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Journal of Food Protection, Vol. 76, No. 3, 2013, Pages 456–461 doi:10.4315/0362-028X.JFP-12-301 Copyright G, International Association for Food Protection
Mating-Disruption Trials for Control of Mediterranean Flour Moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), in Traditional Flour Mills
P. TREMATERRA* AND G. SPINA
Department of Agriculture, Environmental and Food Science, University of Molise, via de Sanctis, 86100 Campobasso, Italy Downloaded from http://meridian.allenpress.com/jfp/article-pdf/76/3/456/1684342/0362-028x_jfp-12-301.pdf by guest on 01 October 2021
MS 12-301: Received 10 July 2012/Accepted 17 August 2012
ABSTRACT In the present work, 3 years of field trials (from 2007 to 2009) were carried out to evaluate use of the pheromone (9Z,12E)- tetradecadienyl acetate (TDA) for mating disruption (MD) of the Mediterranean flour moth, Ephestia kuehniella Zeller. Dispensers containing the pheromone TDA were placed in two traditional flour mills. Pheromone-baited funnel traps were used to monitor the population fluctuations of E. kuehniella males throughout the entire experimental period; female oviposition was assessed by placement of petri cups containing wheat germ–semolina flour bait. According to our results, the use of MD dispensers does not interfere completely with the reproduction of E. kuehniella. However, looking at the overall data, there was a significant reduction in both adults and larvae in treated mills after the MD application. According to hazard analysis and critical control point procedures, treatment should be accompanied by general cleaning of the facilities, including corners and inside machinery, where insects can hide and reproduce. In integrated pest management programs, the use of mating disruption can lead to a drastic reduction in the need for chemical treatments, with improvement in food quality.
Mating disruption is one of the most promising pest whether it was possible to disrupt the mating of the control methods in crop protection. The method is based on Mediterranean flour moth, Ephestia kuehniella Zeller, in the deployment of a synthetic pheromone of the target two different mills. species that ‘‘covers’’ release of the pheromone by unmated females. This interferes with the males’ ability to find MATERIALS AND METHODS females, resulting in reduced mating and, consequently, egg Flour mills. The two mills, one old (built in 1950, flour mill laying by females (10, 28). Pheromones are generally A) and one new (built in 2002, flour mill B), were selected because considered to be relatively safer and environmentally more they are representative of traditional flour mills in Central Italy. acceptable than conventional pesticides, because they occur naturally, are species specific, possess low acute mamma- Flour mill A. The mill was concrete with some wooden lian toxicity, and are usually volatile chemicals that do not sections (e.g., cleaning section, flour milling section, and storage leave harmful residues (10). silos) and consisted of two floors, the ground floor and the first floor (6 by 8 m each). No chemical treatments were applied during In recent years, considerable progress has been made the experimental period. Previous monitoring indicated high not only in monitoring, but also in directly controlling some infestation levels of E. kuehniella at this location. stored-product insects with pheromones (1, 4, 5, 7, 11, 12, During 2007 and 2008, after 4 weeks of pretreatment 14, 26–28). monitoring, CheckMate SPM MD dispensers (Suterra, Bend, Because there will be a need to adopt approaches that OR) were applied to the mill. The MD dispensers consisted of a consider all risks from pests and from pest control cellulose pad loaded with 50 mg of the pheromone (9Z,12E)- treatments in the integrated pest management (IPM) context, tetradecadienyl acetate (TDA) and sealed in a polyethylene sachet. the use of these substances can lead to a drastic reduction in In 2007, dispensers were placed at a density of one dispenser 2 chemical treatments. When managing stored-product insect per 9 m , and attached by hand to machinery, shelves, and other pests, further reductions in chemical control can be achieved suitable stationary structures by using hangers (walls, wooden by replacing pesticides with biological and physical control parts, etc.); in 2008, MD dispensers were applied at a density of one dispenser per 6 m2. Dispensers were placed in mid-August in methods, and biorational approaches (13). both years. In 2009, MD dispensers were applied at a density of In such a context, the aim of this study was to see, by one dispenser per 6 m2 and were placed on 4 May (when the using mating-disruption techniques from 2007 to 2009, annual flight of Ephestia begins) and were replaced once, on 13 July. * Author for correspondence. Tel: z39-0874-404655; Fax z39-0874-404855; In 2008 and 2009, a trial was also performed to flour mill A to E-mail: [email protected]. investigate if there was a benefit from increasing the number of J. Food Prot., Vol. 76, No. 3 MATING DISRUPTION OF MEDITERRANEAN FLOUR MOTH 457
