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Abundance of Chlamydastis platyspora (Elachistidae) on its host plant Roupala montana (Proteaceae) in relation to leaf phenology PDF

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Preview Abundance of Chlamydastis platyspora (Elachistidae) on its host plant Roupala montana (Proteaceae) in relation to leaf phenology

JournaloftheLepidopterists'Society 57(4),2003,291-294 ABUNDANCE OF CHLAMYDASTIS PLATYSPORA (ELACHISTIDAE) ON ITS HOST PLANT ROUPALA MONTANA (PROTEACEAE) IN RELATION TO LEAF PHENOLOGY Aurora Bendicho-Lopez,1 Ivone Rezende Diniz2and John DuVall Hay3 InstitutodeCienciasBiologicas, UniversidadedeBrasilia,ICC Sul,Terreo,SalaATI49,CampusUniversitarioDarcyRibeiro,AsaNorte, 70910-900,Brasilia,DistritoFederal,Brazil ABSTRACT. Chlamydastisplatyspora (Elachistidae) is abivoltine specieswhose larvais aspecialistonRoupala montana Aubl. (Pro- teaceae),acommontreeinthecerrado.Westudiedthepresenceoflarvaeinrelationtoleafphenologyofitshostplantinthecerradosensu stricto(savannah-likevegetation) oftheFieldStationAguaLimpa,belongingtothe UniversityofBrasilia, Federal District, Brazil. Weexam- ined3600plantsinanareaof16.2habetween November1999andOctober2000.Thehostplantproducesleavesasynchronouslyduringthe year,andindividualspresentoneofthreeleafphenologicalphasesatagiventime: (1)newleavesonly,(2)matureleavesonlyand(3)bothma- tureandoldleaves.Larvaewerefoundon273oftheexaminedplants. LarvaewereencounteredbetweenJanuaryandMarch(firstgeneration) andwerefoundonlyonplantsofthethirdphenologicalgroup. LarvaewerealsoencounteredbetweenJuneandOctober(secondgeneration) andoccurredpredominantlyonthethirdgroup.AlthoughthehostplanthasahighabundanceinthecerradoareathepresenceoflarvaeofC. platysporaisapparentlylimitedbytheabundanceofplantsthatsimultaneouslyhavematureandoldleaves. Additionalkeywords: Brazil,caterpillar,cerrado,feedingspecialist. The center ofdistribution ofthe family Proteaceae Materials and Methods is South Africa and Australia. This family contains 72 Study area and its host plants. The studyarea, a genera and about 1400 species but onlythree genera cerrado sensu stricto, was located on the Field Station occur in Brazil: Qrevillea, Euplassa andRoupala (Joly Fazenda Agua Limpa (15°55'S, 47°55'W) ofthe Uni- 1993, Mendonca et al. 1998). The species ofRoupala versity ofBrasilia, Federal District, Brazil, at 1100 m occur mainly in the cerrado, but also are found in in elevation. This regionhas twowell-defined seasons, otherbiomes, such as the Atlantic forest. adryone from Mayto Septemberandawetone from Roupala montana Aubl. is common in the cerrado OctobertoApril (Fig. 1A). sensu stricto and is found from the APA of Curiau Samplingtookplace threetimespermonthover 12 (Amapa) (00°20'N 51°03'W) to Jaguariaiva (Parana) months from November 1999 to October 2000. Over (24°09'S 50°18'W) (Ratter et al. 2000). The highest the study period we inspected 3600 individuals (300 production ofleaves ofR. montana in the cerrado of permonth) ofR. montana inan areaofapproximately central BraziloccursduringSeptemberandOctober, a 16ha. Allindividuals inspectedwere between 0.5 and transitional period from dry to wet season (Franco 1.5 m inheightandtherewasno repetitionofindivid- 1998). However, leafproduction may occur in some uals over the samplingperiod. As C. platyspora is bi- individuals during the whole year, corresponding to voltine this period comprised both generations. The the pattern found in severalwoodyspecies ofthe cer- first generation occurs in the wet season and the sec- rado (Morais et al. 1995). ond occurs during the transition period from the end In a study conducted in a cerrado near Brasilia in ofthe dry season to the beginning ofthe wet season central Brazil, Diniz & Morais (1995) showed that (Bendicho-Lopez2000). On eachcollection date, 100 Chlamydastis platyspora (Meyrick, 1932) (Elachisti- individuals ofR. montanawere surveyedforthepres- dae) was locally restricted to R. montana. Chlamy- ence of larva of C. playtspora. When encountered, dastis platyspora is bivoltine and its first generation the