ww w . r b e n t o m o l o g i a . c o m
REVISTA
BRASILEIRA
DE
Entomologia
AJournalonInsectDiversityandEvolutionBiological
Control
and
Crop
Protection
Biology
and
reproductive
capacity
of
Spodoptera
eridania
(Cramer)
(Lepidoptera,
Noctuidae)
in
different
soybean
cultivars
Bruna
Magda
Favetti
a,∗,
Alessandra
Regina
Butnariu
b,
Luís
Amilton
Foerster
caProgramadePós-graduac¸ãoemProtec¸ãodePlantas(Agronomia),UniversidadeEstadualPaulistaJúliodeMesquitaFilho(UNESP),Botucatu,SP,Brazil bProgramadePós-graduac¸ãoemMeioAmbienteeSistemadeProduc¸ãoAgrícola,UniversidadedoEstadodeMatoGrosso(UNEMAT),TangarádaSerra,MT,Brazil
cProgramadePós-Graduac¸ãoemZoologia,LaboratóriodeControleIntegradodeInsetos,DepartamentodeZoologia,UniversidadeFederaldoParaná(UFPR),Curitiba,PR,Brazil
a
r
t
i
c
l
e
i
n
f
o
Articlehistory: Received18March2014 Accepted26December2014 Availableonline9April2015 AssociateEditor:DanielSosa-Gomez
Keywords: Development Glycinemax Monsoy8757 Southernarmyworms
a
b
s
t
r
a
c
t
Thisstudyaimedtoevaluatethedevelopment,survivalandreproductivecapacityofSpodopteraeridania
infoursoybeancultivars.Theexperimentwasconductedinthelaboratory,inaclimaticchamberat 25◦C±1◦C,70±10%relativehumidityand12hphotophase.Thecultivarsusedwere:FMTTabarana, BRS/MTPintado,FMTTucunaréandMonsoy8757,allconventionalcultivarswithmediumcycles.All cultivarstestedallowedthedevelopmentofS.eridania.However,Monsoy8757wasthecultivarthat mostaffectedtheprolongedinthedurationlarval,pupalandtotalcycle,showedlowerpupalweightas wellasreductionintheintrinsicrateincrease.Theseresultscontributetothemanagementofthisspecies inregionsofoutbreaksinsoybeanareas.
©2015SociedadeBrasileiradeEntomologia.PublishedbyElsevierEditoraLtda.Allrightsreserved.
Introduction
Spodoptera eridania (Cramer) (Lepidoptera, Noctuidae) is a polyphagousdefoliatorthatfeedsonseveralspeciesofeconomic importance, such as apple (Nora and Reis-Filho, 1988), cotton
(Miranda,2010),soybean(Sosa-Gómezetal.,1993),maize(Picanc¸o
et al.,2003), beans(Savoie, 1988), rattlebox (Diasetal., 2009), tomato(Souzaetal.,2013),sorghum,saladgreensandfruit(King
andSaunders,1984),strawberryandgrape(Bortolietal.,2012);
weeds,suchasmorningglory(Santosetal.,2005)andpigweed
(Savoie, 1988); and bracatinga, which is used for reforestation
(MattanaandFoerster,1988).
Although this lepidopteran occurs in soybeans plantations, accordingtoSosa-Gómezetal.(1993)itspresenceinthisculture is incidental,havingthus asecondary role. Thedamage caused bythispestcanbesignificantbecause,aswellasdamagingthe leaves,italsoconsumesthepods,thusassuminganimportantrole atthebeginningofthereproductivephaseofthecrop(Gazzoniand
Yorinori,1995).
Controlof Spodopteraspeciesinsoybeanscanbemadefrom chemical pesticides, resistant varieties and transgenic plants
(Souzaetal.,2012a;Bernardietal.,2014).However,thereisalow
∗ Correspondingauthor.
