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Distribution and hostplant records for Eupackardia calleta from southeastern Texas with notes on mandibular morphology of Attacini (Saturniidae) PDF

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Preview Distribution and hostplant records for Eupackardia calleta from southeastern Texas with notes on mandibular morphology of Attacini (Saturniidae)

Journal of The Lepidopterists' Society Volume 53 2000 Number4 JournaloftheLepidopterists'Society 53(4),1999,133-137 DISTRIBUTION AND HOSTPLANT RECORDS FOREUPACKARDIA CALLETA FROM SOUTHEASTERN TEXAS WITH NOTES ON MANDIBULAR MORPHOLOGYOF ATTACINI (SATURNIIDAE) Valerie A. Passoa 10603BrettridgeDrive,Powell,Ohio43065,USA AND Steven Passoa1 USDA/APHIS/PPQ,TheOhioStateUniversity,MuseumofBiologicalDiversity, 1315KinnearRoad, Columbus,Ohio43212,USA ABSTRACT. Eupackardiacalleta(Westwood)isrecordedonprivet(Ligustrum)fromKingwood,Texas.Thisnewcountyrecordrepresents anortheasternrangeextensionof100milesandconfirmsprivetasahostplantundernaturalconditions.Theroleofthemandibleindigestion ofthehostplantisdiscussed. ThelastinstarlarvalmandibleofmostAttacinieitherlacksteethorhasthecuttingedgebluntlyserratedwithreducedteeth.Incontrast,the distinctivemandibleofE. calletacontainsthreelargeteethanddeepmolarridges,anautapomorphyforthegenus.Well-developedmandibu- larteetharepresentonthelastinstarofafewunrelatedSaturniinaeandCeratocampinae;thereforethischaracterishomoplasticinSaturni- idae.Astructurallycomplexmandibleofthe"sphingid-type"occursinseveralsatumiidssuchasAntheraeapernyi(Gu£r.-Men.)(Saturniidae). Thismandibletypeisillustrated. TherearetwoontogeneticpatternsofmandibulardevelopmentinSaturniidae. Inonecase,teetharepresentinthefirstinstar,thenlostin latermolts.ThemandibulardevelopmentofE. calletarepresentsanalternativescenariowhereteetharepresentthroughoutthelarvalstage. Additionalkeywords: Ligustrum,ontogeneticdevelopment,Sphingidae,Notodontddae. Eupackardiacalleta (Westwood) is amemberofthe morphology to other Attacini, with emphasis on re- Attacini, usually considered the sister group to Roth- lated nearctic taxa. Ontogenetic and phylogenetic ob- schildia (Ferguson 1972, Peigler 1989, Friedlanderet servations are alsoincluded. al. 1998). This species is distributed from Central Materialsand Methods America (Honduras, Guatemala) northtoArizona and southernTexas (VictoriaCo., Calhoun Co.) (Ferguson In order to survey saturniid mandibles, preserved 1972, Wolfe 1995, Tuskes et al. 1996). The systematics larval specimens or exuviae associated with cocoons andbiologyofE. calletahavebeenreviewedbyseveral and emerged adults were taken from the authors' col- authors including Ferguson (1972), Weast (1989), lection and dissected. The mandible was either slide Miller(1976), Lemaire (1978),andTuskesetal. (1996). mounted in Euperal or preserved in alcohol. For Weast (1989) suggestedthatsystematicstudies ofE. analysiswithaJ.O.E.L. JSM 820scanningelectronmi- calleta are needed. Tuskes et al. (1996) illustrated croscope, each samplewas attachedto astubwith car- threelarvalformsofthe UnitedStatespopulations,but bon paint and coated with a thin conductive layer of did not study their mandibular morphology. In this gold-palladium. Mandibular terminology is based on paper we present biological data on E. calleta from Godfrey (1972). southeastern Texas, then compare the mandibular Material examined: Eupackardia calleta: TEXAS (variouslocalities, exovafromfemale mothsinlabcul- 1Addresscorrespondencetothisauthor