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2 Author's personl opy Journl of Stored Produts Reserh 44 (28) Life history prmeters of Lsioderm serriorne (F.) s influened y food soures Rizn M. Mhroof, Thoms W. Phillips 1 Deprtment of Entomology nd Plnt Pthology, 127 Nole Reserh Center, Oklhom Stte University, Stillwter, OK 7478, USA Aepted 24 Deemer 27 Astrt Feundity, egg to dult survivl rte, developmentl time, nd dult ody weight of the igrette eetle, Lsioderm serriorne (Coleopter: Anoiide) were evluted on seven food soures t 28 1C. Ground hili, pprik, yenne pepper, hewing lef too, igr too, ommeril inset it referred to s NOW it, nd whet flour were used to evlute men lifetime feundity. The highest feundity ( eggs/femle) ws oserved in whet flour, wheres the lowest feundity ( eggs/femle) ws oserved in igr too. Among the seven food soures, eetles rered in whet flour showed the highest survivl rte of %. Only 15% of the eggs lid in NOW it developed to the dult stge. In the three food soures ontining Cpsium spp. the survivl rte rnged from 3% to 4%. The egg, lrvl, nd pupl development times vried from 3 to 5, 38 to 92 nd 4 to 18 d, respetively, mong food soures. Body weight nd dult longevity studies showed tht the hevier dults lso hd the longest life spn. Ovipositing femle L. serriorne pper to disriminte mong different food soures. Although L. serriorne lid eggs in ll food soures evluted, lrvl nd pupl survivl were lowest in NOW it. Informtion on the iology nd host use pttern of L. serriorne my help to explin how vrious stored ommodities re ffeted y this speies nd my led to develop pproprite pest mngement strtegies for this inset pest. Pulished y Elsevier Ltd. Keywords: Lsioderm serriorne; Cigrette eetle; Development; Survivl; Feundity 1. Introdution The igrette eetle, Lsioderm serriorne (F.) (Coleopter: Anoiide) is known to suessfully reed nd develop on vriety of grin-sed produts, spies, nd too, nd infest these ommodities during storge, mnufturing nd t the retil level (Howe, 1957; Bry et l., 1974; Allotey, 1988; Jo, 1992; Ashworth, 1993; Dimetry et l., 24). Lrvl feeding uses most of the dmge to stored ommodities (Minor, 1979) while dults re known to use dmge y utting holes to penetrte or espe from pkged ommodities (Highlnd, 1991). In ddition to feeding dmge, the presene of ded insets, Corresponding uthor. Current ddress: Deprtment of Biologil Sienes, South Crolin Stte University, 3, College Street NE, Orngeurg, SC 29117, USA. Tel.: ; fx: E-mil ddress: rmhroof@ssu.edu (R.M. Mhroof). 1 Current ddress: Deprtment of Entomology, 123 W. Wters Hll, Knss Stte University, Mnhttn, KS 6652, USA. residues from different life stges suh s st skins or pupl ses, nd frss eome ontminnts in ommodities nd render them undesirle for humn onsumption. The infesttion of ommodity is usully initited y dispersing grvid femles whih tively seek suitle hosts for oviposition nd penetrte through the pkging mterils of ommodities to olonize new food resoures (Howe, 1957). Biology nd life history prmeters of L. serriorne hve een investigted in the pst under vrious temperture nd reltive humidity (r.h.) onditions, generlly using too, yest, or whet s food soures (Jones, 1913; Powell, 1931; Howe, 1957; Lefkovith nd Currie, 1967). Jones (1913) reported tht the verge time for the egg development ws 6 d, the lrvl stge ws 5 d, nd the pupl stge ws 12.5 d, when too served s lrvl food. Powell (1931) determined the length of the life yle of L. serriorne under severl ontrolled temperture nd humidity onditions using too or yest s food X/$ - see front mtter Pulished y Elsevier Ltd. doi:1.116/j.jspr

