Efficacy of Pyriproxyfen for Control of Stored-Product Psocids (Psocoptera) on Concrete Surfaces

University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln

U.S. Department of Agriculture: Agricultural Publications from USDA-ARS / UNL Faculty Research Service, Lincoln, Nebraska

2011

Christos G. Athanassiou University of Thessaly, [email protected]

Frank H. Arthur USDA-ARS, [email protected]

Nickolas G. Kavallieratos Benaki Phytopathological Institute

James E. Throne USDA-ARS, Manhattan, KS, [email protected]

Follow this and additional works at: https://digitalcommons.unl.edu/usdaarsfacpub

Athanassiou, Christos G.; Arthur, Frank H.; Kavallieratos, Nickolas G.; and Throne, James E., "Efficacy of Pyriproxyfen for Control of Stored-Product Psocids (Psocoptera) on Concrete Surfaces" (2011). Publications from USDA-ARS / UNL Faculty. 2031. https://digitalcommons.unl.edu/usdaarsfacpub/2031

This Article is brought to you for free and open access by the U.S. Department of Agriculture: Agricultural Research Service, Lincoln, Nebraska at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Publications from USDA-ARS / UNL Faculty by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. STORED-PRODUCT Efﬁcacy of Pyriproxyfen for Control of Stored-Product Psocids (Psocoptera) on Concrete Surfaces

CHRISTOS G. ATHANASSIOU,1,2,3 FRANK H. ARTHUR,2 NICKOLAS G. KAVALLIERATOS,4 2 AND JAMES E. THRONE

J. Econ. Entomol. 104(5): 1765Ð1769 (2011); DOI: http://dx.doi.org/10.1603/EC10424 ABSTRACT The insect growth regulator pyriproxyfen was evaluated as a surface treatment for control of three stored-product psocid pests Liposcelis bostrychophila Badonnel, Liposcelis decolor (Pearman), and Liposcelis paeta Pearman (Psocoptera: Liposcelididae). Nymphs were exposed for 35 d on a concrete surface treated with 2.3 mg of active ingredient/m2 pyriproxyfen. Exposure to pyriproxyfen signiÞcantly reduced the numbers of both adults and nymphs in comparison with untreated controls. In adults, the greatest reduction (Ͼ90%) was for L. decolor and L. bostrychophila, whereas for L. paeta it was 49%. Few adults of any species were found in the pyriproxyfen treatments. The greatest numbers of nymphs were recorded for L. bostrychophila for both pyriproxyfen treatments and controls. Few adults of any species were found in the pyriproxyfen treatments. The results indicate that pyriproxyfen is effective for control of L. bostrychophila, L. decolor, and L. paeta on concrete, and although complete control was not achieved, the results warrant further long-term study to determine whether pyriproxyfen can completely eliminate psocid populations over time.

KEY WORDS pyriproxyfen, insect growth regulator, Liposcelis, surface treatment, Psocoptera