TABLE 1. Flour mills involved in the experiments, MD dispensers, and oviposition traps Flour mill Year Dispenser rate (m2) Size (m2) MD dispensers (n) Pheromone monitoring traps (n) Oviposition cups (n)
A 2007 9 110 12 4 inside 4 (a, b, c, d) 2008 6 110 16 4 inside and 2 outside 4 (a, b, c, d) 2009 6 110 16 4 inside and 2 outside 4 (a, b, c, d) B 2007 9 252 26 2 inside 2 (a, b) 2008 9 352 39 4 inside and 2 outside 4 (a, b, c, d) 2009 9 352 39 4 inside and 2 outside 4 (a, b, c, d)
MD dispensers in a given space, in which there was an increase Flour mill A. During 2007, pheromone-monitoring from 12 dispensers (one dispenser per 9 m2) to 16 dispensers (one traps caught 9,180 E. kuehniella males (4,654 on the ground
2 Downloaded from http://meridian.allenpress.com/jfp/article-pdf/76/3/456/1684342/0362-028x_jfp-12-301.pdf by guest on 01 October 2021 dispenser per 6 m ) (Table 1). floor and 4,526 on the first floor of the mill). In 2008, 10,724 males were trapped (7,136 on the ground floor and Flour mill B. The building was made of concrete and 3,588 on the first floor); outside the mill, traps caught an consisted of two floors, the ground floor (14 by 18 m) and the first additional 417 males. During 2009, 7,346 males were floor (8 by 12.5 m). The Mediterranean flour moth was also the dominant moth species at this mill location. No chemical trapped inside the mill (4,738 on the ground floor and 2,608 treatments were applied during the experimental period. During on the first floor); outside the mill, 156 adult males were 2007 and 2008, after 4 weeks of pretreatment monitoring, found (Figs. 1 and 2). CheckMate SPM MD dispensers, each with 50 mg of TDA, were In both 2007 and 2008, E. kuehniella male captures were applied inside the mill. MD dispensers were placed at a density of extremely high before the beginning of the mating-disruption one dispenser per 9 m2. In 2007, the experiment was conducted on program, with a rapid decrease in adults observed soon after the ground floor; dispensers were placed on 9 August and were the placement of the MD dispensers (Fig. 1). As can be seen, replaced once, on 11 September (26 dispensers were distributed on in the subsequent months there was a clear decrease in the the ground floor). In 2008, MD dispensers were placed on 19 presence of adults in the environment. The percent reductions August (26 on the ground floor and 13 on the first floor); in 2009, in moth captures after implementing mating disruption were the same quantity of MD dispensers was applied on 27 April and 49.2% in 2007 and 72.2% in 2008 (Table 2). During the replaced once, on 13 July (Table 1). premating disruption period of 2008, captures outside the mill Monitoring of adult males. Monitoring of E. kuehniella varied from 13 to 81 males per trap and tended to decrease in adults was conducted with blackstripe funnel traps baited with 1 mg the fall, perhaps because of the presence of the indoor MD of TDA (AgriSense-BCS, Pontypridd, UK), hung 1.80 m above the dispensers or seasonal patterns of activity (Fig. 2). Compared floor. From 2007 to 2009, to give an indication of population with previous years (2007 and 2008), during 2009, when levels, monitoring traps were checked on a weekly basis from the control with mating disruption had been achieved for the beginning of each experiment until December. The number of entire season (from March to December), the number of Mediterranean flour moth males was recorded. The distribution of males trapped inside and outside the mill was lower, and a monitoring traps, inside and outside the flour mills, is reported in large proportion of the captures were observed from late June Table 1. to late August (Figs. 1 and 2). The total number of adults collected 4 weeks before and In trials after the placement of MD dispensers, the 4 weeks after the pheromone MD application in treated mills A and presence of larvae in the oviposition cups was observed B were compared by using contingency tables and chi-squared occasionally during all weeks of the monitoring period. In analysis, performed with SPSS software, version 13.0 (SPSS, Chicago, IL). The Yates correction was applied when an observed 2007, cups a and b were found infested three times and one