3 Author's personl opy 22 R.M. Mhroof, T.W. Phillips / Journl of Stored Produts Reserh 44 (28) soures. Completion of the life yle required typilly 18 2 d longer in too thn in yest, for exmple, t 28 1C nd 75% r.h. the development time ws 36 d in yest nd 55 d in too. Howe (1957) studied the developmentl time with rnge of tempertures tht vried from 2. to C in inrements of 2.5 1C, nd t rnge of r.h. tht vried from 2% to 1% with inrements of 1%, using whet s lrvl food. Under different temperture nd r.h. onditions, development times vried from 5.3 to 2.4 d for eggs 18.2 to 11 d for lrve, 6.4 to 25.9 d for pupe, nd 18 to 46 d for dults. Lefkovith nd Currie (1967) showed tht t 3 1C nd 6% r.h., the feundity nd survivl of L. serriorne dults were dependent upon their ody weight when newly emerged. Lsioderm serriorne is well known s stored-produt inset tht n reed nd develop in vriety of hemilly defended food soures suh s spies nd too (Dowd nd Shen, 199; Jo, 1992; Dimetry et l., 24). LeCto (1978) reported the time required for development of L. serriorne in 19 different spies nd the fresh ody weights of eetles tht emerged from eh spie. Among the 19 spies evluted, yenne pepper (Cpsium nnuum L. vr. glriusulum) nd pprik (C. nnuum L.) were the most fvorle diets for the development of L. serriorne. Dimetry et l. (24) ompred six spies nd onluded tht orinder seeds (Corinder stivum L.) were the most suitle host for oviposition. In reent study to evlute the vrition in responses of mle nd femle L. serriorne to vrious food soures, produts ontining Cpsium spp., two vrieties of too, whet, nd proessed lmond eliited signifintly higher ttrtive responses y femle L. serriorne thn mle eetles (Mhroof nd Phillips, 27). The urrent study follows from Mhroof nd Phillips (27) in whih we evluted voltiles from food soures for their effet on orienttion of dult L. serriorne. We found vrition in responses y femles ompred to mles, long with vrition in responses mong foods, nd thus hypothesized tht some food soures my e etter for survivl nd development of L. serriorne thn others. The following three mjor ojetives were then developed to evlute the life history prmeters of L. serriorne in seleted food soures. (1) To determine the oviposition preferene nd egg-to-dult survivl rtes of L. serriorne in food soures previously shown to ontin hemil ttrtnts for L. serriorne dults; (2) to exmine the developmentl time for vrious life stges of L. serriorne in seleted food soures; nd (3) to reord the longevity nd dult ody weight of L. serriorne rered on vrious food soures. 2. Mterils nd methods 2.1. Lsioderm serriorne olony Adult L. serriorne were tken from lortory olonies mintined sine 25 t the Stored Produts Reserh Lortory in Oklhom Stte University, Deprtment of Entomology nd Plnt Pthology (Stillwter, OK, USA). Four to six dys fter emergene, dults were olleted from the olony, were introdued to lehed whet flour in 94-ml glss Mson jrs nd mintined t 28 1C nd 6% r.h. After 2 d, the ontents of the jrs were sifted using two sieves, 85-mm mesh (USA No. 2) (Seeduro Equipment Co., Chigo, IL) nd 212-mm mesh (USA No. 7; Fisher Sientifi, Mdison, WI) to ollet the eggs. Colleted eggs were rered on medium of 5% dried ground rewer s yest nd 95% lehed whet flour t 28 1C, 6% r.h. nd 16:8 (L:D) h photoperiod Food soures Seven tegories of food soures to whih L. serriorne were previously strongly ttrted s mted femles (Mhroof nd Phillips, 27) were seleted to evlute different life history prmeters. Food mterils were from group of eptle host mterils inluding Cpsiumderived produts, ured too, n lmond-derived produt, nd grin. Produts ontining Cpsium spp. inluded in the experiment were (1) dried red hili (Cpsium frutesens L.) (Bolner s Fiest Produt In., Sn Antonio, TX), (2) pprik (Adms Extrt nd Spie LLC, Gonzles, TX), nd (3) yenne pepper (Ztrins, New Orlens, LA). Two types of ured nd proessed too tested in the experiment were (4) hewing lef too (Niotin tum L.; Tylor s Brothers, Winston Slem, NC) nd (5) mild nturl igr too (Bkwoods, Fort Luderdle, FL). Commerilly ville (6) nvel orngeworm, Amyelois trsitell (Wlker) it omposed of pressed lmond (Prunus dulis (Mill.) D.A.We) flkes nd idulted lmond oil (Tre é IV NOW it, Tréé Inorported, CA; herefter referred to s NOW; the rtio for different ingredients in NOW is proprietry informtion), nd (7) lehed whet flour (Tritium estivum L.) were the two other food soures inluded in the study. Red hili, hewing lef too, nd igr too were pulverized using mehnil grinder. The three pulverized mterils, pprik, yenne pepper, NOW nd whet flour were sifted using 425-mm meshed sieve (U.S. No. 4, Seeduro Equipment Co., Chigo, IL). The moisture ontent of ll powdered mterils used t the eginning of the study vried from 9% to 11% on dry weight sis Feundity, egg-to-dult survivl, nd lrvl survivl Vrition in the lifetime feundity in L. serriorne femles ws ompred in the seven food soures. Mles nd femles were first seprted sed on the externl genitl hrteristis of the pupe (Hlsted, 1963). To omplish this, 3 4-d old pupe were tken from the olony, the protetive pupl se ws refully removed, nd pupe were oserved under mirosope for the shpe of the genitl ppille. Mle nd femle pupe were then