Stored-product psocids (Psocoptera) belonging to Insecticides that are registered to control stored- the genus Liposcelis are important emerging pests of product beetles should be evaluated because of the stored products. They feed on a wide variety of foods, increasing pest status of psocids. One such insecti- but they are particularly associated with amylaceous cide is the newly registered insect growth regulator commodities, such as grain and ßour (Nayak 2006, (IGR) pyriproxyfen, which is labeled for use as a Throne et al. 2006). One of the most important char- surface or spot treatment in food facilities in the acteristics of these psocids is their high tolerance to United States. Several studies have documented its many of the insecticides currently used for control of high efÞcacy against stored-product insect pests. stored-product beetles (Nayak et al. 1998). Moreover, Arthur et al. (2009) reported that pyriproxyfen was the bacterium-derived grain protectant spinosad was superior to hydroprene for control of several insect found to be ineffective against several Liposcelis spp. pests on wood, metal, and concrete surfaces, with a (Nayak et al. 2005, Athanassiou et al. 2009). Several considerable level of persistence. Kostyukovsky et organophosphate, carbamate, and pyrethroid insecti- al. (2000) reported that pyriproxyfen was able to cides have been evaluated for control of stored-prod- control pirimiphos-methyl-resistant Tribolium cas- uct psocids on concrete surfaces, but they failed to taneum (Herbst) (Coleoptera: Tenebrionidae) pop- provide long-term protection (Collins et al. 2000; ulations and that it was superior to methoprene for Nayak et al. 2002a,b, 2003). control of the rice weevil, Sitophilus oryzae (L.) (Coleoptera: Curculionidae), and the lesser grain Mention of trade names or commercial products in this publi- borer, Rhyzopertha dominica (F.) (Coleoptera: cation is solely for the purpose of providing speciÞc information Bostrychidae). Hubert et al. (2007) found that a and does not imply recommendation or endorsement by the U.S. combination of permethrin ϩ benzyl benzoate ϩ Department of Agriculture, the University of Thessaly or the Benaki Phytopathological Institute. pyriproxyfen was superior to chlorpyriphos, delta- 1 Laboratory of Entomology and Agricultural Zoology, Depart- methrin, beta-cyßuthrin, and a combination of del- ment of Agriculture, Crop Production and Rural Development, tamethrin ϩ bioallethrin for control of several University of Thessaly, Phytokou str., 38446, Nea Ionia Magnissia, Greece. stored-product mites. However, there are no data 2 USDAÐARS, Center for Grain and Animal Health Research, 1515 available on efÞcacy of pyriproxyfen for control of College Ave., Manhattan KS 66502-2736. stored-product psocids. The objective of our study 3 Corresponding author, e-mail: [email protected]. was to determine whether pyriproxyfen applied at 4 Laboratory of Agricultural Entomology, Department of Entomol- ogy and Agricultural Zoology, Benaki Phytopathological Institute, 8 the label rate would be an effective surface treat- Stefanou Delta str., 14561, KiÞssia, Attica, Greece. ment for control of psocids. 1766 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 104, no. 5

Table 1. Numbers of adults and nymphs in control and pyriproxyfen-treated arenas (individuals/arena ؎ SE) 35 d after application for each psocid species

L. decolor L. paeta L. bostrychophila Treatment Adults Nymphs Adults Nymphs Adults Nymphs Control 9.4 Ϯ 1.0a 6.9 Ϯ 1.5a 3.9 Ϯ 0.7a 4.7 Ϯ 1.7a 8.5 Ϯ 0.3a 60.1 Ϯ 5.3a Pyriproxyfen 0.9 Ϯ 0.2b 2.9 Ϯ 1.2b 2.0 Ϯ 0.4b 0.9 Ϯ 0.6b 0.6 Ϯ 0.3b 11.1 Ϯ 4.2b t Ϫ8.3 Ϫ2.1 Ϫ2.6 Ϫ2.1 Ϫ18.5 Ϫ7.3 P Ͻ0.01 0.02 0.01 0.02 Ͻ0.01 Ͻ0.01

Within each column, means followed by the same letter are not signiÞcantly different (df ϭ 70; two-tailed t-test).