E-mail:favettibruna@yahoo.com.br(B.M.Favetti).
efficiencyintheuseofchemicals(Carvalhoetal.,2013),aswellas tolerancetosomeBtproteins(Bernardietal.,2014)bySpodoptera. However,duetofrequentoutbreaksofS.eridaniaintheCerrado regionofBrazil(Santosetal.,2005;Buenoetal.,2007),itshigh capacitydefoliation(Buenoetal.,2011)andpopulationdensities, thisspeciesbecameanimportantpestofsoybean,causingdamage andeconomiclosses.
The Mato Grosso state, considered the largest producer of soybeaninBrazil(IBGE,2014),withlargecultivatedareas,the pop-ulationincreaseofvoraciousinsectsthatconsumesseveralsoybean genotypes(Souzaetal.,2012a), maybea factorinreducingthe productionofthiscrop.
Although this pest eating of various soybean genotypes, morphological and chemical differences between cultivars may interfereinbiologyandbehaviorandinducepestdegreesof resis-tance. This was verified by Souza et al. (2012a) evaluated the feedingpreferenceofS.eridaniafor23soybeangenotypes,where thecultivarIAC100showedahighdegreeofnon-foodpreference infreechoiceandnochoicetest.
Acommonagriculturalpracticeinmanyareasofsoybean culti-vationinthestateofMatoGrossoistheplantingofseveralcultivars indifferentplotsinthesamearea,orevensharingthesamefield. However,therearenoreportsintheliteratureaboutthe interfer-enceofthispracticeinpopulationdynamicsofS.eridaniainthe field.Thus,thisworkevaluatedtheinfluenceoffoursoybean culti-varsonthedevelopment,survivalandreproductionofthisspecies inthelaboratory.
http://dx.doi.org/10.1016/j.rbe.2015.03.002
Materialandmethods
S.eridaniawasrearedintheEntomologyLaboratoryofthe Cen-trodePesquisas,EstudosandDesenvolvimentoAgro-Ambientais, oftheUniversidadedoEstadodeMatoGrosso,campusofTangará daSerra-MT,aftercollectingspecimensinsoybeanplantations.
The larvae were maintained at room temperature (approx-imately 26◦C±2◦C) in 500mL polyethylene pots, lined with
absorbentpaper,anddailyfedwithsoybeanleavesuntilthe pre-pupalstage.Thepupaewereremovedfromthepotsandplacedin Petridishesandsexingofthepupaewascarriedoutaccordingto
Zenkeretal.(2007).
Afteremergence,theadultsweremaintainedatroom temper-ature(approximately26◦C±2◦C)inPVCtubes(10cmindiameter
by21.5cmhigh),withaPetridishonthetopandthebottom,and coatedinternallywithbondpaperfunctioningasasubstratefor oviposition.Asolutionof 10%honey, offeredasfoodinsidethe tubeswasplacedinaplasticcontainercontainingcottonwool.Eggs depositedonthebondpaperwereremovedbycuttingthepaper andthen transferredtoaPetri dishkeptina climaticchamber (25◦C±1◦C, 70%±10% RHand 12hphotophase).Tube
mainte-nanceandexchangeoffoodwerecarriedoutdaily.
Themediumcyclesoybeancultivarswerechosenbecausethey aresomeofthemoreconventionalgenotypesusedinthestateof MatoGrosso.Thecultivarsusedwere:FMTTabarana,BRS/MT Pin-tado,FMTTucunaréandMonsoy8757,providedbyaseedresale company.
Theplantsweregrowninplasticbuckets,withcapacityfor8kg ofsoil,inagreenhouselocatedinUNEMAT,campusofTangará daSerra(MT).Thecommercialfertilizerusedintheexperiments wasNPKataconcentrationof10%foreachcompound.Whenthe V8developmentstagewasreached(approximately35daysafter planting),theleaveswerecollectedandimmersedfor20minina1% solutionofcommercialsodiumhypochloritetoeliminatepossible entomopathogensand,soonafter,offeredtothelarvae.
Forty-ninenewlyhatchedlarvae,foreachsoybeancultivar,were maintainedindividually in 145mL plastic potslined withfilter paperandfeddaily.Theyweremaintainedinaclimaticchamberat 25◦C±1◦C,70%±10%relativehumidityandphotoperiodof12h.