4 Author's personl opy R.M. Mhroof, T.W. Phillips / Journl of Stored Produts Reserh 44 (28) kept in seprte jrs without ny food soure, t 28 1C, 6% r.h. nd 16:8 (L:D) h photoperiod until dult emergene. Soon fter dult emergene ( 24 h) mle eetles were mrked using titnium white ryli rtisti olor (Liquitex high visosity, Binney nd Smith, Winfield, KS) to distinguish mles from femles when mixed together. A minute mount of olor ws pled externlly on the elytr of the mle eetle nd llowed to dry for 2 min. Mrked mle eetles were trnsferred to the jrs ontining femle eetles nd mintined t 28 1C, 6% r.h. nd 16:8 (L:D) h photoperiod for totl period of 3 d to otin mted femles. Two grm quntities of eh food soure were trnsferred to glss shell vils fitted with plugs (14 45 mm; Fisherrnd Pittsurgh, PA). The plugs were puntured with shrp needle to form holes to filitte ir movement. A single femle from the mixed olony ws introdued into eh of the shell vils ontining one of the food soures. Femles were llowed to ly eggs for 3 d in the shell vils. Every third dy the femle ws refully trnsferred to new vil ontining 2 g of fresh food soure. Eggs lid in the medium were ounted under stereomirosope (Crl Zeiss In., Oerkohen, West Germny). Vils ontining whet flour were emptied into two sieves; 425-mm meshed sieve on top nd 212-mm meshed sieve on ottom (U.S. Nos. 4 nd 7, respetively), nd eggs olleted on the ottom sieve were oserved under the mirosope. Egg ounts were ontinued for eh femle until she died. The experiment ws onduted for seven food soures with totl of 35 shell vils tht represented five shell vils for eh food soure. The entire experiment ws replited four different times. Egg-to-dult survivl of L. serriorne ws studied in the seven seleted food soures. Eggs (24-h-old) olleted under the mirosope from dult eetles ged 4 5-d-old were used in the study. Five eggs were introdued to shell vil ontining 5 g of food nd were rered to the dult stge t 28 1C nd 6% r.h. Egg-to-dult survivl ws lulted from the numer of dults tht emerged out of five eggs. The experiment ws onduted for seven food soures with totl of 35 shell vils tht represented five shell vils for eh food soure. The entire experiment ws replited four times. Lrvl survivl ws studied y introduing five eggs trnsferred to shell vil ontining one of the vrious food soures. The methods used in this experiment were the sme s those desried for egg-to-dult survivl, exept tht during the study of lrve the shell vils were heked periodilly to oserve the lrve. Contents of the vil were emptied onto lk filter pper nd oserved for live nd moving lrve. Lrve tht did not move when stimulted with mel hir rush or tht were immoile nd disolored were onsidered ded. Contents were trnsferred k to the vil nd mintined t 28 1C nd 6% r.h, nd oserved for lrvl mortlity t intervls of 5 d. Oservtions were ontinued until ll lrve strted to onstrut pupl se nd were onfined inside the ses. Survivl of lrve ws sed on the numer of lrve ded out of the totl eggs hthed. The lrvl survivl study ws onduted for ll seven food soures with totl of 35 shell vils t time tht represented five shell vils for eh food soure. The entire experiment ws replited four different times Developmentl time for different life stges Egg development, lrvl development, pupl period nd the totl developmentl time for L. serriorne were evluted. Two grms of powdered food mterils were trnsferred to eh shell vil. One-dy-old eggs were olleted from the lortory olony. A single egg ws trnsferred to the stiky re of retngulr lk filter pper (.5 m wide nd 4 m long) folded into n L shpe, with piee of doule stiky tpe in one end. The filter pper ws slowly inserted into the vil ontining food soure. The vils were mintined t 28 1C nd 6% r.h. Vils were oserved every dy until egg hth, nd dys