frequency was less than 5. time, respectively. In 2008, cups a, b, c, and d were found infested two, four, one, and four times, respectively. During Oviposition monitoring. E. kuehniella female oviposition 2009, larvae in the oviposition cups were observed two was assessed by placing petri cups containing wheat germ– times in each of cups a, b, and d, and three times in cup c. semolina flour bait close (1.5 m) to the pheromone-monitoring traps. The bait cups were replaced at weekly intervals. Bait cups Flour mill B. In 2007, pheromone traps positioned were kept in an incubator for an additional period of 3 to 4 weeks inside the mill captured 3,506 males of E. kuehniella on the (at 27uC and 75% relative humidity). Cups were then examined for ground floor. During 2008, 4,660 males were trapped (1,551 presence or absence of larvae. The distribution of oviposition cups on the ground floor and 3,109 on the first floor of the mill); in the flour mills is reported in Table 1. outside the mill, a further 463 males were found. In 2009, 11,115 males were trapped (4,155 on the ground floor and RESULTS AND DISCUSSION 6,960 on the first floor of the mill); outside the mill 325 Figures 1 through 4 outline the captures in the traps males were trapped (Figs. 3 and 4). installed inside and outside the experimental flour mills A In 2007 and 2008, captures of E. kuehniella adult males and B. Significant differences were noted, suggesting a during the premating-disruption period were high, but different population trend after the MD dispensers were decreased soon after placement of MD dispensers, although deployed (Table 2). the reduction in moth captures was less in 2007 (21.3%) 458 TREMATERRA AND SPINA J. Food Prot., Vol. 76, No. 3 Downloaded from http://meridian.allenpress.com/jfp/article-pdf/76/3/456/1684342/0362-028x_jfp-12-301.pdf by guest on 01 October 2021
FIGURE 1. Flour mill A. Mean number of Ephestia kuehniella male adults caught in the pheromone traps during treatment with mating disruption (MD): inside the mill from 2007 to 2009. Arrows indicate start of MD treatment. FIGURE 2. Flour mill A. Mean number of Ephestia kuehniella male adults caught in the pheromone traps during treatment with mating disruption (MD): outside the mill in 2008 and 2009. than in 2008 (86.8%) (Fig. 3 and Table 2). In 2008, outside It was observed that when MD dispensers were the mill, the male numbers were rather stable from the first suspended during late August or even early autumn, the week in August until early September (Fig. 4). In 2009, constant reduction in captures in pheromone-baited traps when control by mating disruption had been achieved for could be partially attributed to the reduced temperatures, a the entire season (March through December) the population factor that reduces male flying activity. The fact that mating of Ephestia remained low during the first months of disruption was applied once per year (and only for a certain implementation of the mating disruption (in March through period) somewhat late in the season could have partially June); subsequently, an increase was observed in males enabled the increase in moth populations early in the season. trapped in early July, and a decrease just after replacing the The application of the MD dispensers earlier in the season, MD dispensers in August. Thereafter (until December), the or throughout the entire year, is likely to provide increased number of males captured by traps remained rather high, efficacy in comparison with the application of the MD probably because the favorable temperatures present inside dispensers late in the season, even if this is the critical the mill (from 20 to 34uC) did not interrupt the biological season for the infestation. cycle of the Mediterranean flour moth. Outside the mill, In flour mill A, it was observed that placing the MD male numbers were low and fairly stable from the first week dispensers at the beginning of the annual flight of Ephestia of May until early October (Fig. 4). (in March) and increasing the density of the MD dispensers In this mill, larvae were occasionally observed in the (from one dispenser per 9 m2 to one dispenser per 6 m2) oviposition cups during all weeks of the 2007 monitoring reduced pest populations both inside and outside the mill period: cups a and b were infested two and three times, (Figs. 1 and 2). Moreover, it was noted that in the unheated respectively. Similarly, during 2008, cups a and b were mill during the cold months, low temperatures acted found to be infested zero and twice; cups c and d were not negatively on the population of Ephestia, with reductions found to be infested. In 2009, larvae were observed two in the