developmental stage of each larva was recorded occurs between November and April and its second and an evaluation of the phenological stage of the between MayandOctober(Bendicho-Lopez2000). In leaves on the individual was also made. Detailed in- spite ofR. montana beingcommoninthecerradonear formation on the identification ofthe developmental Brasilia (Ratter 1980), the larvae ofthis moth are not stages of the larvae is given in Bendicho-Lopez & foundingreatnumbers, orwithhighfrequency(Diniz Diniz (in press). The phenological stage ofthe leaves etal. 2001). Since resources maybeconcentratedboth was basedonthe densityoftrichomes ontheirabaxial in time and space plant phenology may affect the dis- surface. This characteristic was used as indicator of persionofherbivores (Solomon 1981). Theobjectiveof the relative age of the leaves, classifying them into diepresentstudywas: toquantifydieabundanceoflar- three categories: vaeofC. playtspora in relationtoplantleafphenology. — Newleaves expandingleaves or recentlyexpanded leavesstilltotallycoveredbytrichomesonbodisur- 1Pos-graduacaoemBiologiaAnimal. 2DepartamentodeZoologia. faces. This stage lasts for less than seven weeks 3DepartamentodeEcologia. (Fig. 2A); 292 Journalofthe Lepidopterists' Society xioo X "40 « Month •NL B •ML MOL £ 250-i 2 200H > ® 100« 50 > o Z h s Month FlG. 1. Meanmonthlyprecipitationandrelativehumiditydur- ingthestudyperiodandleafphenologyofthehostplant.A, Mean monthly precipitation (Precip) and relative humidity (R.H.) (Nov/1999-Oct/2000),datafromtheIBGE Meteorological Station, Brasilia; B,Variationinfoliarphenophasesoftheindividualsexam- inedofRoupala montana. NL = newleaves, ML = matureleaves, MOL= matureandoldleaves. Fig. 2. Relative age ofRoupala montana leaves showing the Matureleaves—expandedleaves,whichhadalready abaxialface:A,Newleaves;B,MatureleavesandC,Oldleaves. begunto losetheirtrichomes. This stagelasts be- procedurewas repeatedinJune usinglarvae foundon tween eight and nine months (Fig. 2B); Old leaves—leaves lacking trichomes. This stage another 15 individuals of jR. montana to accompany the residence period and development oflarvae from lasts forupto two months (Fig. 2C). the secondgeneration. Individuals of R. montana were classified in three Statisticalanalyseswere done usingStatistix7(Ana- phenologicalgroups: (1)plantswithallnewleaves (NL), lytical Software 2000). (2) plants with only mature leaves (ML) and (3) plants Results with mature andoldleaves atthesametime (MOL). To compare the residence period and the develop- Leaf phenology of Roupala montana. January, menttime ofalarvaon its hostwe followedthe devel- middleofthewetseason, hadthelargestproportionof opment of larvae on marked host plants. The resi- plants with new leaves; while the maximum level for dence period and development time oflarvae in the mature leaves was May (beginningofthe dry season). first generation was studied using eggs or first instar Also atthe beginningofMay, therewas an increase in larvae found on 15 individuals ofR. montana in Janu- the number of plants of the group mature and old ary. These larvaewere observedtwice aweekthrough leaveswith the maximum numberrecordedin August thepupalstate untilthe emergence ofthe adults. This (Fig. IB). Volume 57, Number4 293 nologicalgroups (Fig. 3B). Atestofproportionsshowed a significant difference in die proportion ofplants with larvaebetweengenerations (z = -7.42, p = 0.000). Also 600 IEfl 50° the abundance oflarvae differed among die tiiree leaf 3. 400 phenophasesinbotiithefirst(%2 =45.75;p = 0.000) and juj 300 second(x2 = 115.72;p = 0.000) generations. a 200 All monitored larvae used in the studv ofresidence g 53 100 periodanddevelopmenttime, from the firsttothelast instars, remained on their monitored plants. In both NL ML MOL trials, all of the monitored plants belonged to the Leafphenologicalstage MOLphenologicalgroup. B Discussion MOL LarvaeofC.platyspora usedthe phenological 600i 3n 500 group ofplantswiththehighest frequency. Therefore, 3. 