: 108 Journalofthe Lepidopterists’ Society irrigated zones in association with naturalized weedy Scott,J.A. 1986.TheButterfliesofNorthAmerica.StanfordUniver- hosts (Shapiro, unpublisheddata). Itisnotknownifthis sityPress,Stanford,CA.583pp. Shapiro,A.M. 1997.Impactosantropogenicossobrelafaunademari- butterflyis native to the region or is itselfnaturalized; it posas (Lepidoptera, Rhopalocera) de PatagoniaaustralyTierra istheonlymemberofitsgenusinthe SouthernConeof delFuego.AnalesInstitutodelaPatagonia(PuntaArenas,Chile), SouthAmerica. sec. Cs. Nat.,25: 117-126. 2006. Useofanexoticweedasanovipositionsubstrateofthe . high-AndeanPieridPhulianymphula. Lepid.Soc.60: 100-101. Literature Cited J. AravPernoav,inRc.iOa.d1e98L3.aIPnasemcptao.s qVueCopenrgjruedsiociaFnoraesltaaflloArargleenntoisnao,deLlaa Arthur M. Shapiro, Centerfor Population Biology, Pampa,4:315-321. University of California, Davis, CA 95616; e-mail Pastrana, J.A. 2004. Los Lepidopteros Argentinos: Sus Plantas [email protected] Hospedadorasyotros Substratos Alimenticios. South American BioControlLab, USDAandSociedadEntomologicaArgentina, Receivedforpublicationl9February2008, revisedandaccepted3 BuenosAires.334pp. July2008. JournaloftheLepidopterists'Society 62(2),2008,108-110 COMMENTS ON LARVAL SHELTER CONSTRUCTION AND NATURAL HISTORY OF URBANUS PROTEUS LINN., 1758 (HESPERIIDAE: PYRGINAE) IN SOUTHERN FLORIDA. Additional keywords. Eggstacking,hostplant,oviposition,clutchsize. The Bean Leaf Roller (Urbanus proteus Linn.) is a Tllis observation prompted us to search foliage ofother common and widespread skipper (Hesperiidae) found V. luteola plants, and resulted in the discovery of 26 from southern United States south to Argentina (Smith additionalclutches ofhatched andunhatchedeggs. et al. 1994). Early observations on its natural history At hatching, larvae consume only the top portion of (Seudder 1889) have been supplemented with details the eggs (pers. obs.), and we were able to use the from various parts of its range (Greene 1970, 1971a; remainingegg fragments to determine clutch size from Kendall 1965; Moss 1949; Riley 1975; Skinner 1911; all27clutches (mean = 2, SD = 1.1, range = 1-5). Most Smith et al. 1994; Young 1985), particularly in Florida clutches were located on the under surface of mature (Quaintance 1898),where it is apest onleguminaceous leaves (n = 24), butoccasionallyon leafpetioles (n = 3). crops (Green 1971b; Quaintance 1898; Watson &Tissot Withinaclutch, eggswereplacedadjacentto (touching) 1942) and where there exist documented seasonal or up to 1 mm from other eggs. One exception was a movements (Urquhart & Urquhart 1976). Like most clutch ofthree eggs found stacked endto end such that skippers (Greeney & Jones 2003), the larvae of U. only the bottom egg was attached to the leaf surface proteus construct and live in shelters made from the (Fig. 1). Similarly, Quaintance (1898) reportedaclutch leaves of the food plant, but only two authors have size of1-6andnotedthateggswere frequentlylaidin a described or pictured these shelters in any detail stackedfashion, 3-4 eggs high. Young (1985), however, (Quaintance 1898; Young 1985). In fact, detailed recordedonlysingle eggclutches in Costa Rica. knowledge of larval shelter construction for most In addition to the eggs, we found a total of50 larvae skippers is weak or nonexistent for all but one widely