number of adults trapped (Figs. 1 and 2). times in cup a, a single time in cup b, and two times in cup In flour mill B, it was observed that after MD c, whereas cup d was never found to be infested. application, the number of males captured by monitoring Regarding the overall data in mill A and mill B, there traps decreased; however, if the application of the MD were significant reductions in both adults and larvae in the dispenser was not done also against autumn populations of treated mills after MD application. Ephestia, infestation tended to increase again (as observed J. Food Prot., Vol. 76, No. 3 MATING DISRUPTION OF MEDITERRANEAN FLOUR MOTH 459 Downloaded from http://meridian.allenpress.com/jfp/article-pdf/76/3/456/1684342/0362-028x_jfp-12-301.pdf by guest on 01 October 2021
FIGURE 3. Flour mill B. Mean number of Ephestia kuehniella male adults caught in the pheromone traps during treatment with mating disruption (MD): inside the mill from 2007 to 2009. Arrows indicate start of MD treatment. FIGURE 4. Flour mill B. Mean number of Ephestia kuehniella male adults caught in the pheromone traps during treatment with mating disruption (MD): outside the mill in 2008 and 2009. in 2009) (Fig. 3). In addition, favorable temperatures (Walker); the Mediterranean flour moth, E. kuehniella present inside the mill lent to increased population counts. Zeller; the tobacco moth, Ephestia elutella (Hu¨bner); When MD dispensers were applied at the beginning of the the raisin moth, Ephestia figulilella (Gregson); and the season, or throughout the entire year, it was observed that Indianmeal moth, Plodia interpunctella (Hu¨bner) (2, 28). the population of E. kuehniella remained at fairly low levels This ‘‘multispecies pheromone’’ has been evaluated as all year (Fig. 3). In any case, the moth captures in each trap a mating-disruption agent, with preliminary results in both underlined their different individual activities, based on their laboratory and field tests. Brady and Daley (3) and Hodges location and on the greater or lesser quantity of food debris et al. (9) conducted studies of mating disruption in C. present in the immediate vicinity. cautella, with promising results for the wider use of this The procedures adopted for the application of mating- technique in large storerooms. Sower et al. (22) showed a 90 disruption pheromones inside the mill also influenced the to 95% reduction in mating in small rooms when P. size of the Ephestia population found outside the structure interpunctella densities were low. Sower and Witmer (23) (Figs. 2 and 4). and Prevett et al. (15) noted a significant reduction in mating The pheromone TDA attracts males of several pyralid and population growth of both P. interpunctella and C. moths, including the almond moth, Cadra cautella cautella in warehouse rooms treated with high TDA
TABLE 2. Flour mills and changes in the mean trap catches between 4 weeks before and 4 weeks after pheromone mating disruption treatment Adult males caught (n)
Flour mill Year Interval I (4 weeks before) Interval II (4 weeks after) % x2 df a
A 2007 854.5 434 249.2 211.16**b 3 2008 1,675 465 272.2 355.50** 3 B 2007 1,029 810 221.3 32.65** 3 2008 870 115.3 286.8 50.18** 3 a df, degrees of freedom. b **, significantly different from random at P , 0.01. 460 TREMATERRA AND SPINA J. Food Prot., Vol. 76, No. 3 concentrations. Fadamiro and Baker (8) reported that In the future, the effect of mating-disruption phero- pheromone puffs suppressed mating of P. interpunctella in mones on population growth or decrease could be measured an infested corn store. Shani and Clearwater (20) noted that by the presence of spermatophores in bursa copulatrix of an increased presence of pheromone disrupted mating of C. females, which is a good indicator of mating activity. cautella adults and influenced progeny production. More- In our case, the use of the MD dispensers did not over, in the same species, Ryne et al. (16) noticed a 94% completely block the reproduction of E. kuehniella. How- reduction in male captures of C. cautella in pheromone- ever, extrapolation of the recorded data suggests that the use baited traps in a chocolate factory in Sweden. Sieminska et of mating disruption in a traditional mill for a longer time al. (21) and Trematerra et al. (26, 29) reported interesting should dilute the population of the Mediterranean flour results on control of E. kuehniella by attracticide and moth even further. mating-disruption methods in flour mills. Savoldelli et al. Pheromone-based methods need to be considered as (19) reported results of field trials evaluating mating one part of an overall IPM program in food systems. disruption of C. cautella in