Journal of the Lepidopterists' Society 40(3), 1986, 138-157 EVOLUTION AND IDENTIFICATION OF THE NEW WORLD HAIRSTREAK BUTTERFLIES (LYCAENIDAE: EUMAEINI): ELIOT'S TRICHONIS SECTION AND TRICHONIS HEWITSON ROBERT K. ROBBINS Department of Entomology, MRC NHB 127, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560 ABSTRACT. I revise the lycaenid genus Trichonis Hewitson (Theclinae: Eumaeini), establish the new combination Trichonis hyacinthus Cramer, and make Papilio thenaus Cramer 1777 a junior synonym of P. hyacinthus Cramer 1775. The genus consists of two species, T. hyacinthus and T. immaculata, which differ in wing pattern, forewing shape, color of androconial scales, and length of the third palpal segment. I then assess Eliot's higher classification of the Eumaeini. I examine leg, genitalia, and wing mor- phology, and conclude that Eliot's Trichonis Section is diphyletic. The unusual male foretarsus of Trichonis appears to have evolved independently three times in the Eu- Eliot (1973) and Clench (1964, pers. comm. 1978) proposed different higher classifications for the "New World hairstreaks" (Lycaenidae: Theclinae: Eumaeini). Eliot divided them into a Trichonis Section• for genera Trichonis Hewitson and Micandra Schatz•and an enor- mous Eumaeus Section•for the remaining genera (64 available names). Clench also divided these butterflies in two groups; in one he isolated Eumaeus Hiibner (his Eumaeini), and in the other he lumped the remaining genera (his Strymonini). The purpose of this paper is to assess the evidence for Eliot's pro- visional higher classification of the Eumaeini. Specifically, I consider whether the Trichonis Section is a monophyletic group. As basic in- formation needed to answer this question, I revise Trichonis. Mican- dra, which Clench (1971) treated preliminarily, needed little work for the purpose of this paper. I then discuss the evidence for Eliot's clas- sification. GENUS TRICHONIS Trichonis consists of two species known only from males, T. theanus Cramer (Fig. la, b) and T. immaculata Lathy (Fig. 2a, b). In 1865, Hewitson named Trichonis, primarily on the basis of an unusual male foretarsus in T. theanus, which he described as "exarticulate, robust, and broad beyond the middle" (Fig. 7a, b). Hewitson also illustrated the "female" of T. thenaus, and described its foretarsus as "of the usual form, jointed and spined." Lathy (1930) pointed out that Hew- itson's female was the male of a second species, which he named T. immaculata. Lathy did not, however, compare foreleg morphology of VOLUME 40, NUMBER 3 139 FIG. 1. Adult Trichonis hyacinthus. (a) male upperside, (b) male underside, (c) fe- male upperside, (d) female underside. T. theanus and T. immaculata, nor did he discuss whether Hewitson's characterization of Trichonis was valid. No females have been asso- ciated with either species. I propose that Papilio hyacinthus Cramer (Fig. lc, d)•Cramer named all butterflies in Papilio•and a second phenotypically similar species (Fig. 2c, d) are the females of T. theanus and T. immaculata. Neither "female species" has been associated with males, but both share with T. theanus and T. immaculata a pastel blue or blue-green color on the frons and ventral wings that is unique among the Eumae- ini. Both sexes have truncate forewings, and share similar geographical distributions. Further, genital morphology, which is discussed more fully later, indicates that both males and females belong to a similar section of the Eumaeini. I associate female T. hyacinthus with male T. theanus and the new female with male T. immaculata. The bases for this action are distri- bution (Fig. 3) and forewing shape (Fig. 4). Briefly, T. hyacinthus and T. theanus are known only from the Guianas and Lower Amazon while the new female and T. immaculata occur there and in the Upper Amazon; the new female and a male of T. immaculata were collected 140 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY * -_•--' FIG. 2. Adult Trichonis immaculata. (a) male upperside, (b) male underside, (c) female upperside, (d) female underside. at the same locality in northern Peru. As detailed below, the forewing apex is more truncate in male T. immaculata than in male T. theanus and in the new female than in female T. hyacinthus. Characters unique to Trichonis include a pastel blue or blue-green frons and ventral ground color, and the genitalia, whose structure (Figs. 5 & 6) differs quantitatively from other eumaeines. I do not know which species or species group is its closest relative. Trichonis may be enlarged as our understanding of eumaeine phylogeny increases. Key to Trichonis Species and Sexes Ventral hindwings without transverse brown lines (Fig. lb, 2b) (males) ... 2 Ventral hindwings with transverse brown lines (Fig. Id, 2d) (females) ... 