ture): 10 eggs, 10 first instars, 27 second to fourth in- 134 Journalofthe Lepidopterists' Society Table1. MandibularmorphologyofselectedlateinstarAttacini as an artificial laboratoryhost (Ferguson 1972,Tuskes larvaeoccurringintheWesternHemisphere.Localitydatadesignate etal. 1996). Oursisthe secondpublishedrecordforE. sourceofthestudyspecimens. calleta eating ornamental privet under natural condi- Source Cuttingedgeofmandible tions, supporting the suggestion ofWeast (1989) that Callosamiaangulifera (Wlk.) USA Teethreduced,margin privetisanimportanthostplantofthisspeciesnearur- bluntlyserrate ban areas of Texas. In fact, E. calleta appears to be Callosamiapromethea (Drury) USA Teeth reduced,margin oligophagous on several generaofOleaceae, including bluntlyserrate Callosamiasecurifera(Maas.) USA Teethreduced,margin privet in Texas (present study) and Mexico (Peigler bluntlyserrate pers. comm.), Fraxinus greggii A. Gray in western Eupackardiacalleta (Ww.) Texas, Teethpresent,well- Texas (Peiglerpers. comm.), andForestieraangustifo- Mexico developed Hyalophoracecropia (L.) USA Teethabsent lia Torrey (Stone 1991). A mature larva of£ . calleta HyalophoraColumbia USA Teethabsent onprivetfrom Kingwood,Texas, isillustratedin Fig. 1. Hya(Sl.oIp.hSomriathe)uryalus(Bdv.) USA Teethreduced,margin At maturity, the larva lacks long and dark scoli which bluntlyserrate are characteristic ofearlierinstars (Fig. 2). In spite of Hyalophoragloveri (Stkr.) USA Teethreduced,margin the availability ofceniza nearby, E. calleta was found bluntlyserrate onlyon privetin the seniorauthor's backyard. Prob.Rothschildiacincta Prob.Baja Teedireduced,margin (Tepp.) Cali- bluntlyserrate Examination of E. calleta mandibles from Texas fornia showed thatthese structures are atypical comparedto Rothschildiaerycina (Shaw) USA Teethreduced,margin other saturniids described in the literature. Bernays bluntlyserrate Rothschildiaforbesi Benj. Texas Teethabsent and Janzen (1988) characterized late instar saturniid Rothschildialebeau Honduras Teethabsent mandibles as "short, with a broad base, and without (Guer.-Men.) Rothschildiaorizaba (Ww.) Ecuador Teethreduced,margin obvious teeth". This definition does not fit E. calleta bluntlyserrate whichhaslargeteeth inboththeearly(Fig. 3) andlast Samiacynthia(Drury) USA Teethreduced,margin instars (Figs. 4, 5, and 6). bluntlyserrate Except forE. calleta, members ofthe Attacini typi- cally lackwell-developed mandibular teeth in the last instar or have the cutting edge ofthe mandible ser- stars, 6pupalcases (with larvalexuviae), det. V. A. Pas- ratedwith bluntlobes (Table 1). Mandibularteeth are soa, mandible slides #581, 582 S. Passoa collection; also absent in mature larvae of two Indo-Australian Harris Co., Kingwood, XII-1995, on Ligustrum, V. A. Attacini, Attacus atlas (L.) (Heppner et al. 1989) and Passoa, 6 mature larvae, det. V. A. and S. Passoa, Coscinocera hercules (Miskin) (specimens in Passoa mandible slide#616 S. Passoacoll. collection). MEXICO: various localities, collection data un- Ourdataon saturniid mandibles agreewith results known, 2firstinstars, 18 secondtofourthinstars, 3pu- of a survey of notodontid mandibles published by pal cases (withlarval exuviae), det. V. A. Passoa; Mich. Godfrey et al. (1989) in several respects. Last instar [Michoacan], Urupan (sic) [Uruapan], 2-X-1941, coll. mandibles of the Notodontidae usually lack teeth, [D. M.] Delong, det. V. A. and S. Passoa (1 last instar except for a few apparently unrelated exceptions. larva with mandibles slide mounted (The Ohio State This is also true in Saturniidae. Outside of the Universitycollection). Attacini, mature larvae ofActias selene (Hbn.), A. One to ten mandibularpairs ofeach species (Table luna (L.), Argema mittrei (Guer.