400 for this species the results do not corroborate what is ® 300 commonforinsectherbivoresofmoisttropicalforests, a 200 namely that the majority use new leaves (Coley & Zs 100- Barone 1996). Herbivores that can use old leaves, with low nutri- NL ML MOL tional quality, may be able to take advantage ofa pe- Leafphenologicalstage riod of low predator density (Moran & Hamilton 1980). They also avoid physical defenses such as leaf withlarvae trichomes that are present on new leaves (Purlin & D withoutlarvae Gilbert 1989, Paleari & Santos 1998). As reported by Fig.3. AbundanceoflarvaeofChlamydastisplatyspora found Morais et al. (1999) previous studies in the cerrado onleavesoiRoupalamontanaindifferentphenologicalcategoriesin have shown a lower density of predators and para- acerradoincentral Brazil. NL = newleaves, ML = matureleaves, MOL = mature and old leaves. A, First generation (January to sitoids duringthe dryseason. March);B,Secondgeneration(JunetoOctober). The nutritional quality of leaves varies among species andovertheirlifecycle, andyoungleavesgen- Over the whole year the phenological phase with erallyhave ahighercontentofnitrogenandwaterthan highest frequencywas ML (37%), NLwas next (36%) mature ones, which are more fibrous. Marquis et al. and the least abundant was MOL (27%). Therewas a (2001) showedthese trends for25 plant species in the significant difference amongthesevalues (%2 = 18.79, cerrado. Herbivores areaffectedbynutritionalquality, p = 0.0001). by the content ofwater and fiber, and by leaftough- Larvalphenologyand relationship withR. mon- ness (Coley & Barone 1996). Mature and old leaves tana. Overallwefound475larvaeon273ofthe3600ex- used by larvae ofthe first generation are physiologi- amined plants (7.58%). Among die plants used by die cally "younger" than those used by the larvae ofthe larvae, 4 (2%) belonged to the group with only NL. 43 second generation since tiiese leaves have been pro- (19%) on plants witii OL and 226 (79%) occurred on duced more recently. The leaves used bv the second plants MOLgroup. LarvaeofC.platysporawerefound generation could have a lower nutritional content and inonly8of12mondisanddierewasnooverlapbetween tiiis couldhave aninfluenceonthe duration ofthelar- dietwogenerations. Firstinstarlarvaeofdiefirstgener- val development. Foliar analyses of R. montana ationwerefoundinJanuaryanddevelopeduntil March, (Medeiros & Haridasan 1985) showed higherconcen- whendieypassedtodiepupal stage. The second larval trations ofK andPandlowerconcentrationsofAl, Mg generation began in June and extended into October. and Ca in younger leaves compared to older leaves. Thus, thelarvalphasewas longerindie dryseason than Additionally, the larvae that develop in the diyseason inthewetseason. Indie firstgeneration, 70larvaewere face more extremeclimatic conditions, such asdie ab- found on 46 ofthe 900 inspected plants (5.1%) and all sence ofrain, andlowrelative humidity, as well as the individualswith larvaewere members ofthe thirdphe- lowesttemperatures ofdievear. nological group (Fig. 3A). In the period ofthe second The slower larval development observed in die dry generation,405larvaewerefoundon227of1500exam- seasonversusdiewetseasonisnotexclusivetoC.platy- inedplants (15.1%) andwere present on all tiiree phe- spora. Indie same studyarea, similargrowthrateswere 294 Journalofthe Lepidopterists' Society recorded for larvae of Cerconota achatina (Zeller) Bendicho-Lopez,A. C. &I. R. Diniz. Inpress. Lifehistoryand (Elachistidae),whichfeedsonByrsonimacoccolobifolia, immature stages ofChlamdastis platyspora (Elachistidae). J. Lepid. Soc. B. pachyphylla (-B. crassa) and B. verbascifolia (Mal- COLEY, P. D. &J.A. Barone. 1996. Herbivoryandplantdefences phigiaceae). Generally, larvaeofthis speciescollectedin intropicalforests.Ann.Rev.Ecol. Syst.27:305-335. thedryseasonandraisedinthelaboratorytooktwicethe Diniz, I. R. & H. C. Morais. 1995. LarvasdeLepidopteraesuas plantas hospedeiras em um cerrado de Brasilia, DF, Brasil. timetodevelopasthosecollectedduringthewetseason Revta. Bras. Entomol.39:755-770. (Moraisetal. 1999). Hereabioticvariabilitywasreduced Diniz,I. R., H. C. Morais&A.J.A.Camargo. 