required for egg development ws reorded. After elosion, the first instr ws refully removed from the filter pper nd dropped into the medium, while the lk filter pper ws permnently removed from the vil. The lrvl developmentl period ws ounted from the dy of emergene of the first instr to the dy on whih the lrv onstruted the protetive se to initite puption, nd ws reorded y oserving the shell vils every other dy. In Cpsium produts lrve did not onstrut protetive pupl se nd under suh irumstnes morphologil differenes were oserved to identify the initition of puption. The pupl development period ws ounted from the time the lrv onstruted protetive se, or when detetle hnges oured in lrvl to pupl morphology to the emergene of the dult. Oservtions were mde y emptying the ontents of the vil onto piee of lk filter pper nd looking t the ontents under mirosope. Bsed on the time required for egg-to-dult development, the totl developmentl time ws lulted. The study ws onduted totl of four times, eh ohort with seven food soures. In ohort totl of 35 vils were oserved tht represented five vils per food soure. The numer of lrve nd pupe in eh ohort vried depending on the survivl of erlier stges Adult longevity nd ody weight Single eggs (24-h-old) were introdued to eh food soure nd the dults were llowed to emerge, then 2 g of food mterils were provided to individul dults in shell vil nd vils were oserved dily. The numer of dys tken from the first dy of emergene to the nturl mortlity of eh dult L. serriorne ws reorded. The study ws onduted for ll seven food soures with totl of 35 shell vils t time tht represented five shell vils for eh food soure. The entire study ws replited four different times. Adults tht emerged from this experiment

5 Author's personl opy 222 R.M. Mhroof, T.W. Phillips / Journl of Stored Produts Reserh 44 (28) nd from the developmentl time studies were olleted to hek oth longevity nd ody weight. One dults were ded they were pled in refrigertor for few hours, nd were then dried in n oven t 6 1C for 48 5 h until onstnt dry weight ws otined. Dry weight ws mesured using n eletroni mirolne type M3P (Srtorius Instruments, MGw Prk, IL; ury.1 mg) nd the dry weight of eh individul dult ws reorded Sttistil nlyses All experiments were onduted s rndomized omplete lok design, with food soures serving s tretment nd replitions serving s loks. Tretments were onsidered s fixed effet in the nlyses nd loks were onsidered s rndom effet. Prior to dt nlyses, numer of eggs, egg development time, lrvl development time, pupl development period, totl developmentl time, the dult longevity, nd ody weight dt were trnsformed to log 1 (X+1) sle to stisfy the ssumption of normlity nd homogeneity of vrines (Zr, 1984). Eggto-dult survivl nd lrvl survivl dt were onverted into perentge of totl eggs per vil nd these perentge vlues were ompred mong different food soures. Perentge egg-to-dult survivl nd lrvl survivl dt were trnsformed to rsine X.5 (Zr, 1984) prior to nlysis. All dt were sujeted to nlysis of vrine (ANOVA) y using the MIXED Proedure of SAS (SAS Institute, 23) to determine the signifint differenes etween tretments. LS mens were used for seprting tretment mens t ¼.5 level. Atul mens nd SEs re presented in the text, tle, nd figures. 3. Results 3.1. Feundity, egg-to-dult survivl, nd lrvl mortlity Newly deposited eggs were trnsluent, olong, nd were deposited s individul eggs etween the glss wll nd the food soure. Most of the eggs were lid during the first 9 1 d in ll food soures. The verge numers of eggs lid per femle were signifintly different mong the food soures (F ¼ 38.5; df ¼ 6, 124; Po.1). Signifintly more eggs ( ) were lid in whet flour when ompred to other food soures (Fig. 1A). For the Cpsium spp., we found signifintly more (Pp.5) eggs were lid on pprik ompred to yenne pepper nd hili. The lest eptle food soure mong the seven food mterils tested, sed on oviposition, ws the igr too (Fig. 1A). Lsioderm serriorne were le to survive from egg to dult stge on ll food soures tested. However, there were signifint differenes (F ¼ 22.6; df ¼ 6, 13; Po.1) in men perentge egg-to-dult survivl, rnging from 91% in whet to 15% in NOW (Fig. 1B). The men egg-to-dult