Materials and Methods were sprayed with water as untreated controls. Ten psocids and Þve kernels of cracked wheat were Insects. Young nymphs (N ÐN ) of the psocids Li- 1 2 placed on each arena, either before or after spray- poscelis bostrychophila Badonnel, Liposcelis decolor ing. The 24 arenas were divided into four categories (Pearman), and Liposcelis paeta Pearman (Psocop- (six replicate dishes per category): 1) arenas that tera: Liposcelididae) were used in the tests. L. bostry- contained psocids before spraying, with the kernels chophila generally is parthenogenetic (males have added after spraying; 2) arenas that contained pso- only recently been discovered; Mockford and Krush- cids and kernels before spraying; 3) arenas that did elnycky 2008), whereas the other two species repro- not contain psocids before spraying (the psocids duce after mating. Psocids were reared on a mixture of were added immediately after spraying) but con- 97% cracked wheat kernels, 2% crisp rice, and 1% tained Þve cracked kernels of wheat; and 4) arenas brewerÕs yeast at 30ЊC and 70% RH. To obtain indi- that did not contain psocids or kernels before spray- viduals of standardized age for experiments, 30 female ing (the psocids and the kernels were added imme- adults were left to oviposit in 35-mm petri dishes diately after spraying). containing red-colored psocid diet for 3 d, and then This procedure was replicated three times, with the females were removed. new arenas each time (3 by 24 ϭ 72 dishes for each After an additional 7 d, the newly emerged nymphs species). After spraying, all arenas were placed in were removed with a Þne brush (Opit and Throne black plastic boxes containing saturated sodium chlo- 2008). The colored diet was prepared by mixing 100 g ride solution below a false ßoor to maintain the rela- of crisp rice with a solution of 5 ml of red food dye tive humidity at Ϸ75%. The boxes were placed in (Global ChemSources, Inc., Cedar Grove, NJ) in 300 incubators maintained at 30ЊC and 70% RH. All arenas ml of water. Afterward, the mixture was dried in a were checked after 35 d of exposure, and the numbers mechanical convection oven (HTM 85, Precision Sci- of adults and nymphs were recorded separately for entiÞc, Inc., Chicago, IL) for 2 d, and then the dried each arena. mixture that resulted was grinded in a Wiley Mill by Data Analysis. For each species, the numbers of using a #20 sieve (0.85-mm openings; ScientiÞc Ap- adults and nymphs in the treated concrete arenas were paratus, Philadelphia, PA). compared with those in the respective control dishes Insecticide. A formulation of pyriproxyfen (Ny- using a two-tailed t-test with n-2 df to indicate whether Guard, McLaughlin Gormley King [MGK] Company, pyriproxyfen affected the number of psocids. Then, Minneapolis, MN) that contains 10% active ingredient for each species and life stage, the data from the (AI) was used in the experiments at the label rate of pyriproxyfen-treated dishes were analyzed using a 2.3 mg (AI)/m2. one-way analysis of variance (ANOVA) to determine Bioassays. All tests were conducted in petri dishes differences in number of psocids among the four treat- (9 cm in diameter by 1.5 cm in height), which had ment categories with JMP software (Sall et al. 2001). a surface area of Ϸ62 cm2. The bottoms of the dishes Means were separated by the honestly signiÞcant dif- were covered with driveway patching material ference (HSD) test at ␣ ϭ 0.05 (Sokal and Rohlf 1995). (Rockite, Hartline Products Co., Inc., Cleveland, OH) to create the concrete surface (hereby re- ferred to as arenas). The arenas were prepared 1Ð2 Results d before the tests. The internal sides of the arenas were coated with Fluon (polytetraßuoroethylene, Adults. Numbers of adults were signiÞcantly re- Northern Products, Woonsocket, RI) to prevent duced for all species in the pyriproxyfen-treated are- escape of psocids. The solutions of pyriproxyfen nas compared with the numbers of adults in the con- were prepared with distilled water. Dishes were trols (Table 1). The greatest reduction (Ͼ90%) was sprayed with a Badger 100 artistÕs airbrush (Badger for L. decolor and L. bostrychophila. For these species, Corporation, Franklin Park, IL) to treat each indi- less than one adult per arena was recorded. The num- vidual concrete arena with 0.25 ml of formulated ber of L. paeta adults was reduced by only 49%. Sig- solution per 62 cm2 area of the arena, which is niÞcantly more L. decolor adults were recorded in equivalent to the label spray rate 3.8 l/94 m2. Twen- arenas in which the kernels were treated Þrst and the ty-four arenas were prepared for each psocid spe- nymphs added afterward in comparison with the are- cies: 12 were sprayed with pyriproxyfen, and 12 nas in which the psocids were treated Þrst and the October 2011 ATHANASSIOU ET AL.: PYRIPROXYFEN FOR CONTROL OF PSOCIDS 1767

Table 2. Numbers of L. decolor adults and nymphs in py- Table 4. Numbers of L. bostrychophila adults and nymphs in riproxyfen-treated arenas (individuals/arena ؎ SE) 35 d after pyriproxyfen-treated arenas (individuals/arena ؎ SE) 35 d after spraying spraying