2001. Hostplants sodifferenceswereduetodifferences inleafquality. oflepidopterancaterpillarsinthecerradoofthe DistritoFed- eral. Revta. Bras. Entomol.45:107-122. The numberoflarvae in the second generationwas Franco,A.C. 1998. Seasonalpatternsofgasexchange,waterrela- seven times higher than that of the first generation. tions and growth ofRoupala montana, an evergreen savanna This result is similar to that in another study of C. species.Pi. Ecol. 136:69-76. Joly, A. B. 1993. Botanica: Introducao a taxonomia vegetal. lla platyspora larva on R. montana done by another col- edicao,(SaoPaulo: EditoraNacional). 777pp. lector(B. Cabral, unpublished). Theseresultsobtained Marquis, R. J„ I. R. Diniz & H. C. Morais. 2001. Patterns and forC.platyspora coincidedwiththeseasonalpatternof correlatesofinterspecificvariationinfoliarinsectherbivoryand pathogenattackinBraziliancerrado. Trop.Ecol. 17:127-148. the lepidopteran larvae established for the cerrado by Medeiros,R.A.&M.Haridasan. 1985.J.Seasonalvariationsinthe Morais et al. (1999), who showed a largest proportion foliarconcentrationsofnutrients insomealuminiumaccumu- ofplantswith larvae in MaytoJuly (dryseason). latingandnon-accumulatingspecies ofthe Cerrado regionof centralBrazil. PlantSoil88:433-^36. Our results showed the close relationship of C. Mendonqa,R.C, M.Felfili,B. M.T.Walter, M.C. DaSilva platyspora larvae with leafphenology ofR. montana. Jr., A. V. RezeJn.de, T. Filgueiras & P. E. Nogueira. 1998. This associationappearstoaffectthesizeofthepopula- Floravasculardocerrado,pp. 290-456.In Sano, S. M. &S. P. Almeida(eds.),Cerrado: ambienteeflora. Brasilia: Planaltina, tionsofbothgenerationsandcanexplainthelowoccur- DF, Brasil. rence oflarvae on its hostplant, in spite ofthe high lo- Morais, H. C, I. R. Diniz& L. Baumgarten. 1995. Padroes de cal density of R. montana. The limiting resource are eprmoduumcacoerdreadfooldheasBreassiuliaa.utRielvitzaa.caoBrpaos.rBloatr.vas18d:e16L3e-p1i70d.optera plants belonging to the third phenological group (ma- Morais,H.C,I.R. Diniz&D. M.S. Silva. 1999. Caterpillarsea- ture andoldleaves) atthe time ofoviposition (Mayand sonalityin acentral Brazilian Cerrado. Revta. Bras. Entomol. December). Thus, the proportion ofindividuals ofR. Mora4n7,:10N2.5-&10W33.D. Hamilton. 1980. Lownutritive qualityas a mthoentlaownaocbceuarrrienngcdeifoffetrheinstslpeeacfiaplhiestnolaprhvaasienstchaenceerxrpaldaoi.n Paleadreif,enLc.eMa.ga&insFt.hAe.rbMi.voSreasn.toJ.s.The1o9r9.8.BiPola.pe8l6:d2o47i-n2d5u4.mentapi- loso na protecao contra a herbivoria em Miconia albicans Acknowledgments (Melastomataceae).Revta.Bras. Biol.58:151-157. Pullin, A. S. & L. E. Gilbert. 1989. The stingingnettle, Urtica Wewouldlike tothankDr. VitorO. Becker,whoidentifiedthe dioicaincreasestrichomedensityafterherbivoreandmechani- mothspecies,offeredinformationandbibliographyforthecomple- caldamage.Oikos54:275-280. tionofthiswork. Dr. HelenaC. Morais (Universidadede Brasilia) Ratter, A. 1980. Notes on the vegetation of Fazenda Agua andDr. AldicirScariot(EMBRAPA) madevaluablesuggestionson LimpJa. (Brasilia,DFBrasil).DepartmentPublicationSeriesno. anearlydraftofthemanuscriptandmanyvaluablesuggestionswere 1,(RoyalBotanicalGarden, Edinburgh), 111pp. mFaaudsetobGyonacnalavneosnytmoookusthreevpiheowteorg.raMprh.sKCinAiPtiESKiptraoyvaimdaedanadM.MSr.. RatteDra,JS.ilAv.a,.S.2B00r0i.dgEeswtautdeor,prJ.elF.imRiinbaerirdoa,dTi.sAt.riBbuoircgaoesda&sMes.peR-. scholarshiptothefirstauthor. cieslenhosasdafitofisionomiacerradosentidorestritonosesta- dos compreendidospelobiomaCerrado. Bol. Herb. Ezechias Literature cited PauloHeringer5:5-4.3. AnalyticalSoftware. 2000. Statistix7.0.AnalyticalSoftware,Inc. Solosmoounrc,eB.dePn.sit1y9,81r.elRateisvpeonasbeunodfanacheo,st-asnpdecipfhiecnohleorgbyi.vorEecotloorgey- Tallahasse,333pp. C 62:1205-1214. Bendicho-Lopez, A. 2000. Biologia e historia natural de Chlamydastis platyspora (Lepidoptera, Elachistidae) no cer- rado de Brasilia. Masters Thesis in Animal Biology. Universi- Receivedforpublication 14October2002; revisedandaccepted10 dadedeBrasilia,Brasilia,DF. June2003.

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