survivl in produts ontining Cpsium spp. did not Men + SE numer of eggs per femle Men + SE egg-to-dult survivl (%) Men + SE lrvl survivl (%) signifintly differ (P4.5) mong them nd vried etween 35% nd 4%. The men perentge lrvl survivl positively orresponded to the egg-to-dult survivl rte in different food soures. Lrvl survivl ws signifintly different (F ¼ 18.5; df ¼ 6, 13; Po.1) mong food soures, with the gretest survivl of 92% on whet flour (Fig. 1C). Men perentge lrvl survivl on the remining food soures rnged from 33% to 52%, with no signifint differenes (P4.5) mong them, d Chili Pprik Cyenne Lef too Cigr too NOW Whet Fig. 1. Men7SE numer of eggs lid per femle L. serriorne (A) men perentge7se egg-to-dult survivl (B); nd lrvl survivl (C) of L. serriorne in different food soures. Mens re sed on N ¼ 4. Men vlues mong different food soures for given figure ompnied y different letters re signifintly different (Pp.5, LS mens). d

6 Author's personl opy R.M. Mhroof, T.W. Phillips / Journl of Stored Produts Reserh 44 (28) Tle 1 Men7SE for the egg hth period, lrvl period, pupl period, nd totl developmentl period in dys, for L. serriorne rered on different food soures Food soures Egg hth period Lrvl period Pupl period Totl development period Whet flour d d d Lef too Pprik Cyenne pepper d Chili Cigr too NOW Egg hth period did not differ signifintly mong different food resoures. Vlues in olumn followed y different letters re signifintly different (Pp.5, LS mens). Overll immture development time from egg hth to dult emergene. exept for the survivl on NOW (Pp.5) tht showed the lowest survivl rte Developmentl time for different life stges Overll, % of the eggs tested hthed to neonte lrve. There were no signifint differenes (F ¼ 2.; df ¼ 6, 13; P ¼.9) in the men egg development time oserved for different food soures (Tle 1). However, the lrvl development time signifintly differed mong the food soures (F ¼ 1.4; df ¼ 6, 124; Po.1). Development times in hili, yenne pepper, igr too nd NOW were signifintly longer (Pp.5) thn development time in whet flour. The differene etween the longest lrvl development period tht ourred in NOW nd the shortest development period tht ourred in whet ws pproximtely 54 d. The pupl development time in vrious food soures rnged from to d nd ws signifintly different (F ¼ 9.6; df ¼ 6, 12; Po.1) in vrious food soures (Tle 1). The shortest developmentl time in pupe ourred when lrve fed on whet. As summtion of egg development, lrvl development, nd the pupl development times the overll immture development time rnged from 46 to 19 d mong the food soures. Signifint differenes (F ¼ 14.5; df ¼ 6, 128; Po.1) were oserved for overll immture development time. The 46 d required when whet flour ws used s food soure ws signifintly shorter (Pp.5) thn for ny other food soures (Tle 1) Adult longevity nd ody weight The dult longevity of L. serriorne in the study with different food soures showed signifint differenes (F ¼ 58.7; df ¼ 6, 128; Po.1) mong tretments. Overll, dults emerged from lrve fed on whet flour lived signifintly longer ( d) thn those from the other six food soures (Fig. 2A). The shortest dult longevity ws reorded for eetles tht were rered on igr too. Although results from previous experiments indited tht NOW ws reltively poor medium for L. serriorne oviposition or development, dults tht emerged from lrve fed on NOW lived signifintly longer (Pp.5) thn those from produts ontining Cpsium spp. or too types. Similr to dult longevity dt, dult ody weight (dry weight) lso signifintly vried (F ¼ 9.2; df ¼ 6, 183; Po.1) mong individuls rered in different food soures. Adults tht emerged from either NOW or whet flour were signifintly hevier thn dults tht emerged from produts ontining Cpsium spp. or the two types of too (Fig. 2B). Further, results showed tht dult longevity is relted to the ody weight. Correltion nlysis showed signifint positive orreltion etween dult ody weight nd dult longevity, with Person orreltion o-effiient of.744 (Po.1). 