- Men.) (Saturni- 1)were examined, dependingon material available. inae) and Citheronia regalis (F) (Ceratocampinae) (specimens in Passoa collection) have well-devel- Results and Discussion oped mandibular teeth, so the presence ofteeth on DuringNovember 1995,the seniorauthorcollected saturniid mandibles is homoplastic across several seven thirdinstarlarvae ofE. calleta onprivet {Ligus- subfamilies. trum) from her backyard in Kingwood, Harris Co., Largeteeth alsooccurintheAntheraea genus com- Texas. This is a northeastern range extension of ap- plex, forexample,Antheraeapernyi (Guer.- Men.) and proximately 100 miles (161 km) and a new county its purported synonym A. hartii (Moore), A. polyphe- record. Itremainstobe determinedifE. calleta isnow nols (Cram.), and Opodiphthera eucalypti (Scott) establishedin Harris County. (spms. in Passoacoll.). In addition, the mandible ofA. Tuskes et al. (1996) considered Leucophyllum pernyi is unusual because it matches the "sphingid- frutescens (ceniza, purple sage) to be the main host- type"as definedbyBernays andJanzen (1988). Sphin- plant forE. calleta in Texas. Privetis normallytreated gidmandibleswerecharacterizedasbeing"ridgedina Volume 53, Number4 135 Figs. 4-6. Mandible oflast instar Eupackardia calleta (West- mm wood),scaleline= .10 inallfigures.4,oralsurface,ventralview, Figs. 1-3. Eupackardia calleta (Westwood) from Texas. 1, late 70x. 5,teethanddeepmolarridges,ventralview, 150x. 6,teethon instarlarvaonprivet(lateralview).2,mid-instarlarvaonprivet(dor- cuttingmargin,dorsalview,95x. salview). 3, earlyinstar mandible, oral surface, ventral view, pho- tographsunderpolarizedlight, lOOx. 136 Journalofthe Lepidopterists' Society variety ofcomplex ways". The mandible ofA. pernyi (Figs. 7, 8, and9) has aseries ofcomplexridges onthe dorsalsurfacethatappeartoformoverlappingteeth, in additionto several smallertoothlikeprojections onthe oral surface. The mandibles ofE. calleta andA. pernyi suggestitiswisertocharacterize saturniidmouthparts with ecological orphysiological criteria (cuttingversus grinding), instead ofusing aphylogenetic approach at the family level as did Bernays and Janzen (1988) in theirstudyofsaturniid and sphingidmandibles. There appeartobetwoontogeneticpatterns forsat- urniid mandibles. In one pattern, first instar larval mandibles have teeth; subsequently these teeth are lost in later molts. This pattern is typical for Costa Ri- & can saturniid species in four subfamilies (Bernays Janzen 1988). A second pattern is found in E. calleta whereearlyinstars have mandibularteeth (Fig. 3) that are retained throughout larval life (Figs. 4, 5, and 6). This dichotomywas alsonotedbyGodfreyetal. (1989) innotodontidgenera. Godfreyet al. (1989) stated that characterization of certain taxabymandibularmorphologywas "problem- atical"becausein somespeciesthe mandibularmargin was intermediate between toothed and smooth. Inter- mediate conditions are common with Saturniidae, as showninTable 1 bytheterm "marginbluntlyserrate". A bluntlyserrated mandible has round conicalprojec- tions that resemble small teeth, or an irregular man- dibular margin with indentations. Callosamia is an ex- ample ofthe first condition where from 10-15 small teeth coverthe cutting margin ofthe