Treatment Adults Nymphs Treatment Adults Nymphs Without psocids, without kernels 0.8 Ϯ 0.3ab 0.2 Ϯ 0.1a Without psocids, without kernels 2.0 Ϯ 0.8a 30.0 Ϯ 10.5a Without psocids, with kernels 2.1 Ϯ 0.8b 9.4 Ϯ 4.2b Without psocids, with kernels 0.0 Ϯ 0.0b 3.4 Ϯ 3.0bc With psocids, without kernels 0.4 Ϯ 0.2a 1.2 Ϯ 0.6a With psocids, without kernels 0.0 Ϯ 0.0b 0.1 Ϯ 0.1c With psocids, with kernels 0.3 Ϯ 0.2a 0.6 Ϯ 0.4a With psocids, with kernels 0.4 Ϯ 0.4ab 10.7 Ϯ 7.7ab F 4.1 4.2 F 4.3 3.0 P 0.01 0.01 P 0.01 0.04

Within each column, means followed by the same letter are not Within each column, means followed by the same letter are not signiÞcantly different (df ϭ 3, 32; HSD test at 0.05; “without” or “with” signiÞcantly different (df ϭ 3, 32; HSD test at 0.05; “without” or “with” indicate absence or presence during spraying, respectively). indicate absence or presence during spraying, respectively). kernels added afterward or with arenas which con- that pyriproxyfen can reduce numbers of psocids on tained psocids and kernels before spraying (Table 2). concrete but that the level of reduction is mainly For L. paeta and L. bostrychophila, signiÞcantly higher dependent on the target species. There have been numbers of adults were in arenas that did not contain several studies on another IGR, methoprene, for con- psocids or kernels before spraying in comparison with trol of psocids. In a recent study, Athanassiou et al. the arenas in which the kernels were treated Þrst and (2010b) found that methoprene was not able to con- the psocids added afterward or vice versa (Tables 3 trol the psocids L. decolor, L. bostrychophila, Liposcelis and 4). entomophila (Enderlein), and L. paeta on maize (Zea Nymphs. The numbers of nymphs in the controls mays L.), rice (Oryza sativa L.), and wheat (Triticum were signiÞcantly greater than the numbers of nymphs aestivum L.). Also, the combination of methoprene in the pyriproxyfen-treated dishes (Table 1). For both with spinosad was not effective for control of L. bostry- pyriproxyfen-treated and control arenas, the greatest chophila on wheat (Athanassiou et al. 2010a). These number of nymphs was recorded for L. bostrychophila. studies and earlier studies indicate lack of efÞcacy of SigniÞcantly more L. decolor nymphs were recorded methoprene for control of Liposcelis spp. (Nayak et al. in arenas that contained only kernels before spraying, 1998, 2002b). Bucci (1994) reported that L. bostry- in comparison with other treatments (Table 2). In chophila was extremely tolerant to the IGR fenoxy- contrast, no signiÞcant differences were noted in carb, even at very high doses. These studies and our numbers of L. paeta nymphs among the four treat- current Þndings indicate that psocids generally are ments (Table 3), and nymphs were found only in quite tolerant to IGRs, including pyriproxyfen, al- arenas that contained psocids and kernels before though pyriproxifen is the most effective of IGRs eval- spraying. SigniÞcantly more L. bostrychophila nymphs uated thus far. were in arenas that did not contain psocids or kernels Access to food can moderate the efÞcacy of a given before spraying in comparison with the arenas in insecticide (Arthur 1998, 2009; Arthur et al. 2009). which the kernels were treated Þrst and the psocids Arthur (2009) noted that survival of T. castaneum added afterwards or vice versa (Table 4). adults on concrete treated with chlorfenapyr in- creased when ßour was present. Consequently, sani- tation (presence or absence of food material) is an Discussion important variable that should be incorporated into This is the Þrst study in which the IGR pyriproxyfen evaluation of insecticides for stored-product pests. In was evaluated on concrete surfaces for control of previous studies, food was usually added after the stored-product psocids, and this is also one of the few application of the insecticide; thus, food was not studies that examined