two Italian chocolate factories. According to hazard analysis and critical control point Mating disruption to control P. interpunctella in a dried procedures, they should be accompanied by general Downloaded from http://meridian.allenpress.com/jfp/article-pdf/76/3/456/1684342/0362-028x_jfp-12-301.pdf by guest on 01 October 2021 bean storage and processing facility was reported by Burks cleaning of the facilities, including in corners and inside et al. (6). machinery, where insects can hide and reproduce. Particu- Several studies from different parts of the world have larly in traditional mills, it is necessary to clean thoroughly shown more or less similar results for moths infesting and to remove all debris, reducing the opportunities for stored products and food factories (1, 11, 14, 16, 17, 24– nesting by E. kuehniella. 28). These positive results indicate that the use of mating In IPM programs, the use of mating disruption can lead disruption for the control of stored product moths should to drastic reductions in chemical treatments, with economic be further evaluated as a component of an IPM-based advantages and improvements in food quality. If such control strategy. measures are not observed, the pheromones will, at best, However, there are problems in evaluating mating only reduce the number of insecticidal treatments necessary, disruption in practice, such as to define what a replicate is which in such cases will remain essential. Implementation and to estimate control based solely on trap captures (2). of the use of pheromones in IPM programs can be increased Ryne et al. (17) compared two adjacent storage rooms, one through educational and training programs for pest treated with mating disruption, the other not, and found by management service providers and food industry managers. using electrophysiological recordings (male antennal re- sponse), there was leakage of the pheromone into the REFERENCES untreated room. Sieminska et al. (21) reported that each 1. Anderbrant, O., C. Ryne, P.-O. C. Olsson, E. Jirle, K. Johnson, and food processing and storage facility is unique, and this C. Lo¨fstedt. 2007. Pheromones and kairomones for detection and makes finding a ‘‘control facility’’ similar to the treated control of indoor pyralid moths. IOBC-WPRS Bull. 30:73–77. facility extremely difficult. For this reason, mating-disrup- 2. Anderbrant, O., C. Ryne, E. Sieminska, G. P. Svensson, P.-O. C. tion experiments usually use a single or small number of Olsson, E. Jirle, and C. Lo¨fstedt. 2009. Odour signals for detection and control of indoor pyralid moths. IOBC-WPRS Bull. 41:69–74. treatment and control rooms (29). As a result, there is still 3. Brady, U. E., and R. C. Daley. 1975. Mating activity of Cadra inadequate information on mating-disruption effectiveness cautella during exposure to synthetic sex pheromone and related under different microclimates and in different types of compounds in the laboratory. Environ. Entomol. 4:445–447. facilities. 4. Burkholder, W. E. 1990. Practical use of pheromones and other Mating disruption could have a cumulative effect after attractants for stored-product insects, p. 497–516. In R. L. Ridgway, R. M. Silverstein, and M. N. Inscoe (ed.), Behavior-modifying multiple years of application in the same facility. At several chemical for insect management. Marcel Dekker, New York. locations, as verified also in our trial, there was a trend 5. Burkholder, W. E., and M. Ma. 1985. Pheromones for monitoring and toward lower captures in the second year of the study in the control of stored-product insects. Ann. Rev. Entomol. 30:257–272. facilities under mating disruption. 6. Burks, C. S., J. R. McLaughlin, J. R. Miller, and D. G. Brandl. 2011. The number of captured males or the absence of males Mating disruption for control of Plodia interpunctella (Hu¨bner) (Lepidoptera: Pyralidae) in dried beans. J. Stored Prod. Res. 47:216– in pheromone traps might not be a clear indicator of mating 221. disruption and female oviposition suppression. Oviposition 7. Cox, P. D. 2004. Potential for using semiochemicals to protect stored and/or immature emergence should be monitored in products from insect infestation. J. Stored Prod. Res. 40:1–25. conjunction with adult activity. Historical data from 8. Fadamiro, H. Y., and T. C. Baker. 2002. Pheromone puffs suppress previous years concerning both adult captures and larval mating by Plodia interpunctella and Sitotroga cerealella in an presence in the target facilities (if available) could serve infested corn store. Entomol. Exp. Appl. 102:239–251. 