representing the following instars: 36 first, 8 second, 3 distributed North American species, Epargyreus clams third, 2 fourth, and 1 fifth. We removed larvae from Cramer, 1775 (Jones et al. 2002; Lind etal. 2001;Weiss theirshelters and carefullydeterminedtheir ages using et al. 2003). As shelters may prove useful in resolving the prior experience ofHFG with the larvae ofrelated phylogenies (Greeney & Jones 2003), here we present species. We also watched as 3 first-instars constructed our observations of shelters from a population of U. new shelters after removal from their original shelters. proteus in southern Florida. By examining shelter construction and comparing our We made observations at Burns Lake Campground observations to previously constructed shelters, we (25°53’N, 81°13’W) in Big Cypress National Preserve, determined that larvae build 3-5 shelters as they Collier County, Florida. On 30 December 2005, at develop, and that these belong to three shelter types. 14:15, we observed a female U. proteus ovipositing on First through third instars were found inside shelters the under surface of a leaflet of Vigna luteola (Jacq.) built by excising a small triangular portion of the leal Bentham (Leguminaceae). She laid three dull yellow margin and creasing it into a tent-shaped lid (Greeney eggs in an evenlyspacedrow, andthen flewoutofsight. & Jones 2003; group III, type 10, two-cut stemmed Volume 62, Number 2 109 will prove useful in testing phylogenetic hypotheses (Greeney&Jones 2003) onlyifwe examine sheltersand their construction in much more detail than previously reported (but see Greeney & Warren 2003, 2004; Lind etal. 2001; Weissetal. 2003). Secondly, our observations, and observations oflate instars of other pyrgines (HFG unpublished) and coeliadines (Common & Waterhouse 1972), suggest thatthere maybe little difference between the “type 3" and“type 4” shelters distinguished by Greeney& Jones (2003); thesesheltertypesbeingdefinedbythe number ofleaves included in the shelter. In later larval stadia, U. proteus silks together two or more leaves or leaflets into a silk-lined pocket (this study, Quaintance 1898). Young (1985), however, observed only two leaves used We in late instar shelter construction. conclude, therefore, that the number and arrangement of the various leaf parts used is likely related to the relative sizeandshape ofthehostplantleavesratherthantoany innate shelterbuildingbehavior. In otherwords, larvae simplyspin silk, pullingleaves (orparts thereof) around themselvesuntiltheyaresufficientlycovered. Similarly, E. clarus shows variation in the number of leaves used Fig. 1. Three Urbanusproteuseggshellsfoundontheunderside ofaleaf. Burns LakeCampground,CollierCounty, Florida, Decem- in late instar shelters, varyingwith size ofthe hostplant ber2005. leaf (M. Weiss pers. comm.). Based on these shelters). These U. proteus shelters were, in faet, very observations , we suggest that “type 3” and “type 4” similar to shelters described for E. clarus (Weiss et al. shelters, as definedby Greeney & Jones (2003), should 2003), and would be considered the same type under be mergedinto one “multi-leaf’ sheltertype, regardless the classification of Greeney & Jones (2003). Like E. ofwhether the shelter includes two or more leaves or darns, the shelter cuts of U. proteus were always leaflets. oriented in adistinctfashion in