4. Disussion In the urrent study, the experiment to determine the numer of eggs deposited y mted femles of L. serriorne ws done for individul femles. Pst results (Howe, 1957; Flether nd Long, 1971; LeCto, 1978) relting to vrious oviposition experiments were otined y verging the numer of eggs deposited y group of femles or mixed sexed dults. A shortoming of grouping femle L. serriorne in onfined environment is tht individuls my not relize their omplete oviposition potentil, euse pheromone-like sustne, serriorone, produed y the femle, is known to regulte ovipositionl ehvior of onspeifis (Kohno et l., 1986). Femles mrk ovipositionl sites with this hemil, so tht onspeifi femles n reognize nd void those sites. This nturl dpttion ensures dispersl of individuls nd provides vilility of food for the newly hthed lrve (Howlder nd Amdkr, 1995). In our experiment, y using single femle we eliminted or minimized the dverse effets on oviposition used y serriorone or ompetition for resoures. Our results showed tht the verge numer of eggs lid y single femle vried depending on the type of food soures provided s oviposition medium, nd therefore L. serriorne femles my e le to disriminte mong different food soures. In our study, mximl feundity of

7 Author's personl opy 224 R.M. Mhroof, T.W. Phillips / Journl of Stored Produts Reserh 44 (28) Men + SE dult longivity (dys) Men + SE dult weight (µg) d Chili Pprik Cyenne Lef too L. serriorne ws oserved on whet flour. Although too is desried s suitle host for L. serriorne (Howe, 1957), in our study L. serriorne femles deposited the lowest numer of eggs in igr too. This ws further demonstrted y Flether nd Long (1971) who lso showed higher oviposition in whet ompred to too. Quntittive nd qulittive differenes in hemil onstituents in food soures my diretly influene the oviposition ehvior of L. serriorne. Kohno nd Ohnishi (1986) reported tht not only the hemil stimulus ut lso the physil texture of the sustrte is limiting ftor for ovipositionl response. In our study the effet used y vrition in the texture of the sustrte ws minimized y pulverizing ll food soures nd sifting them through 425-mm meshed sieve. Adult L. serriorne emerged from ll of our tested food soures, ut the numer tht survived from egg to dult vried depending on the speifi food soure. The gretest survivl rte ws in the whet flour, whih ws similr to the results otined for the feundity study, nd mny other uthors (Howe, 1957; Flether nd Long, 1971; d e Cigr too NOW Whet Fig. 2. Men7SE of dult longevity in dys (A) nd ody weight/dult in mg (B) of L. serriorne in different food soures. Mens re sed on N ¼ 8. Men vlues mong different food soures for given figure ompnied y different letters re signifintly different (Pp.5, LS mens). LeCto, 1978) hve reported tht whet flour is good host for L. serriorne. Unlike for the feundity study, NOW seemed to e the poorest sustrte for L. serriorne survivl euse out 67% of the lrve died on NOW it. Egg-to-dult survivl in Cpsium nd too ws greter thn survivl in NOW ut lower thn survivl on whet flour. In different study, LeCto (1978) showed tht pprik nd yenne pepper were suitle hosts for L. serriorne egg-to-dult survivl, ut poor survivl ws oserved on hili powder. The survivl rte oserved y LeCto (1978) in hili powder ws out 38% lower nd in pprik nd yenne pepper ws out 37 48% higher thn our oservtion. LeCto (1978) used temperture of C, ut humidity onditions were not reported. The vrition oserved in egg-to-dult survivl in the two studies my possily e due to the differenes in the environmentl onditions. Our results for dult emergene orrespond to results from Powell (1931) whose work ws done t 6% r.h., similr to our experimentl onditions. Ninety-six perent of eggs used in our developmentl studies were vile nd hthed. Powell (1931) reported tht L. serriorne eggs do not hth if eggs re present in low humidity ondition or tempertures re elow C during inution. The time required for egg hth did not vry mong the different food soures, suggesting tht egg hth is not influened y the type of host mterils on whih eggs were lid. However, onsiderle vrition in the lrvl, pupl nd totl developmentl periods mong the tested food soures show tht lrvl development is influened y the speifi food soure, whih ould ffet pupl development. Lsioderm serriorne developed to the dult stge in shorter time on whet flour ompred to the other soures nd this