3 Inner edge of dorsal forewing marginal band "smooth" (Fig. la), not scalloped. Ventral forewing androconial patch almost touching upper part of discal cell (Fig. 4a) male hyacinthus Inner edge of dorsal forewing marginal band scalloped (Fig. 2a). Ventral forewing androconial patch barely enters discal cell (Fig. 4c) male immaculata Ventral hindwing with a white band between major brown transverse lines (Fig. Id); hindwing with a tail female hyacinthus Ventral hindwing with a blue or blue-green (ground color) band between major brown transverse lines (Fig. 2d); hindwing without a tail female immaculata VOLUME 40, NUMBER 3 141 TRICHONIS SYSTEMATICS Nomenclature Trichonis Hewitson (1865): Hewitson described Trichonis in the Lycaenidae with Cramer's Papilio theanus as the only species in the genus. It is the type species by monotypy (Hemming 1967). Lathy (1930) subsequently added T. immaculata. Papilio hyacinthus Cramer (1775): Cramer described Papilio hy- acinthus from the West Indies. He included a brief description and a figure of the underside, but did not mention its sex. Various authors (Fabricius 1782, Butler 1870, Draudt 1919-1920) discussed it, but their text paraphrased the original description, and illustrations were copies of Cramer's (sometimes poorly done, such as Seitz [Draudt 1919-1920]). No one has mentioned P. hyacinthus for more than half a century except to note that it is unknown from the West Indies (Comstock & Huntington 1943). There is no other species with which the figure of P. hyacinthus can be confused; the white band sandwiched between two transverse brown lines on the ventral hindwing is distinctive. All known specimens are females. There are no potential lectotypes in the Artis Collection, Instituut voor Taxonomische Zoologie, Zoologisch Museum, Universiteit van Amsterdam (Hogenes, pers. comm.), at the Rijksmuseum van Natuur- lijke Historie in Leiden (de Jong, pers. comm.), or in the British Mu- seum (Natural History) (BMNH). However, identification of Cramer's P. hyacinthus poses no problems, and a type is not needed. Trichonis hyacinthus is a New Combination. Papilio theanus Cramer (1777): Cramer described Papilio theanus from Surinam with a brief description and a figure of the ventral surface, on which "androconial" patches are evident. Hewitson (1862- 1878) illustrated the male and female, but Lathy (1930) noted that Hewitson's female is the male of a second species (which he named Trichonis immaculata). All known specimens are males. Identification of P. theanus poses a problem. Males of both Trich- onis species have extremely similar ventral wing patterns (Fig. lb, 2b), and Cramer illustrated only the ventral surface. The species can be distinguished, however, by forewing shape and extent of the "polished spot" surrounding the forewing androconia. Unfortunately, the fore- wing shape in Cramer's illustration is inaccurate (the curvature at the forewing apex is too great for either species). The polished spot in the original illustration (in the BMNH library) extends almost to the radial vein, which is not the case in the few specimens of T. immaculata that I examined. However, considering the inaccuracy of other characters 142 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY FIG. 3. Distribution of Trichonis. Solid dots designate exact localities, hollow dots represent generalized localities such as "Surinam" or "Maranham." (a) T. hyacinthus, (b) T. immaculata. VOLUME 40, NUMBER 3 143 FIG. 4. Trichonis wing venation, (a) male T. hyacinthus, (b) female T. hyacinthus, (c) male T. immaculata, (d) female T. immaculata. 144 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY in the original figure, I hesitate to base a specific identification on so minute a detail. A type specimen is desirable. I designate as lectotype of P. thenaus a specimen in the Artis Col- lection (Zoologisch Museum, Amsterdam). This specimen fits Cramer's original figure well, and was probably seen by Cramer even if it was not the model for his figure. There are no potential lectotypes in Leiden or London. The specimen bears two labels: one with the number "16" and one with the text "Trichonis Theanus Cram," written in the hand of Snellen, according to Willem Hogenes, Keeper of Lepidoptera at the Zoologisch Museum. I have added a black-bordered label which reads: "Lectotype Papilio theanus Cramer, 1777; by R. Robbins." The words "Lectotype" and "by" are printed in red and the remainder of the label is hand-written in black ink. This lectotype designation will pre- vent confusion in the future, and maintain the previous identifications of P. theanus used by Hewitson (1862-1878), Staudinger (1884-1888), Schatz and Rober (1885-1892), and Lathy (1930). I already outlined the evidence for considering P. theanus to be the male of P. hyacinthus, and now designate P. theanus Cramer a junior synonym of P. hyacinthus Cramer; New Synonymy. Trichonis immaculata Lathy (1930): Lathy described T. immacu- lata from two males: one without locality data (presumably in the Museum National D'Histoire Naturelle) and one which was the model for Hewitson's (1862-1878) "female" illustration of T. theanus (BMNH). Either can eventually be designated a lectotype, but identi- fication poses no problems. Male T. immaculata can be distinguished from male T. hyacinthus (=T. theanus) by the "smooth" dorsal fore- wing border, as noted in the above key.