this insecticide for control of treated. This may have allowed insects to moderate stored-product insects in general. Our results indicate the toxic effects of the insecticides through feeding on untreated food. In our study, both cases were examined: insecticide-treated and insecticide-free food. Table 3. Numbers of L. paeta adults and nymphs in pyriproxy- There were no general trends in survival for either of ؎ fen-treated arenas (individuals/arena SE) 35 d after spraying food categories for all three species. We hypothesize Treatment Adults Nymphs that the presence of cracked kernels in all dishes might have reduced pyriproxyfen efÞcacy, although treat- Without psocids, without kernels 4.1 Ϯ 1.2a 0.0 Ϯ 0.0a ments without kernels were not evaluated in this Without psocids, with kernels 1.1 Ϯ 0.6bc 0.0 Ϯ 0.0a With psocids, without kernels 0.0 Ϯ 0.0c 0.0 Ϯ 0.0a study. These results suggest that removal of insecti- With psocids, with kernels 1.9 Ϯ 0.6ab 3.7 Ϯ 2.5a cide-treated food residues in a milling or warehouse F 6.2 2.3 facility may be just as important for pest control as P Ͻ0.01 0.10 removal of untreated food residues. Differences among psocid species in their suscep- Within each column, means followed by the same letter are not signiÞcantly different (df ϭ 3, 32; HSD test at 0.05; “without” or “with” tibility to insecticides applied on concrete is an addi- indicate absence or presence during spraying, respectively). tional parameter that should be taken into consider- 1768 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 104, no. 5 ation when evaluating insecticides. Among the species activity of pyriproxyfen can gradually eliminate the tested here, L. paeta is considered to be tolerant to psocid population. several insecticides, such as chlorpyriphos-methyl, In conclusion, pyriproxyfen showed activity against pirimiphos-methyl, and fenitrothion, whereas L. L. decolor, L. paeta, and L. bostrychophila on concrete, bostrychophila is considered susceptible (Collins et al. a characteristic that should be further evaluated to 2000, Nayak et al. 2002a). In our study, there were determine whether pyriproxyfen can eliminate psocid considerably greater numbers of L. bostrychophila populations over time. In addition, the combined ap- nymphs, in both treated and untreated dishes, in com- plication of pyriproxyfen with a reduced-risk contact parison with the other two species. For example, the neurotoxic insecticide, such as chlorfenapyr (Guedes number of L. bostrychophila nymphs was 13 and 7 et al. 2008, Arthur 2009), may further increase its times higher than the numbers of L. paeta and L. efÞcacy. decolor nymphs, respectively. In contrast with the other two species, L. bostrychophila typically repro- duces parthenogenetically, with high potential for Acknowledgments production of progeny (Wang et al. 2000). Athanas- siou et al. (2010c) found that L. bostrychophila adult We thank Ann Redmon and Ngunza Kisangani for tech- females generally produced considerably more F1 nical assistance, MGK for the pyriproxyfen sample, and progeny than L. paeta or L. decolor in a wide variety Manoj Nayak for comments on an earlier version of this of grain commodities. Even in insecticide-treated manuscript. wheat and rice, Athanassiou et al. (2009) reported that L. bostrychophila produced greater numbers of off- spring than L. paeta. Greater capacity for progeny References Cited production may counteract the insecticidal effect, es- Arthur, F. H. 1998. Effects of food source on red ßour beetle pecially in the case of IGRs, where this effect is pri- (Coleoptera: Tenebrionidae) survival after exposure on marily focused on the immature life stages. Neverthe- concrete treated with cyßuthrin. J. Econ. Entomol. 91: less, pyriproxyfen reduced L. bostrychophila numbers 773Ð778. in comparison with the control, but the level of re- Arthur, F. H. 2009. EfÞcacy of chlorfenapyr against adult duction varied according to the treatment. In a pre- Tribolium castaneum exposed on concrete: effects of ex- vious study on wheat, rice, and maize, methoprene did posure interval, concentration and the presence of a food not effectively control L. bostrychophila, L. decolor, or source after exposure. Insect Sci. 16: 157Ð163. Arthur, F. H., S. Liu, B. Zhao, and T. W. Phillips. 2009. L. paeta, although that progeny production was re- Residual efÞcacy of pyriproxyfen and hydroprene ap- duced for all species. plied to wood, metal and concrete for control of stored Few adults of any species were found at the end of product insects. Pest Manag. Sci. 65: 791Ð797. the 35-d exposure interval. It is well documented that Athanassiou, C. G., F. H. Arthur, and J. E. Throne. 2009. 35 d is a sufÞcient interval for young psocid nymphs to EfÞcacy of grain protectants against four psocid species reach the adult stage at the conditions examined here on maize, rice and wheat. Pest Manag. Sci. 65: 1140Ð1146. (Rees and Walker 1990, Leong and Ho 1995, Wang et Athanassiou, C. G., F. H. Arthur, N. G. Kavallieratos, and J. E. al. 2000, Opit and Throne 2008). Therefore, we can Throne. 2010a. EfÞcacy of spinosad and methoprene, conclude that there is an effect of pyriproxyfen on the applied alone or in combination, against six stored-product insect species. J. Pest Sci. 84: 61Ð67. species tested. This was evident even in the case of L. Athanassiou, C. G., F. H. Arthur, and J. E. Throne. 2010b. bostrychophila where the greatest number of nymphs EfÞcacy of methoprene for control of Þve species of was recorded on the treated substrates, yet the lowest psocids (Psocoptera) on wheat, rice, and maize. J. Food number of adults was found. The exposure interval Prot. 73: 2244Ð2249. was selected to examine whether the exposed nymphs Athanassiou, C. G., G. P. Opit, and J. E. Throne. 2010c. would reach the adult stage and the capacity of these Inßuence of commodity type, percentage of cracked ker- adults for progeny production. However, we do not nels, and wheat class on population growth of stored- know whether the nymphs found on the pyriproxyfen- product psocids (Psocoptera: Liposcelidae). J. Econ. En- treated surfaces had been arrested at that stage and tomol. 103: 985Ð990. Bucci, R. 1994. Effects of two insect growth regulators on were unable to become adults. For Plodia interpunc- the booklouse Liposcelis bostrychophila. J. Stored Prod. tella (Hu¨ bner) (Lepidoptera: Pyralidae), Mohandass Res. 30: 157Ð161. et al. (2006) found that hydroprene prolonged imma- Collins, P. J., M. K. Nayak, and R. Kopittke. 2000. Residual ture development or caused molting failure. Arthur et efÞcacy of four organophosphate insecticides on con- al. (2009) found that pyriproxyfen reduced adult crete and galvanized steel against three liposcelid psocid emergence of the cigarette beetle, Lasioderma serri- species (Psocoptera: Liposcelidae) infesting stored prod- corne (F.) (Coleoptera: Anobiidae); the confused ucts. J. Econ. Entomol. 93: 1357Ð1363. ßour beetle, Tribolium confusum Jacquelin du Val Guedes, R.N.C., J. F. Campbell, F. H. Arthur, G. P. Opit, K. Y. (Coleoptera: Tenebrionidae); the sawtoothed grain Zhu, and J. E. Throne. 2008. Acute lethal and behavioral sublethal responses of two stored-product psocids to sur- beetle, Oryzaephilus surinamensis (L.) (Coleoptera: face insecticides. Pest Manag. Sci. 64: 1314Ð1322. Silvanidae); T. castaneum; and P. interpunctella after Hubert, J., V. Stejskal, Z. Munzbergova, J. Hajslova, and F. H. 56 d of exposure, whereas, at the same conditions, Arthur. 2007. Toxicity and efÞcacy of selected pesti- hydroprene lost its effectiveness after 28 d. Testing cides and new acaricides to stored product mites (Acari: longer exposure intervals would indicate whether the Acaridida). Exp. Appl. Acarol. 42: 283Ð290. October 2011 ATHANASSIOU ET AL.: PYRIPROXYFEN FOR CONTROL OF PSOCIDS 1769

