10 January 2000 PROC. ENTOMOL. SOC. WASH. 1021, 2000, pp. 162-169

THE NEW WORLD HAIRSTREAK KAYE : : :

ROBERT K. ROBBINS

Department of Entomology, National Museum of Natural History, Smithsonian Insti tution, Washington, DC 20560-0127 U.S.A. e-mail [email protected]

Abstract-Even though Arawacus Kaye is widely used in works on North American , its characterization has been inconsistent and its included have not been listed. Subterminally constricted tips of the papillae anales are proposed to be the best way to characterize Arawacus. They are consistent with the higher classification of the Section of the Eumaeini, to which Arawacus belongs. They are consistent with patterns of oviposition specificity. This characterization re-confirms that Polyniphes Kaye and Dolymorpha Holland are junior synonyms of Arawacus. Tigrinota Johnson is synonymized with Arawacus, new synonym. All nomenclaturally available specific names that belong to Arawacus are listed. Key Words: Thereus, , Contrafacia, Dolymorpha, Polyniphes, Tigrinota, Solana ceae

The generic name Arawacus Kaye was two terminal cornuti, "one of which has an rarely used until 1981. The original descrip abrupt, larger disc-like terminal expansion tion was based on superficial wing pattern with peripheral teeth" Fig. 1. He later elements of the South American type spe made Dotytnorpha a junior synonym of Ar cies, A. aetolus Sulzer Kaye 1904. As a awacus using the same two character states result, besides being cited in nomenclatural an unpublished manuscript that is being lists e.g., Comstock and Huntington 1959- deposited in the Archives of the Carnegie 1964, Eliot 1973, Arawacus was used only Museum of Natural History, Pittsburgh, for species with wing patterns similar to the Pennsylvania, U.S.A.. Miller and Brown type Brown and Mielke 1967; Robbins 1981, who received a copy of the manu 1980, 1981. But after Miller and Brown script after Clench's death, published this 1981 made Arawacus a senior synonym synonymy in their catalog without expla of Dolytnorpha Holland, whose type spe nation. cies ranges into the United States, the use Arawacus has been characterized in dif of Arawacus in general works ferent ways since 1991. Because Clench mushroomed e.g., Scott 1986, Bailowitz mistakenly lumped five distinct species in and Brock 1991, Stanford and Opler 1993, the A. aetolus complex Robbins, Hudson, Cassie et a!. 1995. and Clench, in preparation, he did not re Arawacus was characterized by mor alize that the type species of Arawacus phology of its male genitalia. Clench 1961: lacks both male genitalia traits that he had 212 had delimited Dolymorpha primarily proposed Fig. 2. Consequently, when by a "ventro-lateral conical, acuminate pro Robbins 1991: 3 briefly treated Arawacus cess at middle" of the valves and by the as an outgroup of Rekoa Kaye, he provi VOLUME 102, NUMBER 163

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Figs. 1-2. Male genitalia in ventral aspect with penis in lateral aspect, posterior end towards top of the page. 1, , with arrows pointing to the posterior pointing processes on the valves and to the flattened cornutus with terminal teeth. 2, A. aetolus, which lacks the posterior pointing processes and has an arrowhead shaped cornutus. sionally re-characterized it by "I papillae not note either trait or mention that Tigri anales constricted subterminally, and 2 nota might be closely related to Arawacus. larvae that feed on the foliage of Solanum Further, he treated Dolymorpha as a distinct Solanaceae," but did not illustrate the first genus without mentioning the previous syn character or list the included species. He onymy with Arawacus. Note: Some copies confirmed the synonymy with Dolymorpha, of Johnson [1993] have Johnson and added Po/yniphes Kaye as a second junior Kroenlein listed as the authors. I follow the synonym of Arawacus, and placed Arawa citation in Lamas et al. [1995] with Johnson cus in the Thereus Section of the Eumaeini. as the sole author. Johnson 1992, 1993 described and re The purpose of this paper is to stabilize vised Tigrinota for species that possess usage of Arawacus and to provide a generic both traits that Robbins had proposed for for revision at the species level. characterizing Arawacus, but Johnson did Specifically, this paper shows that the con- 164 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON stricted papillae anales occur in the type membranous dorsally again illustrated in species of Arawacus, Dolymorpha, Polyni Robbins 1991. This classification is a phes, and Tigrinota, but not in other genera framework within which the monophyly of of the Thereus Section, to which Arawacus Arawacus can be assessed. belongs. It is further shown that Tigrinota Type species nomenclature.