Journal of the Le-pidopterists' Society 55(3), 2001, 85-100 AN OVERVIEW OF STRYMON HUBNER (LYCAENIDAE: THECLINAE: EUMAEINI) ROBERT K. ROBBINS Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560-0127, USA AND STANLEYS. NICOLAY 1500 Wakefield Drive, Virginia Beach, Virginia 23455, USA ABSTRACT. North American Strymon Hiibner was revised about 40 years ago, and the significantly larger Neotropical Strymon is now incorporated into this classification. Strymon is characterized by anteriorly directed teeth on the posterior dorsal surface of the valvae. These teeth were first noted by Clench and appear to be modifications of the sockets of setae that normally occur on eumaeine valvae. This charac- terization is most consistent with past usage and appears to represent the best evidence for monophyly. As characterized, Strymon contains 48 described species. Variation in morphology of the male genitalia, female genitalia, wings, and head is documented, and male behaviors and lar- val foodplant records are summarized. We tentatively divide Strymon into species groups, one of which is unusual in its use of Bromeliaceae as its sole larval foodplant. One recently described genus, Heoda Johnson, L. Miller & Herrera 1992, and one recently resurrected genus, Eiseliana Toledo 1978, are made junior synonyms of Strymon Hiibner 1818. Six names recently described in Strymon are transferred to other genera: Strymon angulus Le Crom & Johnson, 1997 to Thereus Hiibner; Strymon daplissus Johnson & Salazar, 1993 to Ministrymon Clench 1961; Strymon carmencitae Le Crom & Johnson, 1997 and Strymon cryptogramus Johnson, