Mating Disruption of Paralobesia viteana in Vineyards Using Pheromone Deployed in SPLAT-GBM Wax Droplets

J Chem Eol (28) 34:189 195 DOI 1.17/s1886-8-9497- Mating Disruption of Paralobesia viteana in Vineyards Using Pheromone Deployed in SPLAT-GBM Wax Droplets Paul E. Jenkins & Rufus Isaas Reeived: 5 Otober 27 /Revised: 1 Deember 27 /Aepted: 9 April 28 /Published online: 26 June 28 # Springer Siene + Business Media, LLC 28 Abstrat A paraffin wax formulation releasing pheromone for mating disruption of insets was tested during 25 and 26 in Vitis labrusa vineyards infested by grape berry moth, Paralobesia viteana (Lepidoptera: Tortriidae). In early May of eah year, 1-ml droplets of SPLAT-GBM wax ontaining 3% sex pheromone of P. viteana were applied to every wooden post at a rate of 4/ha in repliated 1.3-ha plots. Moth aptures in sex pheromone baited traps plaed at the vineyard borders and interiors revealed signifiant disruption of male moth aptures in treated plots, with ativity of one appliation lasting over 1 weeks during both years. Treatment with SPLAT- GBM did not affet the proportion of lusters infested until the end of the seond growing season, when infestation was 27% lower in the treated plots than in the ontrol plots. Comparisons of moth aptures in traps plaed inside 15.2 16.5 m vine plots that were untreated or reeived varying densities of.2-ml wax drops or Isomate- GBM hand-tied dispensers at the reommended rate of 45/ha indiated that orientational disruption inreased with droplet density. Similar numbers of moths were aptured in plots that reeived 1 or 3 drops per vine as were trapped in plots with twist ties spaed at.4 per vine. Moth aptures in monitoring traps baited with inreasing sizes of wax droplets (.2,.5, or 1-ml drops) or red septa ontaining P. viteana sex pheromone suggest dereasing ability of male moths to reah traps with inreasing pheromone loading. This study indiates that wax-deployed pheromone an redue rop infestation by P. viteana after 2 years of deployment, and that the inreasing of pheromone release P. E. Jenkins (*) : R. Isaas Department of Entomology, Mihigan State University, East Lansing, MI 48824, USA e-mail: jenki132@msu.edu by using appliation of greater droplet densities or by using larger droplets will improve the level of disruption ahieved. Keywords Grape berry moth. IPM. Vineyard. Endopiza viteana. Lepidoptera. Tortriidae Introdution Vineyards in the eastern United States are at risk of infestation by the grape berry moth, Paralobesia viteana (Clemens), a tortriid moth that lays eggs diretly onto lusters, and whose larvae then bore into the fruit (Hoffman and Dennehy 1989; Tobin et al. 23). This inset is the most eonomially important diret pest of eastern US and Canadian vineyards, and protetion of fruit is ahieved typially by using broad-spetrum insetiides. Effetive alternative ontrol strategies are needed for the ontrol of P. viteana to enhane adoption of IPM programs and to help minimize the risk of resistane to insetiides (Jenkins and Isaas 27a, b). The use of sex pheromones for mating disruption of P. viteana is one approah that has been tested in researh trials but whih has relatively limited ommerial adoption. The major omponent of the sex pheromone of P. viteana was identified by Roelofs et al. (1971) as(z)-9 dodeenyl aetate with the minor omponent, (Z)-11 tetradeenyl aetate, being identified by Witzgall et al. (2). These disoveries led to evaluations of mating disruption using hand-applied rope dispensers (subsequently referred to as twist ties) and sprayable formulations of this pheromone, that provided promising results in New York and Ontario vineyards (Dennehy et al. 199; Trimble 1993; Trimble et al. 1991, 23). Despite these findings, adoption

19 J Chem Eol (28) 34:189 195 of hand-applied rope dispensers for season-long ontrol of this pest by grape growers has been low, and no ompanies urrently produe sprayable pheromones for P. viteana ontrol. Among the reasons for the low adoption of this tehnology is the 2- to 3-week duration of pheromone release by sprayable formulations (R. Isaas, unpublished data) and the labor required to apply rope dispensers to the rop. Reently, a paraffin wax emulsion has been developed, into whih inset sex pheromones an be mixed at the required onentration (Atterholt 1996; Atterholt et al. 1998; de Lame 23). This wax is a visous homogenate that hardens one applied to rop foliage or branhes and it an at as a long-lasting disrete soure of pheromone emission. The wax an be applied into rop systems by using manual methods that are more rapid than manual appliation of twist ties (de Lame 23; Stelinski et al. 25; Epstein et al. 26), or it an be applied by mehanial methods (Stelinski et al. 25) that enable appliation to large orhards or vineyards from motorized vehiles (Stelinski et al. 26). The optimization of mating disruption requires that the density and size of droplets, as well as the pheromone release rate and duration, are appropriate for the biology of the targeted pest (Miller et al. 26a, b). Beause the size, density, and distribution of wax droplets an be easily manipulated, this delivery system also provides researhers with a flexible tool to aid investigations of how moths an be optimally disrupted. For example, Stelinski et al. (25) doumented inreased mating disruption of male oriental fruit moth, Grapholitha molesta, in apple orhards as droplet density inreased, thus leading to inreased suppression of mating in virgin female moths. The authors ompared mating disruption in plots treated with wax droplets or the standard twist-tie dispenser, and found superior performane when wax was used. This suggests that the typial deployment density of pheromone twist ties may not provide suffiient point soures of pheromone for optimal disruption of G. molesta. More reent studies that evaluated mating disruption of the odling moth, Cydia pomonella, in apple orhards treated with the odlemone sex pheromone in wax droplets provide further evidene that higher densities of pheromone release soures improve disruption of mating (Epstein et al. 26). The suess of these studies and others in otton, pear, and walnut rops has led to the development of a ommerial produt alled SPLAT (Speialized Pheromone and Lure Appliation Tehnology) that ats as the arrier for sex pheromones or other semiohemials. To asertain the potential for wax-based formulations for mating disruption of P. viteana, we determined the duration and effiay of pheromone disruption of P. viteana in Mihigan vineyards over two growing seasons by using the GBM-SPLAT ommerial formulation of the wax matrix. We also tested the hypotheses that orientation of P. viteana to droplets would derease with inreasing pheromone release, and that mating disruption would be improved by inreasing the density of droplets. Methods and Materials Study Sites All studies were onduted in mature Vitis labrusa var. Conord vineyards in Van Buren Co., Mihigan. These vineyards reeived standard fungiide and insetiide programs applied by the growers, but low rop levels due to a late spring freeze in 26 led to a redued pestiide program and higher ativity of P. viteana in the seond year. Large-sale Mating Disruption Study This study was onduted during 25 and 26 in four vineyards at the same farm, eah of whih was split into two equal-sized plots [a. 1.3 ha (1 13 m)]. The plots were assigned to be treated with pheromone in wax, or not treated, in a randomized omplete blok design with four repliates. Plant spaing in these plots was 2.7 m between vines and 3 m between rows, with posts distributed every three vines, resulting in 12 vines and 4 posts per hetare. In both years, 1-ml droplets of GBM-SPLAT (ISCA Tehnologies, In., Riverside, CA, USA) were deployed by hand on the north side of eah wooden post in the treated plots with a 3-ml syringe. The appliations were made on 9 May in 25 and 1 May in 26, with the same plots treated in both years with wax ontaining 3% (v/v) of P. viteana pheromone (12 ml AI/ha) at a 1:1 ratio of (Z)-9 dodeenyl aetate: (Z)-11 tetradeenyl aetate (Shin-Etsu Chemial Co., Tokyo, Japan). In 25, the formulation also ontained 3% (v/v) ypermethrin, but no moths were trapped near these droplets when plaed on stiky traps, suggesting that P. viteana moths did not ontat the droplets (R. Isaas, unpublished data). Adult male P. viteana were monitored in eah plot with large plasti delta traps (Suterra LLC, Bend, OR, USA) baited with rubber septa that ontained female sex pheromone (9:1 ratio of (Z)-9 12A and (Z)-11 14A) (Suterra LLC, Bend, OR). Two traps were suspended from the top trellis wire at a height of 1.5 m at both the border and the interior of eah vineyard, with at least 33 m between traps. Those in the interior were 65 m from the border, in the same rows as the border traps. The number of male P. viteana aptured was monitored weekly, and moths were removed or stiky inserts were replaed. Pheromone lures were replaed every 4 wk, with lures from the same lot throughout eah season. Infestation by P. viteana was quantified by visually examining 5 lusters (25 lusters at two sampling sites) at

J Chem Eol (28) 34:189 195 191 the border and interior of eah plot. Both the number of lusters, and the number of berries infested were reorded in the first generation of GBM on 29 June 25 and 6 July 26, and during the seond generation on 15 August 25 and 4 August 26. Droplet Release Rates To determine the release of pheromone from wax droplets, five 1-ml droplets of GBM- SPLAT were applied to eah of five 3.5 2.9 m untreated wood garden stakes (Dayton Garden Labels, Dayton, OH, USA) and deployed on 24 May 26 at one orner of eah of the four pheromone-treated plots desribed above. Garden stakes were sored every 3.8 m with a utility knife to allow for ease of individual drop olletion. Eah stake was nailed to the north fae of a post loated farthest from monitoring traps. One drop was olleted from a randomly seleted stake in eah of the four treated plots eah week until the termination of the experiment. Eah sample was plaed into a 6-ml glass bottle (Qorpak, Bridgeville, PA, USA) and stored at 2 C until it was extrated using the proedure of Stelinski et al. (25), whih was modified from Meissner et al. (2). Fifty milliliters of an internal standard solution of 232 m1/l methyl myristate (99%, Aros Organis, Geel, Belgium) in aetonitrile (HPLC grade, EMD Chemials In., Gibbstown, NJ, USA), were added to eah sample. The bottles were plaed into a water bath shaker (model 4615, AO Sientifi Instruments, Keene, NH) at 7 75 C without shaking for 1 min, followed by 1 min of shaking, then briskly agitated by hand for 1 se, before an additional 3 min of shaking in the water bath. Samples were then hand-agitated for 1 se and frozen at 2 C to preipitate the wax. After 2 hr, samples were thawed, vortexed, and 1 ml of the solution was removed and filtered into a 2-ml GC vial (Agilent Tehnologies, Santa Clara, CA, USA) through a disposable glass Pasteur pipette fitted with a paper plug (Kimberly-Clark Corp., Roswell, GA, USA) at the tapered end. The pheromone in eah sample was quantified by gas hromatography (GC) (HP-689, Hewlett-Pakard, Palo Alto, CA, USA) by using a 3-m HP DB-23 polar olumn (model 122-2332, J&W Sientifi, Folsom, CA, USA) with the internal standard method (MNair and Miller 1998). Droplet Density Study This study was onduted during 26 within one large vineyard to determine the effet of droplet density on disruption of male P. viteana flight to pheromone traps. The experiment onsisted of a randomized omplete blok design repliated six times. Treatment plots were 15.2 16.5 m with 8 m between plots and between bloks. Plots reeived either, 1, 3, 1, or 3.2 ml droplets of wax ontaining 3% (v/v) P. viteana pheromone per vine, applied by hand with a 1-ml syringe, or Isomate GBM Plus rope dispensers (Shin-Etsu Chemial Co., Tokyo, Japan) at a density of 45/ha. Treatments were applied on 1 and 3 June 26. A large plasti delta trap (Suterra LLC, Bend, OR, USA), baited with rubber septa that ontained P. viteana sex pheromone (Suterra LLC, Bend, OR) was plaed in the middle of eah plot. Traps were monitored weekly until 24 August 26 to reord the number of male P. viteana aptured, at whih time the moths were removed or stiky inserts replaed. Pheromone lures were replaed every 4 wk with lures from the same lot. Droplet Size Study This experiment used the same protool as the droplet density study, but in this ase the traps were baited with -,.2-,.5-, or 1.-ml wax droplets (approximately 6, 15.9, and 3 mg P. viteana sex pheromone, respetively) applied to aluminum foil suspended inside the traps, or a rubber septum loaded with.1 mg P. viteana sex pheromone. All treatments were applied on 24 May 26, and the traps were monitored weekly until 24 August 26. To help minimize the effet of pest pressure within experimental bloks, treatments were rotated within eah blok every week, and droplets were hanged every 4 wk. Statistial Analyses In the mating disruption study, the signifiane of treatment effet on the number of moths aptured for 13 and 15 wk after appliation in 25 and 26, respetively, was ompared among treatments for both loations (plot interior and border) by using ANOVA (SAS/ STAT 23). For eah sample date, the average number of moths per trap was ompared among treatments within eah vineyard position by using ANOVA (SAS/STAT 23). All data were log-transformed (n+1) for normality prior to analysis, and Tukey s HSD test was used to determine differenes between means at α=.5. Perent disruption values were alulated as the proportional redution in the number of moths aught in the treated plots ompared to the untreated plots. Infestation data were arsine squareroot transformed prior to analysis and ompared between treatments at the border and at the interior positions by using ANOVA for eah sampling date (SAS/STAT 23). To determine the relationship between time after appliation and the amount of pheromone released, residual onentration of pheromone droplets in the release rate study was analyzed by regression analysis (SAS/STAT 23). These values also were ompared to the perent disruption values alulated from moth aptures, to determine a ritial release rate for disruption of P. viteana. The signifiane of treatment effet on total moths aptured in the droplet density and droplet size study was ompared among treatments by using analysis of variane (ANOVA) (SAS/STAT 23). Data were log-transformed (n+1) prior to analysis and Tukey s HSD test was used to determine differenes between means at α=.5.

192 J Chem Eol (28) 34:189 195 Mean number of moth aptures 8 Untreated Border Untreated Interior 7 SPLAT-treated Border 6 SPLAT-treated Interior 5 4 3 2 1 8 7 6 5 4 3 2 1 5/2 Results 5/9 5/16 5/23 5/3 6/6 6/13 Large-sale Mating Disruption Study Before appliation of treatments in 25, there was no signifiant differene in the number of moths trapped between treatments (F=3.67; df=1, 2; P=.2) or positions (F=.5; df=1, 8; P=.83), indiating that pest pressure was similar aross vineyards (Fig. 1). However, following the SPLAT-GBM treatments, the seasonal total number of moths per trap was lower in treated (2.6±.7) than untreated (148.9±28.1) plots (F=96.76; df=1, 3; P=.2) (Fig. 1), a pattern 6/2 Date 6/27 7/4 7/11 25 26 Fig. 1 Average number of adult male Paralobesia viteana aptured in pheromone traps at the vineyard border or vineyard interior of vineyard plots that were either untreated or treated with 1-ml droplets of SPLAT-GBM ontaining 3% pheromone 7/18 7/25 8/1 8/8 observed both within (F=126.6; df=1, 3; P=.2) and at the borders (F=37.55; df=1, 3; P=.9) of plots (Fig. 1). In 26, P. viteana were not trapped before treatment appliations (moths had been trapped at nearby vineyards), but post-treatment the aptures of P. viteana were signifiantly lower in the SPLAT-treated (36.5±9.3) than the untreated (279.6±47.2) vineyards (F=39.82; df=1, 3; P=.8). Most moth aptures in the treated plots ourred in the last 2 wk of the trial as the disruption effiieny of the formulations delined (Fig. 1). In ontrast to 25, in 26 more male P. viteana were aptured at the interior ompared to the border (F=35.4; df=1, 12; P<.1) within plots. However, similar to 25, fewer moths were aptured at the interior (F=33.97; df=1, 3; P=.1) and the border (F=21.22; df=1, 3; P=.19) of treated plots ompared to untreated plots. Perent disruption exeeded 9% at the vineyard interior for approximately 12 wk in 25 and 11 weeks in 26. In 26, when there were no insetiide appliations to the vineyards, the level of disruption had delined to 22.7% after 13 wk. In both years, disruption was greater at the interior traps than in traps plaed at the borders. In 25, neither the total number of larvae found in eah sample (F.53; df=1, 3; P.52) nor the proportion of lusters infested with P. viteana larvae (F 3.71; df=1, 3; P.15) varied signifiantly between treatments or vineyard positions for either sampling date (Table 1). During the first generation samples of 26, there was no signifiant differene between treatments in the proportion of lusters infested or the number of larvae in luster samples at the border (F=2.32; df=1, 3; P=.23) or interior (F=6.91; df=1,3; P=.78) of the vineyard (Table 1). However, during the seond generation, there were differenes in these parameters between the SPLAT-treated and untreated borders (F=12.76; df=1, 3; P=.38). Droplet Release Rates Droplets of SPLAT-GBM released pheromone as a first-order exponential deay funtion Table 1 Average number of Paralobesia viteana larval berry infestations and the perentage of lusters infested with P. viteana (±S.E.) at the borders and interiors of vineyard plots that were untreated or treated with SPLAT-GBM during 25 and 26 Date Position No. of infestations per 25 lusters Perent of lusters infested Untreated SPLAT P value Untreated SPLAT P value 6/29/25 Border 6.1±1.3 7.4±2.2.62 7.8±1. 1.±1.8.15 Interior 1.9±.7 1.4±.9.52 2.8±.9 2.±1.1.48 8/15/25 Border _ 2.±2. 21.5±3.2.64 Interior _ 5.8±3.4 3.5±1.6.33 6/27/26 Border 11.1±2.5 7.5±1.4.23 15.8±4.5 9.5±1.3.23 Interior 6.8±.6 3.4±1.1.78 9.3±1. 6.5±2.2.27 8/4/26 Border 22.5±1.9 18.4±1.5.38 35.5±3.5 25.8±2.4.4 Interior 11.8±3.9 8.6±1.2.43 18.±5.9 15.3±2.2.7

J Chem Eol (28) 34:189 195 193 (Z)9-12 A remaining (mg) 6 5 4 3 2 1 y = 4E+14e -.82x R 2 =.83 5/23 6/12 7/2 7/22 8/11 8/31 9/2 Date Fig. 2 Release of pheromone from 1-ml droplets of SPLAT-GBM during 26. The release profile best fits an exponential deay funtion, and the best-fit urve and equation are presented on the graph (Fig. 2), although the high variability among repliates (shown by the high standard error values), resulted in a relatively low preditive power of the exponential equation. The release funtion suggests that almost 5% of the pheromone in the wax droplets was still bound in the wax in late July when aptures of P. viteana were first observed in the SPLAT-treated plots. Droplet Density Study Signifiantly more moths were aptured in the untreated ontrol plots ompared to all other treatments, but the one and three drops per vine treatments had higher moth aptures than the 1 drops per vine, 3 drops per vine, and Isomate treatments (F=27.6; df=5, 25; P<.1) (Fig. 3). Droplet Size Study The size of eah droplet affeted how many moths were trapped (F=66.1; df=4, 2; P<.1) (Fig. 4). The.2- and.5-ml droplets trapped low numbers Mean number of moths ± S.E. 15 12 9 6 3 a b b 1 3 1 3 Ropes Treatment Fig. 3 Average (+SE) number of adult male Paralobesia viteana aptured in pheromone traps in vineyard plots treated with varying densities of.2-ml droplets of SPLAT-GBM wax that ontained 3% sex pheromone, or pheromone ropes. Bars with the same letter are not signifiantly different at α=.5 Mean number of moths ± S.E. 3 25 2 15 1 5 of moths, and inreasing the droplet size to 1. ml aused a slight derease in the number of moths trapped (Fig. 4). The greatest aptures of moths were in traps baited with the lure that ontained the standard.1 mg of sex pheromone, an amount muh lower than that in any of the applied droplets (approximately 6 mg or greater). No moths were aught when the traps were unbaited. Disussion a d Septum ml.2 ml.5 ml 1 ml Treatment Fig. 4 Average (+SE) number of adult male Paralobesia viteana aptured in pheromone traps baited with different sizes of SPLAT- GBM wax droplets that ontained 3% sex pheromone, or a ommerial pheromone lure ontaining.1 mg of sex pheromone. Bars with the same letter are not signifiantly different at α=.5 Appliation of 1-ml droplets of SPLAT-GBM that ontained 3% pheromone to vineyards before the first generation flight of P. viteana resulted in high levels of disruption of male moth orientation to monitoring traps during two growing seasons, with redued rop infestation in treated plots in the seond growing season. This first report of using a wax formulation for pheromone deployment in vineyards provides evidene for the potential of this formulation to ontrol P. viteana. Wooden vineyard posts provided a pratial target for appliation of the wax, and droplets applied on the north side of these strutures in early May eah year remained in plae throughout the growing season. These droplets were easy to apply by hand with a 3-ml syringe, and their appliation was faster than that of Isomate twist ties at 45/ha (Isaas, unpublished data). The large droplets used in this study released pheromone until late July. The longevity of these droplets is lower ompared with those used for mating disruption of G. molesta (Stelinski et al. 26, 27). Pheromone dispenser density affets the distribution of pheromone that permeates the rop habitat, thus influening the degree of mating disruption ahieved (Rothshild 1975; Flint and Merkle 1983; Lawson et al. 1996; Stelinski et al. 25; Miller et al. 26a, b). A similar pattern was found in this study, as trap athes in untreated plots were greater than one or three droplets per vine (.2-ml volume) >1 and 3 droplets per vine. The higher doses gave similar results b b

194 J Chem Eol (28) 34:189 195 as Isomate GBM ropes applied one every.4 vine. Overall, our results are similar to those observed for G. molesta (Stelinski et al. 25), although they reported higher densities of droplets outperformed twist ties and ompletely prevented mating of tethered female moths. A profile analysis of variation in moth ath with dispenser density, similar to that onduted by Miller et al. (26a), was not possible in our study beause of an insuffiient range of pheromone point-soure densities. A pheromone formulation lasting >2 wk, as ahieved with Isomate-GBM twist ties, is neessary to over the full ativity period of P. viteana in the Great Lakes region of the US and Canada. Our results suggest that this annot be urrently ahieved by using SPLAT-GBM, so one approah to would be to re-treat vineyards in late July. Alternatively, one ould apply an appropriate Lepidopteraspeifi insetiide for ontrol of first-generation P. viteana, followed by appliation of SPLAT-GBM in early July to provide protetion against this pest until harvest. Another promising line of researh would be the development of a wax formulation that provides a pheromone release system lasting >2 wk. Development of formulations that slowly release the pheromone is an important goal for mating disruption, not only to provide a long period of ativity but also to ensure that expensive pheromone is not wasted by remaining bound in the wax matrix (de Lame 23). Residual analysis of wax revealed that approximately 5% of the P. viteana pheromone was still bound in the drops when disruption performane delined in late July. The 1-ml droplets used in this study are larger than those deployed in many reent studies (Stelinski et al. 25, 26, 27), so the use of more, smaller-sized droplets should result in a greater proportion of the pheromone being released (Stelinski et al. 25). This is urrently under investigation, espeially as the amount of pheromone released per hetare in the 1- drops-per-vine treatment (58.1 mg/ha) was lower than the Isomate treatment (99.8 mg/ha). Evaluations of sex pheromone formulations for ontrol of P. viteana have shown that mating disruption an help ontrol this pest (Dennehy et al. 199; Trimble 1993), yet there has been relatively low adoption of this approah in vineyard IPM programs in eastern North Ameria where vineyards are infested by P. viteana. Primary impediments to adoption are the pereption that pheromone formulations have lower effiay, and the ost of pheromone produts and their appliation is greater when ompared with insetiides. An additional hallenge to ontrolling infestation of P. viteana is that this inset typially auses higher levels of rop infestation at the vineyard borders than in the interiors (Hoffman and Dennehy 1989; Botero-Garés and Isaas 23). Our results showed that while appliation of SPLAT-GBM aused an immediate shutdown of traps, a signifiant redution in luster infestation at the vineyard borders was observed only in the seond year of treatment, suggesting that multiyear appliation of pheromone may be required for mating disruption of GBM. Aknowledgments We thank Agenor Mafra-Neto and Reg Coler at ISCA Tehnologies, In. for supplying the SPLAT-GBM used in these trials. Piera Siegert and Zsofia Szendrei provided tehnial assistane, and we thank Lukasz Stelinski, Larry Gut, and Jim Miller for input. We also thank the anonymous reviewers for onstrutive omments on the previous version of this manusript. This researh was funded in part by the National Grape Cooperative, the USDAVitiulture Consortium East, and the Mihigan Agriultural Experiment Station. Referenes ATTERHOLT, C. A. 1996. Controlled release of inset sex pheromones from sprayable biodegradable materials for mating disruption. Ph.D. dissertation, University of California-Davis. ATTERHOLT, C. A., DELWICHE, M. J., RICE, R. E., and KROCHTA, J.M. 1998. Study of biopolymers and paraffin as potential ontrolledrelease arriers for inset pheromones. J. Agri. Food Chem. 46:4429 4434. BOTERO-GARCÉS, N., and ISAACS, R. 23. Distribution of grape berry moth, Endopiza viteana (Lepidoptera: Tortriidae), in natural and ultivated habitats. Environ. Entomol. 32:1187 1195. de LAME, F. M. 23. Improving mating disruption programs for the Oriental fruit moth, Grapholita molesta (Busk): Effiay of new wax-based formulations and effets of dispenser appliation height and density. M.S. thesis, Mihigan State University. DENNEHY, T. J., HOFFMAN, C. J., NYROP, J. P., and SAUNDERS, M.C. 199. Development of low-spray, biologial, and pheromone approahes for ontrol of grape berry moth, Endopiza viteana Clemens, in the eastern United States., pp. 261 282, in N. J. Bostanian, L. T. Wilson, and T. J. Dennehy (eds.). Monitoring and Integrated Management of Arthropod Pests of Small Fruit CropsInterept Ltd., Andover, NH. EPSTEIN, D. L., STELINSKI, L. L., REED, T. P., MILLER, J. R., and GUT, L. J. 26. Higher densities of distributed pheromone soures provide disruption of odling moth (Lepidoptera: Tortriidae) superior to that of lower densities of lumped soures. J. Eon. Entomol. 99:1327 1333. FLINT, H. M., and MERKLE, J. R. 1983. Pink bollworm (Lepidoptera: Gelehidae): Communiation disruption by pheromone omposition imbalane. J. Eon. Entomol. 76:4 46. HOFFMAN, C. J., and DENNEHY, T. J. 1989. Phenology, movement, and within-field distribution of the grape berry moth, Endopiza viteana (Clemens) (Lepidoptera: Tortriidae), in New York vineyards. Can. Entomol. 121:325 335. JENKINS, P. E., and ISAACS, R. 27a. Cutting wild grapevines as a ultural ontrol strategy for grape berry moth (Lepidoptera: Tortriidae). Environ. Entomol. 36:187 194. JENKINS, P. E., and ISAACS, R. 27b. Redued-risk insetiides for ontrol of grape berry moth (Lepidoptera: Tortriidae) and onservation of natural enemies. J. Eon. Entomol. 1:855 865. LAWSON, D.S.,REISSIG, W.H.,AGNELLO, A.M.,NYROP, J.P.,and ROELOFS, W. L. 1996. Interferene with the mate-finding ommuniation system of the obliquebanded leafroller (Lepidoptera: Tortriidae) using syntheti sex pheromones. Environ. Entomol. 25:895 95. MCNAIR, H. M., and MILLER, J. M. 1998. Basi Gas Chromatography. Wiley, New York, NY.