-Synonymy was delimited by characters that are incor of the type species of Arawacus, Contra rectly described or that occur in other gen facia, and Tigrinota is a bit confused. The era of the Thereus Section. Finally, the spe type of Arawacus is Papi/io linus Fabricius, cific taxa that belong to Arawacus are listed which is a junior synonym of P. aetolus for the first time. Sulzer Comstock and Huntington 1959- 1964. The type of Contrafacia is C. inex MATERIALS AND METHODS icana Johnson 1989, a species that is indis This study was based upon the approxi tinguishable from C. imnia Prittwitz 1865 mately 3,250 specimens belonging to the Robbins 1991. The type of Tigrinota is Thereus Section genera Arawacus. Rekoa, Thee/a el/ida Hewitson, which occurs from Thereus Hubner, Contrafacia Johnson, and northern Venezuela to central Argentina. Its the "" ligurina species group an Un- wing pattern varies slightly over this wide described genus whose species are placed, range with virtually no geographical varia following convention, in Thecla E, a genus tion in genitalic structures Robbins, in that does not belong to the Eumaeini, Eliot prep., contrary to the treatments in John 1973 in the National Museum of Natural son 1992, 1993. History, Smithsonian Institution, Washing ton, DC, USA USNM. Included were all RESULTS species that belong to these genera as de Monophyly and Arawacus.-The papil limited by Robbins 1991. Preparation of lae anales "ovipositor valves" were ex genitalia for examination using light and amined for 57 of the approximately 63 spe scanning electron microscopes follows the cies that belong to the Thereus Section. Fe procedures in Robbins 1991. The male males are not known for the other 6 species. and female genitalia of all species in the The tips of the papillae anales ventral as Thereus Section were dissected except for pect for the type species of Arawacus Fig. those few species known from only one 3, Do/ymorpha Fig. 4, Tigrinota Fig. 5, sex, including more than 120 genitalic dis and Polyniphes Fig. 6 have a subterminal sections of those species placed in Arawa constriction, which might alternately be de cus in this paper. scribed as terminally expanded. The tips of the papillae anales of Rekoa Figs. 7-9 are PREVIOUS RESULTS notched, not subterminally constricted. The The Thereus section.-Robbins 1991 papillae anales of Thereus Figs. 10-11 are proposed two phylogenetic hypotheses in similar to those of Rekoa, but contain a volving Arawacus. First, Arawacus, Rekoa, sclerotized patch that is otherwise unre The reus, Cantrafacia, and the "Thecla" ported in the Eumaeini Robbins 1991. ligurina group form the The reus Section of The tips of the papillae anales of Contra the Eumaeini because they share a unique facia Fig. 12 and the "Thecla" ligurina process of the vinculum abutting the brush group Fig. 13 are regularly tapered. Al organs, when present illustrated in Robbins though the papillae anales in the Thereus 1991. Second, the first three genera form Section vary in shape, only those of Ara a monophyletic group. The ductus semin wacus have a conspicuous subterminal con alis arises from a pouch of the posterior striction. corpus bursae. This pouch is dorsal of the Larval food plant specificity.-Among ductus bursae, sclerotized laterally, and the eumaeine genera, larval feeding on the VOLUME 102. NUMBER 1 65

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Figs. 3-13. Papillae anales in ventral aspect, posterior end towards top of the page. 3, Arawacus aetolus; arrow points to the subterminal constriction. 4. A. jada. 5. A. ellida. 6, A. du,nenilii. 7, . 8, R. palegon. 9, It marius. 10, Thereus lausux; arrow points to scierotized patch. Ii, T. praxis; arrow points to scierotized patch. 12. Contrafacia im,na. 13, "Thecla" lyde. leaves of Solanum as opposed to flowers Leguminosae Robbins, in press. Leaf occurs only in Arawacus, but the larvae of feeding on So/anuin has been recorded for three Arawacus species have recently been 11 of the 17 Arawacus species noted in the reported to eat plants in the Compositae and list at the end of this paper, including the 166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON type species of Arawacus, Dolyrnorpha, Ti grinota, and Polyniphes Guppy 1904, 1914; Bourquin 1945; Kendall 1975; Rob- bins, in press. However, larvae of A. el/ida Hewitson, which have been reared re peatedly from the leaves of So/an urn, and A. binangula Schaus were both recently recorded eating flowers of Compositae-a larval food plant used in the related genus Rekoa-and the larvae of A. tarania were recorded on legumes Brown 1993. Con sequently, although larval feeding on the leaves of Solanurn is widespread in Ara wacus and is unreported in other Eumaeini, it cannot be used to delimit the genus. The larvae of other The reus Section spe cies primarily use plants in families other than the Solanaceae as larval food Robbins 1991, in press. Rekoa larvae are extremely polyphagous on buds, flowers, and shoots of plants in 16 families, including Solana ceae, but most records are in the Compos itae, Leguminosae, and Malvaceae. In con trast, larvae of Thereus specialize on Lor Fig. 14. Scanning electron micrograph of the anthaceae, with one species eating Mal valves of Arawacus ellida in ventral aspect, posterior end towards the top of the page. Note the lack of rims, pighiaceae and Chrysobalanaceae. microtrichia, and "caudo-lateral sculptures." Contrafacia larvae are recorded eating plants in the Compositae and Leguminosae, and the only food plant record in the "The 1991, these bands would appear to be a e/a" ligurina group is in the Erythroxyla symplesiomorphy and are questionably di ceae. agnostic. Monophyly and Tigrinota.-The only al The diagnosis of the male genitalia of Ti ternative to the proposed classification in grinota in Johnson 1992: 186 is cited in this paper is that of Johnson 1992, 1993. its entirety because it is confusing. "Ven It is the purpose of this section to assess the trally, valvae appearing as paired and rather evidence supporting the monophyly of Ti smoothly scierotized oblongate lobes sepa grinota. rated by a thin transparent fissure which at The diagnosis of Tigrinota Johnson each end, shows sclerotized ridges forming 1992, 1993 is similar in both works, but 1 prominent rims about the bibbed area difficult to interpret. The diagnosis of the and 2 caudo-lateral sculptures of the val wings refers to the hindwing "with concen val terminus from which emerge clusters of tric bands," but bands on the wings of these robust microtrichia." The valves of the type species are not concentric. It refers to the species of Tigrinota are illustrated in ven "ovate brand," but the type species lacks tral aspect Fig. 14. There are no rims. an ovate brand or other scent patch. It men There are no microtrichia, robust or not. tions alternating bands on the hindwing, but Perhaps setae were mistaken for microtri these also occur in Cramer chia, but if so, their occurrence on the ven and R. pnalina Hewitson. Since Rekoa is tral valves is true for virtually all eumaeines a close relative of Arawacus Robbins e.g., Clench [1961] characterized Ca/b- VOLUME 102, NUMBER 167 phrys Westwood by the unusual lack of Se phyly of Tigrinota. Either its diagnostic tae on the valve tips. It is unclear to what traits are incorrect, such as the concentric the "caudo-lateral sculptures" refer Fig. hindwing bands and rims on the valves, or 14. Johnson 1993: 2 repeats these char appear to be symplesiomorphic, such as the acters and adds "brush organ occurrences alternating hindwing bands and "transpar diffential sic", but it is unclear what this ent neck" of the ductus bursae. means or how it might be diagnostic. As Provisional classification of Arawacus.- best I can tell, those parts of this diagnosis The status of Po/yniphes as a synonym that are accurate do not differentiate Tigri Robbins 1991 is confirmed. The status of now from most eumaeines. Dolyinorpha as a synonym Miller and The diagnosis of the female genitalia is Brown 1981, which was not recognized in similarly confusing Johnson 1992, 1993. Johnson 1992, is also confirmed. Tigri The description I presume of the ductus nota is synonymized with Arawacus, new bursae with a "transparent neck" refers to synonym. the ductus bursae of most Rekoa and Ara Johnson 1992, 1993 presented a clas wacus Robbins 1991; Robbins, Hudson, sification of almost half the specific names and Clench, in preparation and is not di listed below. He did not assess intraspecific agnostic. The description of a simple "sub geographical variation and reported "diag cordate incised posterior cavity abbreviat nostic" morphological traits, such as con ed sipc in the diagnosis of the 8th tergite" centric hindwing bands and rims on the probably referring to the shape of the 8th valves, that do not exist. Further, my ex abdominal tergum is difficult to interpret. amination of more than 120 genitalic dis However, the shape of this tergum is similar sections and detailed comparison of andro in Tirgrinota and some Rekoa, such as R. conia and wing patterns from throughout zebina Hewitson, so it is unlikely to be the range of each species failed to confirm diagnostic. most of his conclusions. Rather than await detailed species level revisions, I present an DISCUSSION alternate classification below that is intend Taxonomy.-The genus Arawacus ap ed to be a working hypothesis for species pears to be monophyletic, characterized by level revisions. Although it is unusual to the shape of the papillae anales. Although present a tentative classification, such as it is not desirable to delimit Arawacus by this one, the extensive reporting of charac only one character, this classification is con ters that do not exist Johnson 1992, 1993 sistent with the hierarchical classification of is perhaps a more unusual circumstance that the Thereus Section, and other morpholog creates the need for a more reasonable ical traits support the monophyly of The r working classification. ens and Rekoa Robbins 1991. These char As characterized in this paper, Arawacus acterizations of Arawacus, The reus, and Re contains 39 available specific names repre koa are reasonably consistent with patterns senting 17 biological species. There is also of larval plant use. For these reasons, this one undescribed species Robbins, Hudson, classification appears to be the best option and Clench, in preparation. The female for promoting a stable generic nomencla genitalia of all have been examined except ture in the Thereus Section. for A. euptychia, for which no females are Recognition of Tigrinota is unlikely to be known. However, A. euptychia is placed in a reasonable alternative. Its use leaves Ar Arawacus because its male genitalia are awacus and Polyniphes uncharacterized and barely distinguishable from those of A. du would consequently destabilize the generic menilii and A. tadita. taxonomy of the Thereus Section. There is Those species that have been reared from little, if any, evidence to support the mono- plants in the Solanaceae, Compositae, or 168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Leguminosae are designated respectively in Arawacus. Some information reported in bold with 8, C, or L. this paper was discovered during a joint 1. Arawacus togarna Hewitson, 1867 project with P. Hudson on species level sys 2. Arawacus lincoides Draudt, 1917-S tematics of the A. aeto/us complex. For help 3. Arawacus aetolus Sulzer, 1776-S with the illustrations, I thank S. Braden, W. Papilio linus Fabricius, 1776 Brown, V. Malikul, and 0. Venable. For Papilio amelia Herbst, 1804 reading and commenting on the manuscript, 4. Arawacus separata Lathy, 1926-S I am grateful to J. Glassberg. G. Lamas, J. Thecla paraguayensis Lathy,1926 Scott, and A. Warren. 5. Arawacus aethesa Hewitson, 1867 LITERATURE CITED 6. Arawacus s/to Boisduval, 1836-S Thecla phaenna Godman and Salvin, Bailowitz, R. A. and J. P Brock. 1991. Butterflies of 1887 Southeastern Arizona. Sonoran Stud Arawacus mexicana D'Abrera, 1995 ies. Thcson, 342 pp. Brown, K. S. Jr. and 0. Fl. H. Mielke. 1967. Lepidop Arawacus 7. leucogyna Felder and Felder, rera of the central Brazil plateau. I. Preliminary 1 865-S list of Rhopalocera continued: Lyeaenidae. Pier Thecla phaea Godman and Salvin, idae, Papilionidac, Hesperiidae. Journal of the 1887 Lepidopterists' Society 2 I: 145-168. 8. Arawacus meliboeus Fabricius, 1793-S Brown, Jr., K. 5. 1993. Theclinae endemic to the cer rado vegetation central Brazil, 152. In New, Jolaus eurisides p. HUbner, 1823 T. R, ed,, Conservation Biology of Lycaenidae Thecla barrensis Rosa, 1936 Butterflies, IUCN. Gland, Switzerland. 9. Arawacus jada Hewitson, 1867-S Bourquin, F 1945. Mariposas Argentinas. Edicion del 10. Arawacus el/ida Hewitson, I 867-S,C autor, Buenos Aires, 211 pp., 192 photographs, 3 Thecla toba Hayward, 1949 maps. Cassie, B, J. Glassberg, P. Opler, R. Robbins, and G, Tigrinota perinota Johnson, 1992 Tudor. 1995. North American Butterfly Associa Tigrinota jennfera Johnson, 1992 tion NABA Checklist & English Names of Tigrinota catamarciana Johnson, 1993 North American Butterflies, First Edition. North Tigrinota chaosa Johnson, 1993 American Butterify Association, Morristown, 11. Arawacus hypocrita Schaus, 1913 New Jersey, 43 pp. Clench, H. K. 1961. . 177-220, In Arawacus pp. 12. dolylas Cramer, 1777-S Ehrlieh, P. R., and A. H. Ehrlich, How to Know Pseudolycaena spurius Felder and the Butterflies, Brown Company, Duhuque, Iowa, Felder, 1865 262 pp., 525 figs. Thecla dolosa Staudinger, 1888 Comstock, W. P. and E. I, Huntington. 1959-1964. An Thecla pallida Lathy, 1930, annotated list of the Lycaenidae Lepidoptera: Rhopalocera of the Western Hemisphere, Journal 13. Arawacus duineni/il Godart, 1824-S of the New York Entomological Society 66: 103- Thecla argiva Hewitson, 1877 118, 67: 59-95, 163-212, 68: 105-122, 176-186, Thecla obscura Staudinger, 1888 232-240, 69: 54-58, 105-118, 157-176, 191- Thec/a carteri Weeks, 1906 200, 70: 39-46, 100-118, 177-179, 71: 45-57, 14. Arawacus euptychia Draudt, 1921 115-119, 189-197, 262-264, 72: 120-130, 173- 192. 15. Arawacus tadita Hewitson, 1877-S Eliot, J. N. 1973. The higher classification of the Ly Thecla datitia Jones, 1912 caenidae Lepidoptera: A tentative anangement. 16. Arawacus binangula Schaus, 1902-C Bulletin of the British Museum Natural History Thecla bolima Schaus, 1902 Entomology 28No.6: 371-505. 17. Arawacus tarania Hewitson, 1868-L Gnppy, J. 1904. Notes on the habits and early stages of some Trinidad butterflies, 225-228, 2 Thecla atrana Schaus, pp. 1902. plates. Appendix to: W. J. Kaye. A catalogue of ACKNOWLEDGMENTS the Lepidoptera Rhopalocera of Trinidad. Trans actions of the Entomological Society of London, Mary Clench kindly sent me her late hus pp. 159-228, 2 pl. band's unpublished manuscript on part of 1914. Notes on the habits and eady stages of VOLUME 102, NUMBER I 169

some Trinidad butterflies. pp. 76-79. Appendix to: Miller, L. D, and E M. Brown. 1981. A catalogue/ W. J. Kaye. Fauna of Trinidad. Part I. A catalogue checklist of the butterflies of America north of of the Lepidoptera Rhopalocera of Trinidad. But Mexico. Memoirs of the Lepidopterists' Society, terflies of Trinidad. The Agricultural Society of No. 2. 280 pp. Trinidad and Tobago. Paper No. 558. 79 pp. Robbins, R. K. 1980. The lycaenid "false head" hy Johnson, K. 1992. Genera and species of the Neotrop pothesis: historical review and quantitative anal ical `elfin-like hairstreak butterflies Lepidop ysis. Journal of the Lepidopterists' Society 34: tera, Lycaenidae. Theclinae. Reports of the Mu- 194-208. scum of Natural History, University of Wisconsin 1981. The `false head" hypothesis: predation Stevens Point 22: 1-279. and wing pattern variation of lycaenid butterflies. 1993. Hairstreak butterflies of the genus Ti The American Naturalist 118: 770-775. gnno,rI Lepidoptcra, Lycacnidae, Theclinae. Re 1991. Evolution, comparative morphology, ports of the Museum of Natural History, Univer and identification of the eumaeine butterfly genus sity of Wisconsin Stevens Point 37: 1-2t. Rekoa Kaye Lycaenidae: Theclinae. Smithsoni Kaye, W, J. 1904. A catalogue of the Lepidoptera Rho an Contributions to Zoology No. 498, 64 pp. palocera of Trinidad. Transactions of the Ento In press. Larval Food Plants of the Neotrop mological Society of London, pp. 159-228, 2 pl. ical Eumaeine Hairstreaks Lycaenidae: Thecli Kendall, R. 0. 1975. Larval foodplants for seven spe nac. Memoir No. 5, Lepidopterists' Society. 246 cies of hairstreaks Lycaenidae from Mexico. pp. Bulletin of the Allyo Museum of Entomology 24: Scott, J. A. 1986. The Butterflies of North America, A 1-4. Natural History and Field Guide. Stanford Uni Lamas, G., R. G. Robbins, and W. D. Field. 1995. versity Press, Stanford, California. 583 pp.' 64 p1. Bibliography of butterflies. An annotated bibli Stanford, R. E. and PA. Opler 1993. Atlas of Western ography of the neotrupical butterflies and skippers USA Butterflies, Including Adjacent Parts of Can Lepidoptera: Papilionoidea and Hesperioidea. ada and Mexico. Published by the authors, Denver Atlas of Neotropical Lepidoptera. 124: 463 pp. and Fort Collins, Colorado, 275 pp.