Thenumberofinstarsandtheirdurationwasdeterminedbythe presenceofthecephaliccapsuleandexuviate.Thisprocedurewas repeateduntilthepre-pupalstage,whenthecaterpillarintegument changescolorandstopfeeding.
ThepupaewereplacedinPetridishesand,24hafter forma-tion,theoccurrenceofdeformitieswasevaluated,theyweresexed, andweighedusingananalyticalbalanceof0.0000gaccuracy.The biologicalparametersusedweredurationoflarval,pre-pupaland pupalstages(days), viabilityofimmaturestages (%),number of instars,weightpupal,percentageofdeformedpupae,andsexratio (numberoffemales/(numberofmales+numbersoffemales)).
Foreachtreatment,15coupleswereisolatedinPVCtubecages (10cmindiameterby21.5cmhigh),accordingtothedateof emer-gence.The cages weremaintainedat 25◦C±2◦C, coveredwith
voile,placedonaPetridish,andinternallylinedwithbondpaper asasubstrateforoviposition.
Adultswerefedasolutionof10%honeyoncottonwoolplaced inplasticvialsatthebottomofthecages.Theeggscontainedin thesheets ofbond paperandvoilewereremoveddaily,placed inPetridishes(10cm diameter)andkeptinaclimaticchamber (25◦C±1◦C,70%±10%RHand12hphotoperiod)untilthe
emer-genceofthecaterpillars.
ToevaluatethereproductiveperformanceofS.eridaniainthe fourtreatments,thebiologicalparametersconsideredwere: fecun-dity(dailyand total),egg viability,incubation period,longevity (malesandfemales),pre-oviposition,ovipositionperiodsandtotal cycle(egg-adultemergence).Torecordfecundityandeggviability,
eggswereremoveddailyandisolatedinPetridishesuntil caterpil-laremergence.
Eggswerecountedwiththeaidofphotographicimages. Pho-tographs of each egg clutch were taken with a camera and a stereoscopicmicroscopeandtransferredtoacomputerfor count-ingonthescreen.Fortheeggmassescontainingmorethanone layerofeggs,thenumberofeggsonthetoplayerwerecounted, multipliedbythenumberoflayers,andaddedtothenumberof sur-roundingeggs.Thetotalviabilityoftheeggswasfoundbyrecording larvalemergencethroughouttheovipositionperiod.
Withthedataofsurvivalandovipositionofeachfemalewas preparedthefertilitylifetable.Theparametersfollowedthe for-mulascontainedinMaiaetal.(2000).Wecalculatedaccordingto
Montezanoetal.(2013):Ro(netreproductiverate)=ratiobetween
thenumberoffemalesintwosuccessivegenerations;T(mean gen-erationtime)=meannumberofdaysfromthebirthoftheparents tothebirthofoffspring;rm=dailyintrinsicrateofincrease;=the dailyfiniterateofincrease.
Theexperimentaldesignwascompletelyrandomizedandthe resultsweretestedbyShapiroWilk’snormalitytest(p>0.05).Data thatdidnotreachnormalityweretestedusingtheKruskal–Wallis test(5%)andtherestwereanalyzedbyANOVAat5%.Meanswere comparedusingtheTukeytestat5%probability.Tocomparethe sexratio,thechi-squaretestwasusedwith0.5fortheexpected value.ThesoftwareusedwereRStudioandStatistica7.0.
Results
DevelopmentoftheimmaturestagesofS.eridaniainlaboratory
Therewasnodifferenceinthedurationoftheincubationperiod ofeggsofS.eridaniarearedinthefoursoybeancultivars,theaverage beingaboutthreedays(Table1).Thelarvalperiodrangedfrom15 to19daysandwassignificantlyinfluencedbythecultivarstested, beingsimilaronlyincultivarsFMTTabaranaandFMTTucunaré,and inMonsoy8757,thehighestlarvalperiodwasrecordedcompared totheothercultivars(Table1).