fster development ould e euse of the lned nd essentil nutrients present in the whet. Whet flour is nutritionlly rih in vlule proteins, omplex rohydrtes, vitmins nd minerls (Cornell nd Hoveling, 1998) neessry for growth nd development of L. serriorne, nd the flour ould lso lk the defensive hemils present in other hosts we tested. Developmentl times oserved in other nutritionlly poor nd hemilly defended foods suh s Cpsium nd Niotin were moderte. The ility of L. serriorne to utilize toxin-rih hosts s food my e orrelted to the eologil tendeny of this speies to reed in food soures ontining defensive sustnes, nd lso relte to the ssoition of L. serriorne with yest-like symiont, Symiotphrin kohii (Shen nd Dowd, 1991). The symionts of L. serriorne potentilly help the inset to utilize host mterils whih re rih in plnt llelohemils nd onvert these to nutrients nd ron soures (Dowd, 1989; Dowd nd Shen, 199; Shen nd Dowd, 1991). The reson for prolonged L. serriorne development time in igr too ws not determined. Cigr too onsistently performed s poor ttrtnt for L. serriorne in our pst studies (Mhroof nd Phillips, 27). Developmentl time ws lso prolonged in L. serriorne rered on NOW it. NOW it is omposed of pressed lmond

8 Author's personl opy R.M. Mhroof, T.W. Phillips / Journl of Stored Produts Reserh 44 (28) flkes nd idulted lmond oil, nd is rih in lipids. The reson perhps NOW served s poor host for L. serriorne growth my e tht free ftty ids in proessed lipid-rih foods re oxidized when exposed to mient ir to form peroxide linkges (Love nd Person, 1971). Oxidtion of lipids results in unplesnt flvors nd odors (rnidity) nd my use the deteriortion of the food. Oxidtive rnidity in wlnuts proved to e toxi for egg hth nd development in the Indin mel moth, Plodi interpuntell (Hu ner) (Johnson et l., 1992). There ws positive reltionship etween ge of the dults nd their ody weight. The question of whether or not L. serriorne eetles feed s dults hs not een fully resolved (Lefkovith nd Currie, 1967; Minor, 1979; Shinod nd Fujiski, 21), ut the qulity of food ville to the lrvl stges ould influene dult longevity nd ody size. Adult eetles with lrge ody sizes, espeilly if they do not feed, my hve more nutrient reserves to id in longevity ompred to smller dults. Lefkovith nd Currie (1967) reported tht the life spn of L. serriorne is positively orrelted with the fresh weight of the newly emerged dults nd onluded tht the weight of dults is dependent in prt upon the mount of food onsumed s lrve. Among the seven food soures, whet flour gve the est results for L. serriorne survivl, growth, nd development. Lsioderm serriorne used in ll the experiments were initilly rised in medium of whet flour nd yest, nd therefore survivl on whet flour during the study my relte to the dpttion of our lortory olony to the rering medium. Although, severl different mehnisms in insets nd other higher nimls re sid to ffet the ehvior of host utiliztion, Bernys nd Chpmn (1994) suggest tht these mehnisms re short-term within the life of n individul, nd do not ffet susequent genertions. Thus, lerning nd experiene would e most fvored when within-genertion preditility of the resoures nd etween-genertions preditility is lest fvored. Our experiments with egg-to-dult survivl, lrvl survivl, development time, dult longevity nd ody weight were initited with eggs nd therefore, no opportunity ws ville for lerning to fvor whet s the suitle host, nd therefore genetilly inherited trits for host use re most likely operting. However, in our feundity experiments the dult femles used were initilly rised in whet flour medium in our lortory. Adults tht emerged from the pupl se my retin experiene or onditioning from the remnnts of the lrvl food ssoited with the pupl se. These remnnts my simply use n indued preferene for dults to ly more eggs in host with the sme or similr hemil ues. In the urrent study we found tht red hili, pprik, yenne pepper nd hewing lef too were suitle food soures for the survivl nd development of L. serriorne under lortory onditions. Further we onfirmed the suitility of whet, Cpsium produts nd some types of too s ovipositionl medi nd lrvl food soures for L. serriorne. Knowledge of suh