9. Hodges, R. J., F. P. Benton, D. R. Hall, and R. Dos Santos Serodio. as an internal control, because such information shows 1984. Control of Ephestia cautella (Walker) (Lepidoptera: Phyciti- seasonal patterns of insect activity. dae) by synthetic sex pheromone in the laboratory and store. J. Stored The results of mating disruption are, above all, Prod. Res. 20:191–197. conditioned by the density of the populations present. In 10. Jones, O. T. 1998. The commercial exploitation of pheromones and fact though, the nesting of insects in the remotest corners, other semiochemicals. Pest Sci. 54:293–296. 11. Phillips, T. W. 1997. Semiochemicals of stored-product insects: within tubes and machinery, makes it impossible at present research and applications. J. Stored Prod. Res. 33:17–30. to render sufficiently precise evaluations of the population 12. Phillips, T. W., P. M. Cogan, and H. Y. Fadamiro. 2000. sizes that can be impacted in real life scenarios. Pheromones, p. 273–230. In B. Subramanyam and D. W. Hagstrum J. Food Prot., Vol. 76, No. 3 MATING DISRUPTION OF MEDITERRANEAN FLOUR MOTH 461
(ed.), Alternatives to pesticides in stored-product IPM. Kluwer 21. Sieminska, E., C. Ryne, C. Lo¨fstedt, and O. Anderbrant. 2009. Long- Academic Publishers, Norwell, MA. term pheromone-mediated mating disruption of the Mediterranean 13. Phillips, T. W., and J. E. Throne. 2010. Biorational approaches to flour moth, Ephestia kuehniella, in a flour mill. Entomol. Exp. Appl. managing stored-product insects. Ann. Rev. Entomol. 55:375–397. 131:294–299. 14. Plarre, R. 1998. Pheromones and other semiochemicals of stored 22. Sower, L. L., W. K. Turner, and J. C. Fish. 1975. Population-density- product insects. A historical review, current application, and perspec- dependent mating frequency among Plodia interpunctella (Lepidop- tive needs. Mitt. Biol. Bundesanst. Land- Forstwirtsch. 342:13–83. tera: Phycitidae) in the presence of synthetic sex pheromone with 15. Prevett, P. F., F. P. Benton, D. R. Hall, R. J. Hodges, and R. Dos behavioral observations. J. Chem. Ecol. 1:335–342. Santos Serodio. 1989. Suppression of mating in Ephestia cautella 23. Sower, L. L., and G. P. Witmer. 1977. Population growth and mating (Walker) (Lepidoptera: Phycitidae). J. Stored Prod. Res. 25:147–154. success of Indianmeal moths and Almond moths in the presence of 16. Ryne, C., M. Ekeberg, N. Jonze´n, C. Oehlschlager, C. Lo¨ftedt, and O. synthetic sex pheromone. Environ. Entomol. 6:17–20. Anderbrant. 2006. Reduction in an almond moth Ephestia cautella 24. Su¨ss, L., D. P. Locatelli, and R. Marrone. 1996. Possibilities and (Lepidoptera: Pyralidae) population by means of mating disruption. limits of mass trapping and mating disruption techniques in the Pest Manag. Sci. 62:912–918. control of Ephestia kuehniella (Zell.) (Lepidoptera Phycitidae). Boll. 17. Ryne, C., G. P. Svensson, O. Anderbrant, and C. Lo¨ftedt. 2007. Zool. Agr. Bachic. 28:77–89. Evaluation of long-term mating disruption of Ephestia kuehniella 25. Su¨ss, L., D. P. Locatelli, and R. Marrone. 1999. Mating suppression and Plodia interpunctella (Lepidoptera: Pyralidae) in indoor of the Mediterranean flour moth (Ephestia kuehniella Zeller) Downloaded from http://meridian.allenpress.com/jfp/article-pdf/76/3/456/1684342/0362-028x_jfp-12-301.pdf by guest on 01 October 2021 storage facilities by pheromone traps and monitoring of relative (Lepidoptera: Pyralidae) in a food industry. Boll. Zool. Agrar. aerial concentrations of pheromone. J. Econ. Entomol. 100:1017– Bachic. 31:59–66. 1025. 26. Trematerra, P. 1994. Control of Ephestia kuehniella Zeller by sex 18. Ryne, C., G. P. Svensson, and C. Lo¨fstedt. 2001. Mating disruption pheromones in the flour mills. Anz. Schaedlingskd. Pflanzenschutz of Plodia interpunctella in small-scale plots: effects of pheromone Umweltschutz 67:74–77. blend, emission rates and population density. J. Chem. Ecol. 27: 27. Trematerra, P. 2002. Use of pheromones in integrated pest 2109–2124. management of stored-products, p. 1–4. In D. Pimentel (ed.), 19. Savoldelli, S., L. Su¨ss, and P. Trematerra. Mating disruption of Encyclopedia of pest management. Marcel Dekker, New York. Ephestia cautella (Walker) in chocolate factories. IOBC-WPRS Bull., 28. Trematerra, P. 2012. Advances in the use of pheromones for stored- in press. product protection. J. Pest Sci. 85:285–299. 20. Shani, A., and J. Clearwater. 2001. Evasion of mating disruption in 29. Trematerra, P., C. Athanassiou, V. Stejskal, A. Sciarretta, N. Ephestia cautella (Walker) by increased pheromone production Kavallieratos, and N. Palyvos. 2011. Large-scale mating disruption relative to that of undisrupted populations. J. Stored Prod. Res. 37: of Ephestia spp. and Plodia interpunctella in Czech Republic, 237–252. Greece, and Italy. J. Appl. Entomol. 135:749–762.