ThenumberofinstarsofS.eridaniarangedfromsixtoseven, andonlyindividualsfedwithBRS/MTPintadodidnotreachthe7th instar(Fig.1).Themeandurationofthe1stinstarwasthreedays, withnosignificantdifferencesbetweencultivars.Inthe2ndand 3rdinstarslastedlongerincultivarFMTTucunaré,whencompared totheothers(Fig.1).Inthe4thand5thinstarsthehighestmeans werefoundincultivarMonsoy8757(Fig.1).Thedurationofthe 6thinstarwasnotstatisticallydifferentamongthecultivars(Fig.1). Althoughthenumberofindividualsinthe7thinstarwaslower,the meanforalltreatmentsthatreachedthisinstarwasapproximately fivedays(Fig.1).
ThepercentagesofindividualsofS.eridaniathatreachedthe7th instarwere12.2%,0.0%,6.1%and2.0%,whenfedonFMTTabarana, BRS/MTPintado,FMTTucunaréandMonsoy8757,respectively.The onlylarvalmortalityrecordedwasforthecultivarFMTTucunaré inthe5thand6thinstars(Fig.1).
Thedurationofthepre-pupalstageofS.eridaniawassimilar amongcultivars(Table1).Theaveragedurationofthefemalepupal stage,fedonMonsoy8757duringthelarvalstage,wasgreaterthan theothercultivars(Table1).Thedevelopmenttimeofthemale pupalstagewassimiliarinallcultivars(Table1).
Table1
Mean(±SD)duration(days)ofimmaturestagesofSpodopteraeridaniamaintainedinfoursoybeancultivars.
Cultivars Durationofimmaturestages(days)
Egga Larvaeb Prepupaea Pupa
Femaleb Malea
FMTTabarana 3.2±0.43 15.9±0.70b 1.0±0.24 10.5±0.63b 11.4±0.61 BRS/MTPintado 3.1±0.30 15.0±0.80c 1.0±0.23 10.5±0.69b 11.3±0.57 FMTTucunaré 3.1±0.30 16.4±1.01b 1.2±0.42 10.6±0.59b 11.2±0.62 Monsoy8757 3.2±0.40 18.5±0.93a 1.1±0.48 11.2±0.46a 11.8±0.49
H 3.1 129.5 11.2 18.2 16.6
Df 3 3 3 3 3
p – <0.05 – <0.05 >0.05
aNotsignificantaccordingtoKruskal–Wallisat5%.
bMeansfollowedbythesameletterdonotdifferstatisticallybyKruskal–Wallistestat5%probability.
6
100
80
60
40
Survival (%)
Duration of instars (days)
20
0 5
4
3
2
1
0
FMT tabarana FMT tucunaré Survival
Monsoy 8757 BRS/MT pintado
ns
b b a b b b a b c
b
b
ab ab a
bc a ns ns ns
nsns ns ns ∗ ∗ ∗
1st 2nd 3rd 4th 5th 6th 7th
Instars
Fig.1.Duration(days)andsurvival(%)oflarvalinstarsofSpodopteraeridaniafedondifferentsoybeancultivars.MeansfollowedbythesameletterdifferedbyKruskal–Wallis testat5%.nsNotsignificantaccordingtoKruskal–Wallis(p>0.05).*Duetothelownumberofindividuals,itwasnotpossibletoapplystatisticaltests.
soybeancultivarsevaluated,andthevaluesrangedfrom0.4to0.5
(Table2).
Inalltreatments,theviabilityofthelarval,pre-pupal,andpupal stageswashigh,withnosignificantdifferencesamongtreatments
(Table3).Thetotalviabilityoftheeggswasnotaffectedbythe
cul-tivars,rangingfrom43.0%to53.6%(Table3).Overtheoviposition period,theviabilitywasabove60%,fromthe2ndtothe8thdayof oviposition(Fig.2).
ReproductivecapacityofS.eridaniainlaboratory
ThelongevityofmalesandfemalesofS.eridaniawasnot influ-encedbythedifferentsoybeancultivars;femalelongevityranged from10.1to11.1days,andmalelongevitybetween13.6and17.8
days (Table4).The duration of thepre-oviposition period was
alsonotaffectedbythecultivarstested,withameanoftwodays
(Table4).DifferencesinovipositionperiodofS.eridaniawerenot
Table2
Mean(±SD)totalweightpupal(g)andfemalesandmales,pupaldeformity(%)andsexratioofSpodopteraeridaniainfoursoybeancultivars.
Cultivars Pupalweighta Deformitypupal(%) Sexratiob
Mean Female Male
FMTTabarana 0.2509±0.02a 0.2533±0.02a 0.2474±0.01a 6.1 0.59
BRS/MTPintado 0.2195±0.02b 0.2296±0.02b 0.2122±0.02b 14.2 0.42 FMTTucunaré 0.1978±0.02c 0.2079±0.02c 0.1882±0.02c 10.6 0.48
Monsoy8757 0.1723±0.02d 0.1816±0.02d 0.1639±0.01d 0.0 0.42
F 12.0 95.2 39.9 – –
X2 – – – – 4.0
Df 3 3 3 – 3
p <0.05 <0.05 <0.05 – >0.05
0 0 20 40 60 80 100
1 2 3 4 5 6 7
Oviposition (days)
V
iability of eggs (%)
8 9 10 11 12 13 14 15 16
FMT tabarana
FMT tucunaré Monsoy 8757 BRS/MT pintado
Fig.2.Eggviability(%)offemalesofSpodopteraeridaniaduringtheovipositionperiod.
observedamongcultivars,andthevaluesrangedfrom7.3to7.8
days(Table4).Thetotalcycle(egg-adultemergence)ofS.eridania
waslongerinlarvaefedwithcultivarMonsoy8757,when com-paredwiththeothercultivars,andthisperiodranged,onaverage,
from29.1to33.3days(Table4).
The soybean cultivars tested did not affect the fecundity and egg viability of S. eridania (Tables 3 and 4). Although no differences were observed, total fecundity ranged from 2034.6 to 2766.4 eggs, and the number of eggs produced per day by females ranged from 211.0 to 279.1 (Table 5). The cul-tivars tested did not affect the total number of egg masses of S. eridania, which ranged from 8.8 to 11.1 egg masses
(Table5).
Inallcultivars,thehighestovipositionoccurredonthe3rdand 4thdays.Therewasalsoadownwardtrendinthenumberofeggs andlarvaetowardtheendoftheovipositionperiod(Fig.3).
Duringtheovipositionperiod,thepercentageofeggviabilityof
S.eridania,rearedinfoursoybeancultivars,wasabove60%untilthe 8thand/or11thday,dependingonthecultivar(Fig.2).Afterthis period,adecreaseinviabilitywasobservedinallcultivars(Fig.2). TheaveragenumberofeggsperfemaleofS.eridaniaduringthe ovipositionperiodwashigherinthefirstfewdays(1st–4th),with adecreasetowardtheendofthisperiod,exceptincultivarFMT Tabaranainwhichanincreaseinthenumberofeggswasobserved fromthe12thtothe14thday(Fig.4).
Thefertilitylifetableshowedthatallcultivarsaresuitablehost tocompletedevelopmentofS.eridania.Thenetreproductiverate (Ro)indicatedthatS.eridaniafedwithMonsoy8757mayincrease 558.12 times in each generation, and double in FMT Tabarana (1004.12pergeneration).Themeangenerationtime(T)washigher maintainedinMonsoy8757andlowestinBRS/MTPintado.The intrinsicrateofincrease(rm)wassimilarto0.20forallcultivars,
Table3
Mean(±SD)viabilityofimmaturestagesofSpodopteraeridaniainfoursoybeancultivars.
Cultivars Viabilityofimmaturestages(%)
Egga Larvaea Prepupaa Pupaa
FMTTabarana 43.1±30.97 100.0±0.00 100.0±0.00 93.8±24.22
BRS/MTPintado 43.0±31.97 100.0±0.00 100.0±0.00 85.7±35.35
FMTTucunaré 51.1±34.60 95.9±19.99 100.0±0.00 89.3±31.16
Monsoy8757 53.6±30.14 100.0±0.00 100.0±0.00 100.0±0.00
H 2.0 6.0 0.0 7.7
Df 3 3 3 3
p >0.05 >0.05 >0.05 >0.05
aMeansnotsignificantaccordingtotheKruskal–Wallistestat5%probability.
Table4
Mean(±SD)longevityoffemalesandmales(days),pre-ovipositionperiod(days),oviposition(days)andtotalcycle(days)ofSpodopteraeridaniarearedondifferentsoybean cultivarswhenlarvae.
Cultivars Longevitya Pre-ovipositiona Ovipositiona Totalcycleb
Female Male
FMTTabarana 11.0±2.18 13.6±3.99 2.4±0.63 7.7±1.79 30.1±1.08b BRS/MTPintado 10.3±1.91 15.5±0.46 2.4±0.50 7.8±1.52 29.1±1.25b FMTTucunaré 10.1±1.50 14.8±0.44 2.1±0.51 7.2±1.62 30.4±1.16b Monsoy8757 10.8±3.88 17.8±1.48 2.6±0.82 7.6±2.02 33.3±1.06a
H 1.3 6.4 2.9 1.1 –
F – – – – 77.0
Df 3 3 3 3 3
p – – – – <0.05
Table5
Mean(±SD)ofdailyandtotalnumberofeggmassesandeggsofSpodopteraeridaniarearedindifferentsoybeancultivarswhenlarvae.
Cultivars No.eggmassesa Fecunditya
Daily Total Daily Total
FMTTabarana 1.1±0.42 8.8±7.41 263.2±270.69 2338.0±2890.40
BRS/MTPintado 1.1±0.25 9.7±5.04 261.6±216.76 2478.1±2131.02
FMTTucunaré 1.2±0.42 11.1±5.88 279.1±274.19 2766.4±2754.52
Monsoy8757 1.1±0.19 9.1±5.76 211.0±173.61 2034.6±2032.11
H 1.2 1.1 0.3 0.7
Df 3 3 3 3
p >0.05 >0.05 >0.05 >0.05
aMeansnotsignificantusingtheKruskal–Wallistestat5%.
Table6
Netreproductiverate(Ro),meangenerationtime(T),dailyintrinsicrateofincrease(rm)andthedailyfiniterateofincrease()oftheSpodopteraeridaniainfoursoybean cultivars.
Parameters Cultivars
FMTTabarana BRS/MTPintado FMTTucunaré Monsoy8757
Ro 1004.12 587.08 790.81 558.12
T 33.30 32.22 33.52 36.89
rm 0.21 0.20 0.20 0.17
1.23 1.21 1.22 1.18
exceptinMonsoy8757.Thefiniterateofincrease()corresponding tothenumberofindividualsaddedtothepopulation/female/day ranged1.18–1.23(Table6).
Discussion
TheresultsshowedthatthedevelopmentofS.eridaniawasmost affectedbythecultivarMonsoy8757,whencomparedtotheother cultivars,sinceanincreaseinthedurationofthelarval,pupaland totalcycleoccurred,aswellasareductioninpupalmass.Thefact thatthiscultivaraffectedthesebiologicalparametersisprobably relatedtothelownutritionalqualityoftheingestedfood.
Accord-ingtoPanizziandParra(2009),ingestionintheearlylifestagesis
ofparamountimportanceforphytophagousinsectsbecausesome dietaryrequirementsofthehostplantmustproperlyfilltheir nutri-tionalneeds,inorderforthemtodevelopnormallyintotheadult stage.
Thelengtheningofthelarvaland pupalstages,and increase in number of instarsin cultivars FMT Tabarana, FMT Tucunaré andMonsoy8757mayhaveoccurred,accordingtoMachadoetal.
(1999),asacompensatorystrategyforS.eridaniatoreachtheideal
masswithlessthanadequatefood.
The duration of theinstars of S.eridania maintained in dif-ferent soybeancultivars provided differentresponses,sincethe earlyinstarsshowedreduced periodsand the latteroneswere longer(Fig.1).Thisispossiblyrelatedtothefactthat,attheend ofthelarvalstage,thelarvaeofLepidopteraconsumemorefood
(Grützmacheretal.,1999)tomeettheirnutritionalneeds(Bortoli
etal.,2005),acquiringthenecessaryreservesforthedevelopment ofpupaeand,consequently,adults.
Theadditionofaseventhinstarinthreeofthesoybean cul-tivarstestedwassimilartothatfoundbyParraetal.(1977)and
Santoset al.(2005),inwhich larvaeofS.eridania rearedin
dif-ferenthostplants reachedtheseventhinstaronlywhenfed on soybeanleaves.AccordingtoEsperketal.(2007),theoccurrence ofadditionalinstarsisacompensatorymechanismduringadverse conditions,wheretemperature,photoperiod,qualityandquantity offood,densityofindividuals,andhumidityarethemostcommon factorsthataffectthenumberofinstars.
Esperket al. (2007) showed that in somespecies of
noctu-ids, suchas Anticarsia gemmatalisHüebner, 1818 (Lepidoptera, Noctuidae), Chrysodeixis includens (Walker, 1858) (Lepidoptera,
Noctuidae),SpodopterafrugiperdaandSpodopteralitura(Fabricius, 1775)(Lepidoptera,Noctuidae),thequalityoffoodisreferredto asoneofthepossiblefactorsaffectingtheincreaseinnumberof instars,afactthatmayalsohaveoccurredwiththelarvaeofS. erida-nia,sincetheothervariablesmentionedaboveremainedconstant inthepresentstudy.
ThemeandurationofpupaeofS.eridaniawasgreaterthanthose recordedbyFarahanietal.(2011)forSpodopteraexigua(Hüebner, 1808)(Lepidoptera,Noctuidae)ondifferentsoybeancultivars,in whichthepupalperiodrangedfrom7.0to8.9days.Theresults forthepupaldurationinfemalesandmalesweresimilartothose reportedbyMattanaandFoerster(1988)forS.eridaniainsweet potato(10.0daysformalepupae and9.0daysforfemales)and bracatinga,(11.8daysformalepupaeand12.0daysforfemales). Forthesamespeciesfedonsoybeansduringthelarvalstage,Parra
etal.(1977)reportedlowervaluesforfemale(8.6days)andmale
(9.5days)pupae.
Gazzoniand Tutida(1996)reportedthat highpupalmassin
femaleLepidoptera, as foundinS. eridaniawhen fed withFMT TabaranaandBRS/MTPintado,mayberelatedtoahighlevelof reserves,whicharefundamentalforadequateperformanceinthe reproductivephase,includingmating,oviposition,andegg viabil-ity.
Somedisadvantagesofinadequatefoodintakearesuggested
byBelandandHatchett(1976),suchastheextensionofthelife
cyclewhich,inthefield,inadditiontocausingotherphysiological disorders,exposestheinsectstonaturalenemiesforalongertime, usuallyunderstressandhighersusceptibility.Thehighsurvival ofthedevelopmentalstagesandthehighreproductivepotential ofS.eridaniafoundinthepresentstudyshowsthatithasagreat potentialformultiplicationinsoybean.
Souza et al. (2012b) evaluated the effect of different
culti-varsof cowpea onthebiological parametersof S.eridania,and foundthatdevelopmentwassignificantlyaffectedbythecultivars. OnefactorthatmayhaveaffectedthedevelopmentofS.eridania, whenfedwithcultivarMonsoy8757,isthepossiblepresenceof toxins.
