, Student T-test). 10 cm: 10 cm away from the conducted in Wisconsin, signi cant more males were caught in traps tested attractant source. baited with caged virgin females collected from Wisconsin than in 252 J. Zhu et al. / Journal of Asia-Pacific Entomology 12 (2009) 247–252 those baited with caged New Jersey females, suggested that a possible Cockfield, S.D., Butkewitch, S.L., Samoil, K.S., Mahr, D.L., 1994. Forecasting the flight activity of Sparganothis sulfureana (: Tortricidae) in cranberries. J. Econ. pheromone polymorphism in S. sulfureana. Further work is needed, Entomol. 87, 193–196. however, because although care was taken to equilibrate New Jersey Comeau, A., Roelofs, W.L., 1973. Sex attraction specificity in the Tortricidae. Entomol. females to their new surroundings, the stress of shipping the pupae to Exp. Appl. 16, 191–200. Cross, B., 2003. 2002 Cranberry Statistics. New Jersey Agricultural Statistics Service, Wisconsin could have reduced female attraction due to lower New Jersey Department Agriculture, Trenton, New Jersey. emission rates or frequency of calling posture (gland extrusion), Daterman, G.E., Robbins, R.G., Eichlin, T.N., Pierce, J., 1977. Forest Lepidoptera attracted effects that were not at all related to geographically polymorphic by known sex attractants of western spruce budworms, Choristoneura spp. – blend ratio differences. (Lepidoptera: Tortricidae). Can. Entomol. 109, 875 878. Guerin, P.M., Buser, H.R., Tόth, M., Höbaus, E., Schmid, A., Arn, H., 1986. Sex pheromone The present study has shown that females of S. sulfureana produce of : E- and Z-11-tetradecenyl acetates as essential compo- the primary component, E11–14:OAc along with several other nents. Entomol. Exp. Appl. 40, 137–140. pheromone compounds. A blend of six compounds was highly Hill, A., Carde, R., Comeau, A., Bode, W., Roelofs, W., 1974. Sex pheromone of the tufted apple bud moth (Platynota idaeusalis). Environ. Entomol. 3, 249–252. attractive to male moths in New Jersey. Although traps baited with Mayer, M.S., Mc L aughlin, J.R., 1991. Handbook of Insect Pheromones and Sex this pheromone blend captured the highest number of males in Attractants. CRC Press, Boca Raton, . Wisconsin, the identification of the minor gland constituents Polavarapu, S., Lonergan, G., 1998. Sex pheromone of Choristoneura parallela (Lepidoptera: Tortricidae): components and development of a pheromone lure responsible for S. sulfureana pheromone communication in Wisconsin for population monitoring. Environ. Entomol. 27, 1242–1249. has to be performed. Polavarapu, S., Lonergan, G.C., Peng, H., Neilsen, K., 2001. Potential for mating disruption of Sparganothis sulfureana Clemens (Lepidoptera: Tortricidae) in cranberries. J. Econ. Entomol. 94, 658–665. References Roelofs, W.L., Brown, R.L., 1982. Pheromones and evolutionary relationships of Tortricidae. Ann. Rev. Ecol. Syst. 13, 395–422. Averill, A.L., Sylvia, M.M., 1998. Cranberry Insects of the Northeast. University of Roelofs, W.L., Comeau, A., 1970. Lepidopterous sex attractants discovered by field Massachusetts Extension, Amherst, MA. 112pp. screening tests. J. Econ. Entomol. 63, 969–973. Bjostad, L.B., Taschenberg, E.F., Roelofs, W.L., 1980a. Sex pheromone of the woodbine Roelofs, W.L., Comeau, A., 1971. Sex attractants in Lepidoptera. In: Tahori, A.S. (Ed.), leafroller moth, Sparganothis spp. J. Chem. Ecol. 6, 797–804. Chemical Releasers in Insects. InGordon & Breach, NY, pp. 91–114. Bjostad, L.B., Taschenberg, E.F., Roelofs, W.L., 1980b. Sex pheromone of the chokecherry Zhu, J., Ryrholm, N., Ljungberg, H., Hansson, B.S., Hall, D., Reed, D., Löfstedt, C., 1996. leafroller moth, Sparganothis directana. J. Chem. Ecol. 6, 487–498. Olefinic acetates, Δ-9,11–14:OAc and Δ-7,9–12:OAc used as sex pheromone Cardé, R.T., Minks, A.K., 1995. Control of moth pests by mating disruption: successes and components in three geometrid moths, Idaea aversata, I. straminata and I. biselata constraints. Ann. Rev. of Entomol. 40, 559–585. (Geometridae, Lepidoptera). J. Chem. Ecol. 22, 1505–1526.