Kostyukovsky, M., B. Chen, S. Atsmi, and E. Shaaya. 2000. Liposcelididae), a pest of stored products. Pest Manag. Biological activity of two juvenoids and two ecdysteroids Sci 58: 1223Ð1228. against three stored product insects. Insect Biochem. Nayak, M. K., P. J. Collins, and R. A. Kopittke. 2003. Residual Mol. Biol. 30: 891Ð897. toxicities and persistence of organophosphorus insecti- Leong, E.C.W., and S. H. Ho. 1995. Life cycle of Liposcelis cides mixed with carbaryl as structural treatments against entomophila (Psocoptera: Liposcelididae) and a culturing three liposcelidid psocid species (Psocoptera: Liposceli- regime for liposcelidids. Bull. Entomol. Res. 85: 501Ð506. didae) infesting stored grain. J. Stored Prod. Res. 39: Mockford, E. L., and P. D. Krushelnycky. 2008. New species 343Ð353. and records of Liposcelis Motschulsky (Psocoptera: Li- Nayak, M. K., G. J. Daglish, and V. S. Byrne. 2005. Effec- poscelididae) from Hawaii with Þrst description of the tiveness of spinosad as a grain protectant against resistant male of Liposcelis bostrychophila Badonnel. Zootaxa 1766: beetle and psocid pests of stored grain in Australia. J. 53Ð68. Stored Prod. Res. 41: 455Ð467. Mohandass, S., F. H. Arthur, K. Y. Zhu, and J. E. Throne. Opit, G. P., and J. E. Throne. 2008. Population growth and 2006. Hydroprene prolongs developmental time and in- development of psocid Lepinotus reticulatus at constant creases mortality in wandering-phase Indianmeal moth temperatures and relative humidities. J. Econ. Entomol. (Lepidoptera: Pyralidae) larvae. J. Econ. Entomol. 99: 101: 605Ð615. 1509Ð1519. Rees, D. P., and A. J. Walker. 1990. The effect of temperature and relative humidity on population growth of three Nayak, M. K. 2006. Psocid and mite pests of stored com- Liposcelis species infesting stored products in tropical modities: small but formidable enemies, pp. 1061Ð1073. countries. Bull. Entomol. Res. 80: 353Ð358. In I. Lorini, B. Bacaltchuk, H. Beckel, D. Deckers, E. Sall, J. E., A. Lehman, and L. Creighton. 2001. JMP Start Sundfeld, J. P. dos Santos, J. D. Biagi, J. C. Celaro, L. R. Statistics. A guide to statistics and data analysis using JMP DÕA. Faroni, L. de O. F. Bortolini et al. (eds.), Pro- and JMP IN software. Duxbury Press, Belmont, CA. ceedings of the 9th International Working Conference Sokal, R. R., and F. J. Rohlf. 1995. Biometry, 3rd ed. W. H. on Stored-Product Protection, 15Ð18 October 2006, Freeman and Company, New York. Campinas, Sao Paulo, Brazil. Brazilian Post-harvest As- Throne, J. E., G. P. Opit, and P. W. Flinn. 2006. Seasonal sociation-ABRAPOS, Passo Fundo, RS, Brazil. distribution of psocids in stored wheat, pp. 1095Ð1103. Nayak, M. K., P. J. Collins, and R. S. Reid. 1998. EfÞcacy of In I. Lorini, B. Bacaltchuk, H. Beckel, D. Deckers, E. grain protectants and phosphine against Liposcelis bostry- Sundfeld, J. P. dos Santos, J. D. Biagi, J. C. Celaro, L. R. chophila, L. entomophila and L. paeta (Psocoptera: Lipos- DÕA. Faroni, L. de O. F. Bortolini, et al. (eds.), Pro- celidae). J. Econ. Entomol. 91: 1208Ð1212. ceedings of the 9th International Working Conference Nayak, M. K., P. J. Collins, and R. A. Kopittke. 2002a. Com- on Stored-Product Protection, 15Ð18 October 2006, parative residual toxicities of carbaryl, deltamethrin and Campinas, Sao Paulo, Brazil. Brazilian Post-harvest As- permethrin as structural treatments against three lipos- sociation-ABRAPOS, Passo Fundo, RS, Brazil. celidid psocid species (Psocoptera: Liposcelididae) in- Wang, J. J., J. H. Tsai, Z. H. Zhao, and L. S. Li. 2000. De- festing stored commodities. J. Stored Prod. Res. 38: 247Ð velopment and reproduction of the psocid Liposcelis 258. bostrychophila (Psocoptera: Liposcelididae) as a function Nayak, M. K., P. J. Collins, and H. Pavic. 2002b. Long-term of temperature. Ann. Entomol. Soc. Am. 93: 261Ð270. effectiveness of grain protectants and structural treatments against Liposcelis decolor (Pearman) (Psocoptera: Received 20 November 2010; accepted 30 April 2011.