Journal of the Lepidopterists' Society 57(1 ),2003, 43-46

NOTES ON THE LIFE HISTORY OF THRASO (HESPERIIDAE: PYH.GINAE ) IN ECUADOR

HAROLD F. GREENEY Yanayacu Biological Station and Center for Creative Studies , Cosanga, Ecuador c/o Carrion No. 21-01 y Juan Leon Mera, Quito, Ecuador

AND

ANDREW D . WARREN] Department of Entomology, Oregon State University, Corvallis, Oregon 97331-2907, USA

ABSTRACT. The early stages and larval behavior of Eantis thraso (Hubner) in eastern Ecuador are described. All larval ins tars were found to build leaf shelters on the host plant, which are described and illustrated. A summary of larval host plants for in the Eantis Bois­ duval is provided. Cultivated Citrus L. species are repOlted as the local larval host plants for E. thraso in eastern Ecuador. Additional key words: larvae, larval food plants, larval shelters, life cycles, pupae.

Warren (1996) resurrected the generic name Eantis mature larvae of E. thraso as being '''plain green with a Boisduval from synonymy with Hubner for rotund brown head." Here, we provide a more detailed taxa including species and subspecies of Evans' (1953, description of the early stages of E. thraso in Ecuador, 1955) Fab. group, after a phylo­ as well as notes on its larval shelter-building behavior, genetic analysis of morphological characters observed in Achlyodes. The genus Eantis is entirely Neotropical MATERIALS AND METHODS in its distribution and includes 9 superficially similar The majority of observations were made between species. Material studied by Warren (1996) indicated a September and November of 1997 in north-eastern geographical range for E. thraso from extreme south­ Ecuador at the La Selva Lodge Research Station, 75 ern Chiapas, Mexico, east to southern Belize, and kilometers E.S.E. of Coca, Garzacocha, Sucumbios south throughout tropical Central and South America; Province, at 250 meters elevation. For a detailed de­ recent color illustrations of adult males appear in scription of this site, see DeVries et al. (1999). Subse­ Lewis (1987:80), Brown (1992:176) and Warren quently, several larvae were found and reared under (1996). Eantis thraso is replaced in most of Mexico similar conditions at the Sacha Lodge Research Sta­ and Texas by the similar E. tarnenund W H. Edwards, tion located 10 kilometers up river from La Selva. All Food plant records for Eantis are mostly from the larvae and pupae were collected from cultivated Cit­ , including cultivated Citrus L. plants rus trees located along the edge of seasonally inun­ (Panton 1898, Smyth 1919). Native hosts in most areas dated forest. Over 30 individual larvae of various in­ also appear to be Rutaceae, especially Zanthoxylum L, stars were encountered on the host plants and (Bruner et al. 1945, Poey 1832, Janzen & Hallwachs transferred to the lab for rearing. Head capsule width 2003). Wolcott (1923, 1951 ) described the larva and measurements were taken from shed head capsules pupa of Eantis minor (Comstock) from Puerto Rico on using an ocular micrometer. Voucher material includ­ cultivated Citrus and native Zanthoxylum species. ing preserved larvae of all instars together with their Eantis tamenund has been reared in Texas on native associated shelters is deposited in the collection of the Zanthoxylum species (Kendall 1965). Hayward (1941, senior author. 1948) reported E. thraso from various Zanthoxylum species and Moss (1949) found it on one Zanthoxylum species. Biezanko et al. (1974) found E. thraso on four RESULTS species of Citrus and two species of Zanthoxylum. Early stages. No eggs were encountered, First in­ Most recently, Janzen and H allwachs (2003) found star (n = 7). Head capsule black to dark brown, mod­ E. thraso on six species of Zanthoxylum in northwest­ erately heart-shaped; body parallel sided, slightly flat­ ern Costa Rica. Despite the fact that larvae of tened, roughly dome shaped in cross section, entirely E. thraso have been observed and reared several pale olive-green to clear green with some variation due times, the most detailed description of its larva avail­ to gut contents, dorsum bare but with a ventro-Iateral able is that provided by Moss (1949), who described the fringe of minute, pale setae and four long, stiff, pale setae along the margin of the anal plate. Second instar 1 Research Associate, M useo de Zoologia, Facultad de Ciencias, Universidad Nacional Aut6nOlna de Mexico . Apdo. Postal 70-399, (n = 6) as described for first instar. Third instar (n = Mexico , D. F. 04510 Mexico. 13) head capsule caramel-brown, fading gradually to 44 JOURNAL OF THE LEPTDOPTER1STS' SOCIETY

hours before eclosion, the pupa turns generally more A yellowish, with the wing pads gaining an orange-brown cast and the eyes becoming dark brown. Larval shelters. Larvae of all instars formed a shel­ ter made by modifying the leaves of their host plant. Terminology for discussion of shelters follows Greeney and Jones (in press). First instar shelters (n = 7) were roughly triangular or trapezoidal shaped sections of leaves cut from the leaf margin and flipped onto the dorsal surface of the leaf (Fig. lA). Two major cuts in the leaf were made that ended towards the central por­ tion of the leaf and angled to a narrow shelter bridge. This is termed a two-cut, stemmed fold, Group III, type 10 shelter (Greeney & Jones in press). Second in­ star larvae (n = 6) were found in shelters as described for the first instar. No abandoned first-instar shelters were found at the study sites, suggesting that larvae re­ mained in the shelter built during the first instar. Third instar shelters (n = 13) were similar to those described for the first instar, but were larger. Three of 14 fourth ins tar larvae had built a third shelter formed by silking together two leaves so that the dorsal surface of one FIG. 1. Shelters and head capsule of Eantis thraso from eastern leaf contacted the ventral surface of another leaf and Ecuador: A, Larval shelter type built by first through third (or rarely formed a pocket (Fig. IB). This is termed a two-leaf through fifth ) instar larvae; B, Larval shelter type built by many fourth and fifth instar larvae; C, Head capsule of fifth instar larva. pocket, Group I, type 2 shelter (Greeney & Jones in press). The remaining eleven fourth ins tar larvae were encountered in the second larval shelter as described dark brown basally around ommatidia and mouthparts, for the third instar. Fifth instar (n = 9) larvae remained heart shape more pronounced, average width 1.41 mm in shelters built during the fourth (or third) instar. Two (n = 2); body as described for first instar but some indi­ pupae were found in the field, and both were inside the viduals with a faint supraspiracular line of yellow hatch last feeding shelter built. The pupae were attached, marks from T2 to A8. Fourth instar (n = 14) average face down, to the ventral surface of the shelter by head capsule width 2.62 mm (n = 3), body as described heavy silking at the cremaster, and were supported by a for third instar, but yellow hatch marks more pro­ band of silk across the mid-thorax. nounced, and anal plate still with four stiff, pale setae, which are approximately the same size as in the third DISCUSSION instar, so now appear proportionately smaller. Fifth in­ Despite the relative abundance of larvae at the study star (n = 9) head capsule (Fig. lC) now with distinct site, adult E. thraso were not common. Those encoun­ black area basally giving a bearded appearance, aver­ tered were exclusively associated with disturbed areas. age width 4.37 mm (n = 4); body as described for Other species of Rutaceae, such as Zanthoxylum arner­ fourth instar (also see photos by Janzen & Hallwachs icanum Mill and Ptelea trifoliata L. , have been sug­ 2003). Immediately upon molting, the fresh head cap­ gested as larval host plants for Eantis from other re­ sules of last instar larvae are pale lime green and slowly gions (Kendall 1965, Kendall & McGuire 1975). It is change to an ivory color and finally to caramel over the unknown if these genera are utilized in our area, but course of an hOllr. Pupa (n = 8) stout with a short blunt the occurrence of adults associated with disturbed ar­ horn arising between eyes and projecting foreword and eas, where Citus is often abundant, suggests that Citus slightly upward, ground color entirely lime-green; four is an important larval host in eastern Ecuador. On the tiny black dots behind the head form a small crescent other hand, Janzen and Hallwachs (2003) reported on on the dorsum of the prothorax, fOllr additional small 201 rearings of Costa Rican E. thraso, all from Zan­ brown spots located on dorsal abdomen just anterior to thoxylum species. cremaster. Cremaster dark brown, entire pupa covered The construction of larval shelters by hesperiid lar­ with a light dusting of white waxy flocculence except vae has been known for many years (e.g., Scudder for two small bare patches in the shape of crescents 1889) and has since been reported for a wide variety of along the costal edge of wing pads. Approximately 24 species (e.g., Moss 1949, Miller 1990, Atkins et al. VOLUME 57, NUMBER 1 45

1991). Within the , species of many taxa­ Ecologfa e Pr eserva ~ao de uma Area Florestal no Sudeste do nomic groups build shelters of some type (e.g., Miller Brasil. Universidade Estadual de Campinas, Brazil. 321 pp. BRUNER, S. C. , L. C. SCARAMUZZA & A. R. OTERO. 1945. Catalogo 1983, DeVries 1987, Jones 1999), and there are at least de los insectos que atacan a las plantas econ6micas de Cuba. 10 distinct types built by the Hesperiidae alone Ministerio de Agricultura, La Habana, Cuba. (Greeney & Jones in press). All of those recorded for DEVHIE S, P. J. 1987. The of Costa Rica and their natural history. Vol. 1. Papilionidae, Pieridae, and Nymphalidae. the Hesperiidae involve modifying the host plant with Princeton University Press, Princeton, New Jersey. 327 pp. cuts and/or silk. Ontogenetic changes in the form of DEVRIES , P. J. , T. R. WALLA & H. F. GREENEY. 1999. Species diver­ these shelters are also well documented (Graham sity in spatial and temporal dimensions of fruit-feeding butterflies from two Ecuadorian rainforests. BioI. J. Linn. Soc. 68:.3.3.3-35.3. 1988, Miller 1990). Eantis thraso larvae form shelters EVA NS, W. H. 195,3. A catalogue of the American Hesperiidae indi­ of two basic types. The first, involving cutting of the cating the classification and nomenclature adopted in the leaf (Fig. lA), is a common type seen in many other British Museum (Natural History). Part III (groups E, F, G), Pyrginae, section 2. British Museum, London. 246 pp. species (Young 1991, 1993, HFG pers. obs.). The sec­ --- . 1955. A Catalogue of the Amelican Hespeliidae indicating ond, involving the silking together of two separate the classification and nomenclature adopted in the British Mu­ leaves (Fig. IE), is likely utilized in this case due to seum (Natural History). Part IV (groups H to P), Hesperiinae and Megathyminae. British Museum, London. 499 pp. size constraints imposed by the growing larvae and the GRAHAM , A. T. 1988. The life history of a semi-arid population of relatively small leaves of the Citrus host. Croitam; cmites (Hcwitson) (Lepidoptera: Hesperiidae: vVhile no oviposition events were observed and no Trapezitinae). Aust. Ent. Mag. 15: 123- 126. GR EENEY, H. F. & M. T. JONES. (in press). Shelter bUilding in the eggs were found in the field, it is likely that E. thraso Hesperiidae: a classification scheme for larval shelters. J. Res. oviposit on the meristem leaves of their host. All Lep. early instars were found on pale, fresh leaves, and HAYWARD, K. J. 1941. Plantas alimenticias de hesperidos argenti­ nos. Revta Soc. Ent. Argent. 11:31- 36. later instars were typically on or near new growth. - --. 1948. Insecta, Lepidoptera (Rhopalocera), familia Hcs­ This preference for meristem tissue is known for periidarurn, subfamilia Pyrrhopyginarum et Pyrginaru n.1n De­ other rutaceous feeders (Vaidya 1969, Young 1993, scole, H. R. (ed. ), Genera el Species Anim,i1ium Argentinorum. Guillermo Kraft, Buenos Aires . Vol. 1. .389 pp. HFG pers. obs.), as well as many species of butterflies HENNIG, W. 1966. Phylogenetic systematics. University of Illinois feeding on a variety of host plants (DeVries 1987). Press, Urbana. JA"i ZE"i, D. H. & W. HALLWACHS. 2003. Philosophy, navigation and The value of detailed natural history studies in cre­ usc of a dynamiC database ("AGC caterpillars SRNP") lor an in­ ating and testing phylogenetic hypotheses has been ventory of the macroeaterpillar fauna and its foodplants and noted by other authors (Hennig 1966, DeVries 1987). parasitoids, of the Area de Conservaci6n Gnanacaste (AGC), In general, the morphological and ecological attributes northwestern Costa Rica. http://janzen.sas.llpenn.edu lON ES, M. T. 1999. Functions of shelter bUilding in the IIesperi ­ of hesperiid larvae are very poorly known, and the tax­ . idae. Unpublished Masters Thesis, Georgetown University, onomy of most tropical groups remains con­ Washington, DC. fused; much additional work is needed in these areas. In KENDALL, R. O. 1965. Larval food plants and distribution notes for twenty-four Texas Hesperiidae. J. Lep. Soc. 19:]-33. the face of ever increasing rates of habitat destruction, KENDALL, R. O. & w. w. MCGUIRE. 