mandible. Irregular mandibular margins are found in Hyalo- phora and Rothschildia where it is difficult to discern teeth, butthe mandibularmarginisnotstraight. Dock- ter (1993)warnedthat mandibularwearmustbe taken into account when describing mandibles. It is unclear whether bluntly serrated teeth in Saturniidae are a transition state between toothed and smooth man- dibles, asnotedbyGodfreyetal. (1989) innotodontids, or instead represent worn mandibular teeth as de- scribedby Dockter (1993) in Heterocampa and Soura- kov (1996) in larvae ofsatyridbutterflies. Examination of unworn mandibles on freshly molted larvae are neededtoresolvethisquestion. Thus,itmaybeprema- ture to characterize saturniid mandibles as usually Figs. 7-9. Mandible of last instar Antheraea pernyi (Guer.- toothless (Bernays &Janzen 1988) until more ontoge- Men.). 7,complexpatternofteethondorsalandoralsurfaces,ven- netic studies arepublished. tralview,27x,scaleline= 1 mm. 8,overlappingteethofdorsalsur- In summary, the mandibular shape of Macrolepi- face,ventralview,33x,scaleline= 1mm.9,toothofdorsalsurface, ventralview,70x,scaleline= .1mm. dopteradepends onseveralfactors includingheadsize and food toughness (Bernays 1986), foodparticle size and chemistry (Bernays & Janzen 1988), the need to piercethechorionoftheeggateclosion, andaneedto Volume 53, Number4 137 seal the oral cavity during ingestion (Godfrey et al. Texas. We are grateful to K. S. Johnson (Ohio University, Athens, 1989). Unlike notodontids which use toothed man- Ohio) for Callosamia larvae and pupae, K. L. Wolfe (Escondido, California)foraR. erycinalarvalexuvium,C.A.Conlan(SanDiego, dibles to pierce the leafepidermis (Dockter 1993), E. California) forE. calleta pupal cases, M. D. Schmidt (Springboro, calleta does notskeletonizeleaves duringanyinstar(V. Ohio) for a H. euryalus pupa, and R. S. Peigler for use of his A. Passoapers. obs.). Therefore, it seems likelythat a OleAatcetaheehTohsetpOlhanitorSetcaotredsU.niversity(Columbus,Ohio),wethank toothed mandible is required forotherreasons. C. Mitchellforthescanningelectronmicroscopephotographs,D.J. J. Bernays andJanzen (1988) noted that larvae which Horn and G. D. Keeneyforgreenhouse space usedto growhost- feed on toxic plants tend to be associated with plants,andN. F.Johnsonforuseoftheinsectcollection. mandiblesthatcontain teeth. They(Bernays &Janzen TheVoOuhcihoerStsapteecUinmievenrssiotfyE'.scIanlsleecttaCaonldleitcstihoosntaplnadntHearrbeadreipuoms.itedin 1988) suggested that complete mastication ofthe leaf tissue allows more complete digestion, a strategy not Literature Cited possiblewithhostplants containingtanninsthatinhibit Rernays, E.A. 1986. Diet-inducedheadallometryamongfoliage- digestive efficiency. However, the digestive physiology chewinginsects and its importance forgraminivores. Science 231:495^96. ofE. calleta maybe more complicatedthan other Sat- Rernays, E. A. & D. H. Janzen. 1988. Saturniid and sphingid urniidae. Caterpillars ofE. calleta secrete a pungent caterpillars:twowaystoeatleaves.Ecology69:1153-1160. liquid containing phenolic compounds and biogenic Deml, R. & K. Dettner. 1993. Riogenic amines and phenolics amines from their scoli when disturbed, but it is un- c(hLaerpaicdtoeprtiezrea:tShaetudrenfieindsaei)v:easceocmreptairoantiovfessattuudyr.niidCocamtper.piPlhlyasr.s knownifplanttoxins aresequesteredforthis secretion R 163:123-132. J. (Demi& Dettner 1993). Untilthemetabolismandde- Dockter, D. E. 1993. Developmentalchangesandwearoflarval fensive chemistry ofE. calleta are clarified, it maybe m(aNontdoidbolnetsidaien).HetLeepr.ocSaomc.pa47:g3u2tt^i8v.itta and H. subrotata J. premature to assume mandibular teeth are an aid to Ferguson,D.C. 1972. In Dominick,R.B.etal.