relationto the leafbase; Finally, the three egg shells we found stacked end to the longest cut always being distal. The most obvious end showed a different emergence pattern than and consistent difference we found was the lack of a described in previous observations of lepidopteran “notch” in the cut closest to the leafpetiole in shelters opposition. Several species of the nymphalid genus built by U. proteus larvae (Fig. 2). Earlyinstar shelters Hamadryas Hiibnerare alsoknown to deposit eggs one were still “tented” into a distinct peak, however, by on top ofanother, sometimes in chains of more than 10 pinching together (using multiple silk ties) a small eggs (Muyshondt & Muyshondt 1975b, 1975c). To section along the margin ofthe shelter lid. The result was a shelter similar in appearance to that built by E. clarus, but arrived at by slightly different means (ie. without the notch). Fourth instars were found, one each, inside a shelter created by silking two leaves together (group I, type 4, two-leaf shelter) and one formed by silkingseveral leaves together (group I, type 3, multi-leafshelter). We found the single fifth instar feedingadjacenttoseveralleaves silkedtogether(group I, type 3 shelter) at around 17:45. Our observations bring to light several important Fig. 2. Comparisonofcutpatternsforfirstinstarsheltersof(a) aspects ofegglayingandlarvalshelterbuilding. Firstly, Epargyreusclarus(redrawnfromWeissetal.2003)and(b) Urbanus species building superficially similar shelters may use proteus. Largearrowpointstowardsthebaseofthefoodplantleafto showorientation ofshelters. Small arrows pointto (a) position of slightlydifferentcutpatternsorconstruction techniques notchmadebyE.clarustoaidintentingtheshelterand(b)position to arrive at the finished product. Therefore, shelters ofsilklaiddowntopinchshelterintoatentedpeakbyU. proteus. 110 Journalofthe Lepidopterists’ Society emerge from the eggs, Hamadryas larvae create an Jones, M.T., I. Castellanos,6cM. R.Weiss.2002. Doleafshelters openingin the side ofthe egg. Previous discussions on alwaysprotectcateqrillarsfrominvertebratepredators?Ecol. En- tomol.27:753-757. patterns of egg laying and larval emergence in Kendall, R. O. 1965. Larval foodplants and distribution notes for nymphalids suggest an evolutionary signifigance to the twenty-fourTexasHesperiidae. Lepid. Soc. 19: 1-33. correlation between side emergence and egg stacking: Lind,E. M., M.T.Jones, J. D. LongJ,. &M. R.Weiss.2001. Ontoge- neticchangesinleafshelterconstructionbylarvaeofEpargyreus side emergencebeingnecessarytoavoiddamagingeggs clams(Hesperiidae),thesilver-spottedskipper. Lepid.Soc.54: laid above (Muyshondt & Muyshondt 1975a). Our 77-82. J. observation of egg stacking in U. proteus showed Moss, A. M. 1949. Biological notes on some “Hesperiidae” ofPara andtheAmazon(Lep. Rhop.).ActaZool. Lilloana7:27-79. emergence from the top, suggesting that emergence Muyshondt, A. &A. MuyshondtJr. 1975a. Notes onthelifecycle from the side of the egg is not a necessary adaptive and natural history of El Salvador. IB. - Hamadryasfebrua responsetoeggslaidinstacks. Whilewewereunableto (Nymphalidae-Hamadryadinae). J. NewYork Entomol. Soc. 83: 157-169. clearlyillustrate top-emergence in Figure 2, our direct . 