1975. Larval foodplants lex we hope this study may encourage others to continue twenty-one species of skippers (Lepidoptera: Hesperiidae) publishing observations on the life history of this and from Mexico. Bull. Allyn Mus. 27:1-7. LEWI S, U. L. 1987. Butterflies of the World. Harrison House, New other poorly known (but frequently encountered) taxa. York. 312 pp. M ILLEH, C. G. 1990. The life history of Chaetocneme denitza (He­ ACKl\OWLEDGMENTS wilson) (Lepidoptera: Hesperiidae: Pyrginae). ALlSt. Ent. Mag. 17:97-100. This research was fimded in part by a grant from the Azar Foun­ MILLER, W. E. 1983. Eucosmomorpha albersana (Hiihner), a dation. HFG thanks the owner and staff of Sacha Lodge for their Palaearctic species, collected in North America (Tortricidae, generosity and support. Phil DeVries, George Austin, and Meg Crapholitini). J. Lep. Soc. 37:88-89. Jones provided useful comments on the manuscript and have gener­ Moss, A. M. 1949. Biological notes on some Hesperiidae of Para ously provided information on the biology of various skippers. Fi­ and the Amazon (Lep. Rhop. ). Acta Zoo!. Lilloana 7:27-29. nally, Ryan Hill and Nicolc Gerardo provided a great deal of assis­ PANTON, E. S. 1898. The life history of some Jamaica Hesperiidae. tance in the field. Thanks also to Roy Kendall, Richard Peigler, and J. Inst. Jamaica 2:435-441. Andrew Atkins lor providing some literature cited herein. Addition­ POEY, F 1832. Centurie de lepidopteres de rIle de Cuba, con­ ally, we acknowledge the PBNHS for their ongoing support of our tenant la description et les figures coloriees de cent especes de natural history studies. papillons nouveaux ou peu conn us, representes d'apres nature, LITERATURE CITED sou vent avec la chenille, la chrysalide, et plusieurs detalis mi­ croscopiques. J. Albert Mercklein, Paris. ATKINS, A., R. MAYO & M. MOORE. 1991. The life history of Signef,a SCUDDER, S. H. 1889. The Butterflies of the Eastern United States tymbophora (Meyrick and Lower) (Lepidoptera: Hesperiidae: and Canada, with special reference to New England. Vol. 2, Ly­ Trapezitinae) Aust. Ent. Mag. 18:87- 90. caenidae, Papilionidae, Hesperiidae. S. H. Scudder, Cam­ BIEZA NKO, C. M., A. RUFFINELLI & D. LI"iD. 1974. Plantas y otras bridge, M assacllllsetts. xii + 767-1774 pp. sustancias alimenticias de las orugas de los lepid6pteros SM YTH, H. E. G. 1919. Dominio de inseetos en los dtricos en Urllguayos. Revista do Centro de Cicncias Rurais 4:107-148. Puerto Rico. Revta. Agric. Puerto Rico 3:39-50, .55-62. BROWN , K. S. 1992. Borboletas da Serra do Japi: diversidade, habi­ VAIDYA, V. G. 1969. Investigations on the role of visual stimuli in tats, recursos alimentares e val;a~ao temporal, pp. 142-187. In the egg-laying and resting behavior of Papilio denwleus L. (Pa­ Morellato, L. P. C. (org.), Hist6ria Natural da Serra do Japi, pilionidae, Lepidoptera). Anim. Behav. 17:350-3.55. 46 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY

WARREN, A. D. 1996. Phylogeneti(; revision of the skippers of the YOUNG, A. M. 1991. Notes on the natural history of Quadrus mithridates species group, and their replacement in Eantis Bois­ (Pythonides) contuberalis (Hesperiidae) in Costa Rica. J. Lep. duval, 1836 (Lepidoptera: Hesperiidae: Pyrginae). Entomology Soc. 45:366--37l. Department Honors Thesis, Cornell University, Ithaca, New York. ---. 1993. Notes on the natural history of Achlyodes selva WaLCOTI', C. N. 1923. "Insecatc Portoricensis." A preliminary an­ (Hesperiidae) in Costa Rica. J. Lep. Soc. 47:323-327. notated check-list of the of Puerto Rico, with descrip­ tions of some new species. J. Dept Agnc. Puerto Rico 7:1-313. - --. 1951. The insects of Puerto Rico. J. Dept Agric. Puerto Received for publication 16 January 2002; revised and accepted Rico 32:417-748. 22 August 2002.