(eds.),Themoths coping with plant toxins. Nevertheless, E. calleta lar- ofAmericanorthofMexico. Bombycoidea. Saturniidae. Fasci- vaeBedsoiduetislitzheetonxeicedplatnotsun(dWeerassttan19d89m)a.ndibular mor- FriecPdleleia2gn0ld.e2e.rr,2&7T5Q..pPp.Q.,.,K2F.a2nRcg.o.lHoor1r9es9dt8p,.isTJ..wCo.nRucelgeiaerr,geCn.esMyiitetledrc,onRc.orS-. phology in an ecological context, it should be noted dant relationships within Attacini (Lepidoptera: Saturniidae). Mol.Phyl. Evol.9:131-140. thatasurveyofsaturniidmandibleswouldaididentifi- Godfrey,G. L. 1972. Areviewandreclassificationoflarvaeofthe cation ofthe immatures. Minet (1994:70) mentioned subfamily Hadeninae (Lepidoptera, Noctuidae) of America mandibular secondary setae as an apomorphy ofthe northofMexico. U. S. D.A.Tech. Bull. No. 1450. Lasiocampidae, althoughthese setae are alsoindepen- Godfpraeryt,moGd.ifLi.c,aJt.iSo.nsMiinlllaerrval&NDo.toJd.oCnatritdeare.(L1e9p8i9d.opTtweroa)m:outhtehi-r dently evolved in other macrolepidopteran families. taxonomicdistributionsandputativefunctions. NewYorkEn- J. Mandibularsecondarysetae arepresent (Agapema) or tomol. Soc.97:455-470. Heppner.J. B.,H.Y.Wang&Y.C. Sen. 1989. Larvalmorphology haabssepnotte(nEt.iaclallfeotra)idienntSiafitcuartniioindapeu,rptohsuess.thiMsacnhdairbaucltaerr oMfuTsaeiuwamn4S2a:t8u9r-n9i7l.idae[sic]:Attacusatlas(Linnaeus).J.Taiwan teeth associated with the cast larval exuvium can be LeMaire, C. 1978. Les Attacidae Americains. 1. Attacinae. used to separate cocoons ofE. calleta (teeth present) MillNeeru,ilTl.yA-.sur1-9S7e6i.ne,ObFsrearnvcaet.ioPnusbloifsEhnepdabcykatrhediaautchaolrl.et2a3i8npspo.uth- andRothschildia (teethabsent). Bothgeneraare easily ernTexas(Saturniidae). Lep. Soc.30:127-130. J. confused due to their similarpupal morphology. A fi- Minet,J. 1994. The Bombycoidea: phylogenyandhigherclassifi- cation(Lepidoptera: Glossata). Entomol. Scand.25:63-88. nal example is A. luna andA. pohjphemus. Although Peigler,R. S. 1989. ArevisionoftheIndo-AustraliangenusAtta- these genera are often confused as larvae (Ferguson cus.TheLepidopteraResearchFoundation,Inc. BeverlyHills, 1972:207),theirmandiblesarecompletelydifferent,be- California. 167pp. SOURAKOV,A. 1996. NotesonthegenusCalisto,withdescriptions ingeithersimpleandtoothed(A. luna) orcomplexwith ofthe immature stages (part 1) (Lepidoptera: Nymphalidae: ridges (A. polyphemus). Moredescriptivestudiesofsat- Satyrinae).Trop. Lepid. 7:91-112. urniidmandibleswillno doubtyieldfurtherexamples. Tuskes,P.M.,J.P.Tuttle,&M. M.Collins. 1996. Thewildsilk mothsofNorthAmerica.CornellUniversityPress,Ithaca,New Acknowledgments York.250pp. Weast, R. D. 1989. Saturniidae. Ecologicalandbehavioralobser- Reviews by G. L. Godfrey (Haskell Indian Nations University, vationsofselectAttacini.Self-published.Johnston,Iowa.53pp. Lawrence,Kansas),R.S. Peigler(UniversityoftheIncarnateWord, Wolfe, K. L. 1995. Photo Contest-1995. Enpackardia calleta. SanAntonio,Texas),J. RTurtle(Tucson,Arizona),andR.D.Weast Trop.Lepid. 6:iv(fig.8). (Johnston, Iowa) significantlyimprovedthemanuscript.W. D. Ol- hausen (JesseJonesParkandNatureCenter, Humble,Texas)pro- Receivedforpublication 11 May1997;revisedandaccepted10De- vided keen observations concerning E. calleta in Harris County, cember1999.

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