6cA. Muyshondt Jr. 1975b. Notesonthelifecycleandnatural observations showthatthiswasindeedthecase. Figure history of El Salvador. II B. - Hamadryas guatemalena Bates 2 also shows that the eggs of U. proteus were not laid (Nymphalidae-Hamadryadinae). J. NewYorkEntomol. Soc. 83: 170-180. dMiuryecsthloyncdetnter&ed aMbuoyvsehtohnedetggb(e1l9o7w5,b,as i1l9lu7s5tcr)atedfoinr hi.s6tcorAy. MofuyEslhoSnadltvadJro.r1.97I5lcl.BN.ot-esHoanmtahderlyiafescyacmlephaindnonmateuraL.l Hamadryas. It is possible that this means ofattaching (Nymphalidae-Hamadryadinae).J. NewYork Entomol. Soc. 83: 181-191. stacked eggs represents an alternative adaptation Quaintance, A. L. 1898. Three injurious insects. Bean leaf-roller, allowingeggs to be laidin stacks. Corn delphax, Cannaleaf-roller. FloridaAg. Exp. Sta. Bull. 45: 53-71. Acknowledgements Riley, N. D. 1975.AfieldguidetothebutterfliesoftheWestIndies. DemeterPress, Boston,Massachusetts. ingWoeurtvhiasnitktDoiFalnoeriadan.dTTehrerywSohreklodfonHfForGtihseisruphopsoprittealditiyndpuarr-t ScudCdaenra,dSa.wHi.th18s8p9e.ciTalherebfuetrteernfcleietsooNfetwheEenagsltaenrdn.UPnuibtleidsShteadtebsyatnhde by the Hertzberg Family Foundation, the Population Biology author,Cambridge, Massachusetts. Foundation, Nature 6c Culture International, and a Rufford Skinner,H. 1911.ThelargerborealAmericanHesperiidae,including SmallGrant. The PBNHScontinuestoprovideinspirationand Eudamus, Erycides, Pyrrhopyge and Megathymus. Trans. Am. supportforournaturalhistoryresearch. Theclarityandcontent Entomol. Soc.37: 169-209. ofthis manuscriptwere greatlyimprovedbythesuggestions of Smith, D. S., L. D. Miller, & Y. Miller. 1994.Thebutterfliesof Annette Aiello and MarthaWeiss. This is publication number theWestIndiesandsouthFl|orida.OxfordUniv.Press,NewYork. 96oftheYanayacuNaturalHistoryResearchGroupandisded- Urquhart, F. A. & N. R. Urquhart. 1976. Migration ofbutterflies icatedtoDanielSheldonforhisencouragementofK. Sheldon’s alongtheGulfCoastofnorthernFlorida. Lepid.Soc.30:59-61. academicandscientificpursuits. Watson,J.R.6cA.N.Tissot. 1942.InsectsanJd. otherpestsofFlorida vegetables. FloridaAg. Exp. Sta. Bull.370:33-35. Literature Cited Weiss, M. R., E. M. Lind, M.T. Jones,J. D. Long, 6c J. L. Maupin. CGroememtnroaeln,i,a.GIA..nFLg..uBs.19&&70R.Do.bHFee.ratWdsaotmneerPauhsboulu.r,seemL.eonn1td9so7n2a.,nTdU.hweeK.ibguthttesrfolfietshoefbAeuasn- Young2Bgy0e,r0h3eaA.uv..sUMc1n.6li:fa1o4m9r6s8m55i-(.tH4yeN8sa0opt.feurrliaeialdfahseih)se,tlottrehyrencosotinelsvsteroru-ncsptAoistottnreadbpytskelisaprpavenarde.oUJ.frbIEnapsnaeurc-st leaf roller, Urbanus proteus (Hesperiidae). J. Lepid. Soc. 24: (Hesperiidae)inCostaRica. Lepid. Soc.39:215-223. 47-51. J. 1971a. Instardistributions,naturalpopulations,andbiologyof tir.ebeanleafroller. FloridaEntomol.54:213-219. Harold F. Greeney & Kimberly S. Sheldon, . 1971b.Economicdamagelevelsofbeanleafrollerpopulations Yanayacu Biological Station l? Center for Creative Greeonneys,naHp.bFe.an&s.M.].TE.coJonn.esE.nt2o0m0o3l..S6h4e:lterbuildingintheHesperi- Studies, Cosanga. Ecuador do 721 Foch y Amazonas, idae: aclassificationschemeforlarvalshelters. Res. Lepid. 37: Quito, Ecuador; email: [email protected]. J. 27-36. 6cA. D.Warren.2003. NotesonthenaturalhistoryofEantis . thraso (Hesperiidae: Pyrginae) in Ecuador. Lepid. Soc. 57: 43-46. J. Receivedforpublication 18August2007;revisedandaccepted5 &A.D.Warren.2004.Naturalhistoryandshelterbuildingbe- December2007 . havior ofNoctuana haematospila (Hesperiidae) in Ecuador. J. Lepid. Soc.59:6-9.