host use ttriutes will e vlule in further reserh on host seletion y L. serriorne nd my help in the development of pest mngement tools for this serious inset pest. Aknowledgments The uthors re grteful to Drs. Rihrd Bereret nd Astri Wydnde for omments on the erlier drft of this mnusript. This work ws supported in prt with funds from the USDA CSREES Risk Avoidne nd Mitigtion Progrm (RAMP) funded under Agreement No , nd the Oklhom Agriulturl Experiment Sttion. Referenes Allotey, J., A study of the inset pests in stored plm produe in Port Hrourt, Nigeri. Journl of Stored Produts Reserh 24, Ashworth, J.R., The iology of Lsioderm serriorne. Journl of Stored Produts Reserh 29, Bernys, E.A., Chpmn, R.F., Host-Plnt Seletion y Phytophgous Insets. Chpmn & Hll, New York, 312pp. Bry, R.E., Botright, R.E., Lng, J.H., Dmge to otton textile g sewing thred y lrve of the igrette eetle (Coleopter: Anoiide). Journl of Georgi Entomologil Soiety 9, Cornell, H.J., Hoveling, A.W., Whet: Chemistry nd Utiliztion. Tehnomi Pulishing Compny In., Lnster, PA. Dimetry, N.Z., Brkt, A.A., El-Metwlly, H.E., Rish, E.M.E., Ad El Slm, A.M.E., 24. Assessment of dmge nd losses in some mediinl plnts y the igrette eetle (Lsioderm serriorne (F.)). Bulletin of Ntionl Reserh Center of Egypt 29, Dowd, P.F., In situ prodution of hydrolyti detoxifying enzymes y symioti yests in the igrette eetle (Coleopter: Anoiide). Journl of Eonomi Entomology 82, Dowd, P.F., Shen, S.K., 199. The ontriution of symioti yest to toxin resistne of the igrette eetle (Lsioderm serriorne (F.)). Entomologi Experimentlis et Applit 56, Flether, L.W., Long, J.S., Influene of food odors on oviposition y the igrette eetles on nonfood mterils. Journl of Eonomi Entomology 64, Hlsted, D.G.H., Externl sex differenes in stored-produts Coleopter. Bulletin of Entomologil Reserh 54, Highlnd, H.A., Proteting pkges ginst insets. In: Gorhm, J.R. (Ed.), Eology nd Mngement of Food Industry Pests. Assoition of Offiil Anlytil Chemists, Arlington, VA, pp Howe, R.W., A lortory study of the igrette eetle, Lsioderm serriorne (F.) (Col., Anoiide) with ritil review of the literture on its iology. Bulletin of Entomologil Reserh 48, Howlder, A.J., Amdkr, P.M., Oviposition deterring influene of femle ody wsh in too eetle, Lsioderm serriorne (F.) (Coleopter: Anoiide). Journl of Stored Produts Reserh 31, Jo, S., Host food preferene of the igrette eetle, Lsioderm serriorne (F.) to few stored spies. Plnt Protetion Bulletin 44, Johnson, J.A., Wofford, P.L., Whithnd, L.C., Effet of diet nd temperture on development rtes, survivl nd reprodution of the Indinmel moth (Lepidopter: Pyrlide). 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9 Author's personl opy 226 R.M. Mhroof, T.W. Phillips / Journl of Stored Produts Reserh 44 (28) Kohno, M., Mohizuki, K., Chumn, T., Ohnishi, A., Pheromonelike sustnes ffeting the oviposition ehvior of the femle igrette eetle, Lsioderm serriorne (F.) (Coleopter: Anoiide). Applied Entomology nd Zoology 21, LeCto, G.L., Infesttion nd development y the igrette eetle in spies. Journl of the Georgi Entomologil Soiety 13, Lefkovith, L.P., Currie, J.E., Ftors ffeting dult survivl nd feundity in Lsioderm serriorne (F.) (Coleopter: Anoiide). Journl of Stored Produts Reserh 3, Love, J.D., Person, A.M., Lipid oxidtion in met nd met produts. Journl of the Amerin Oil Chemists Soiety 48, Mhroof, R.M., Phillips, T.W., 27. Orienttion of the igrette eetle Lsioderm serriorne (F.) (Coleopter: Anoiide) to plnt derived voltiles. Journl of Inset Behvior 2, Minor, M.F., Do dult igrette eetle feed? Too Siene 23, Powell, T.E., An eologil study of the too eetle, Lsioderm serriorne (F.) with speil referene to its life history nd ontrol. Eologil Monogrphs 1, SAS Institute, 23. The SAS System Version 9.1 for Windows. SAS Institute, Cry, NC. Shen, S.K., Dowd, P.F., Detoxifition spetrum of the igrette eetle symiont Symiotphrin kohii in ulture. Entomologi Experimentlis et Applit 6, Shinod, K., Fujiski, K., 21. Effet of dult feeding on longevity nd feundity of the igrette eetle, Lsioderm serriorne F. (Coleopter: Anoiide). Applied Entomology nd Zoology 36, Zr, J.H., Biosttistil Anlysis, seond ed. Prentie-Hll, Englewood Cliffs, NJ.