J Chem Eol (28) 34:189 195 195 MEISSNER, H. E., ATTERHOLT, C. A., WALGENBACH, J. F., and KENNEDY, G. G. 2. Comparison of pheromone appliation rates, point soure densities, and dispensing methods for mating disruption of tufted apple bud moth (Lepidoptera: Tortriidae). J. Eon. Entomol. 93:82 827. MILLER, J. R., GUT, L. J., de LAME, F. M., and STELINSKI, L.L. 26a. Differentiation of ompetitive vs. non-ompetitive mehanisms mediating disruption of moth sexual ommuniation by point soures of sex pheromone (Part I): Theory. J. Chem. Eol. 32:289 2114. MILLER, J. R., GUT, L. J., de LAME, F. M., and STELINSKI, L.L. 26b. Differentiation of ompetitive vs. non-ompetitive mehanisms mediating disruption of moth sexual ommuniation by point soures of sex pheromone (Part II): Case studies. J. Chem. Eol. 32:2115 2143. ROELOFS, W. L., TETTE, J. P., TASCHENBERG, E. F., and COMEAU, A. 1971. Sex pheromone of the grape berry moth: Identifiation by lassial and eletroantennogram methods, and field tests. J. Inset. Si. 17:2235 2243. ROTHSCHILD, G. H. L. 1975. Control of the Oriental fruit moth (Cydia molesta) (Busk) (Lepidoptera: Tortriidae) with syntheti female pheromone. Bull. Entomol. Res. 65:473 49. SAS INSTITUTE. 23. SAS/STAT User's Manual, Version 9.1. SAS Institute, Cary, NC. STELINSKI, L. L., GUT, L. J., MALLINGER, R. E., EPSTEIN, D. L., REED, T. P., and MILLER, J. R. 25. Small plot trials doumenting effetive mating disruption of oriental fruit moth by using high densities of wax-drop pheromone dispensers. J. Eon. Entomol. 98:1267 1274. STELINSKI, L. L., MILLER, J. R., LEDEBUHR, R., and GUT, L. J. 26. Mehanized appliator for large sale field deployment of paraffin wax dispensers of pheromone for mating disruption in tree fruit. J. Eon. Entomol. 99:175 171. STELINSKI, L. L., MILLER, J. R., LEDEBUHR, R., SIEGERT, P., and GUT, L. J. 27. Season-long mating disruption of Grapholita molesta (Lepidoptera: Tortriidae) by one mahine appliation of pheromone in wax drops (SPLAT-OFM). Journal of Pestiide Siene 8:19 117. TOBIN, P. C., NAGARKATTI, S., and SAUNDERS, M. C. 23. Phenology of grape berry moth (Lepidoptera: Tortriidae) in ultivated grape at seleted geographi loations. Environ. Entomol. 32:34 346. TRIMBLE, R. M. 1993. Effiay of mating disruption for ontrolling the grape berry moth, Endopiza viteana (Clemens) (Lepidoptera: Tortriidae), a ase study over three onseutive growing seasons. Can. Entomol. 125:1 9. TRIMBLE, R. M., PREE, D. J., VICKERS, P. M., and KER, K. W. 1991. Potential of mating disruption using sex-pheromone for ontrolling the grape berry moth, Endopiza viteana (Clemens) (Lepidoptera: Tortriidae), in Niagara peninsula, Ontario vineyards. Can. Entomol. 123:451 46. TRIMBLE, R. M., VICKERS, P. M., NIELSEN, K. E., and BARINSHTEYN, G. 23. Sprayable pheromone for the ontrolling the North Amerian grape berry moth by mating disruption. Agri. For. Entomol. 5:263 268. WITZGALL, P., BENGTSSON, M., and TRIMBLE, R. M. 2. Sex pheromone of grape berry moth (Lepidoptera: Tortriidae). Environ. Entomol. 29:433 436.