8000 FMT tabarana 7000
6000
Fecundity Fertility
5000 4000 3000 2000 1000
0 1 2 3 4 5 6 7
Oviposition (days)
T
o
tal number of eggs an larvae
8 9 10 11 12 13 14 15 16 17
8000 BRS/MT pintado
7000 6000
Fecundity Fertility
5000 4000 3000 2000 1000
0 1 2 3 4 5 6 7
Oviposition (days)
T
o
tal number of eggs an larvae
8 9 10 11 12 13
8000 FMT Tucunaré
7000 6000
Fecundity Fertility
5000 4000 3000 2000 1000
0 1 2 3 4 5 6 7
Oviposition (days)
T
o
tal number of eggs an larvae
8 9 10 11 12 13
8000 Monsoy 8757
7000 6000
Fecundity Fertility
5000 4000 3000 2000 1000
0 1 2 3 4 5 6 7
Oviposition (days)
T
o
tal number of eggs an larvae
8 9 10 11 12 13 14
Fig.3.TotalnumberofeggsandlarvaeofSpodopteraeridaniaduringtheovipositionperiodincultivarsTMGTabarana,BRS/MTPintado,FMTTucunaréandMonsoy8757.
glucosides,cyanogens) or proteinaceous (lectin, arcelin, vicilin, sistemin,chitin,glucanaseandenzymeinhibitors)(Francoetal., 2002),withpre-andpost-ingestiveeffects,hinderingthe devel-opment of insects (Bortoli et al., 2012). Piubelli et al. (2005), whenstudyingthebiologyofA.gemmatalisafteringestionof soy-beangenotypescontainingisoflavones,observedhighmortality, decreasedweightoflarvaeandpupae,andaprolongedcycle.
The ingestion of substances produced by soybean that can providedifferentdegreesofresistancetodefoliatorscauses neg-ativeeffectsonthebiologyofinsects,suchasantibiosis.Thus,as therewerenodrasticchangesinthedevelopmentofS.eridania, exceptfortheadditionofoneinstar(seventh),greaterdurationof thelarvalandpupalstages,andsignificantlysmallerpupalmass,it isbelievedthattheseantibioticeffectsarebeingonlymoderately expressed(VendramimandGuzzo,2009).
Ingeneral,althoughnoobservesignificantdifferencesbetween variousbiologicalparametersofS.eridaniaamongcultivars, Mon-soy8757prolongedthelarval,pupalandtotalcycle,andshowed lower pupal weight as well as reduction in the intrinsic rate increase.Thesecharacteristicsmaybefavorabletotheproducer, since theextensionof duration thecycle canreduce the num-berofgenerationsin thefield,andthedecreasein thenumber offemales produced per female,in nextgeneration, willresult infewerofeggmassesandconsequentlylesscaterpillarsinthe field.
The increase in the density of pest insects, from the high capacityofmultiplicationis acrucialfactorof outbreaks.Inthe presentstudy,themultiplicationwasnotaffectedbytested soy-bean cultivars, suggesting that population outbreaks no occur onlybyagriculturalpracticeusedinMatoGrossostate.Thegood
0 0 200 400 600 800 1000
1 2 3 4 5 6 7
Oviposition (days)
Number of eggs/female
8 9 10 11 12 13 14 15 16 17
FMT tabarana
FMT tucunaré Monsoy 8757 BRS/MT pintado
developmentandreproductionofS.eridaniaintestedcultivarsmay beoccurredbypolyphagous habit. Thisdemonstrating thatthe speciesisadapted toconsumeplantswithdistinct morphologi-calandphysiologicalcharacteristics,thusprovidinghigherfeeding plasticityandpromotingthemaintenanceofthereproductivecycle ofthepest.Thus,studiesthatconsiderthepopulationdynamicsof thispest,inthefield,arefundamentaltoseekwaystopreventand controloutbreakspopulation.
Conflictsofinterest
Theauthorsdeclarenoconflictsofinterest.
Acknowledgements
The authors thank the Universidade Federal do Paraná and Universidadedo EstadodeMatoGrosso(UNEMAT)forthe sup-portintheaccomplishmentofthis research.We alsothankthe Coordenac¸ão de Aperfeic¸oamentode Pessoal de Nível Superior (CAPES)forprovidingscholarships.
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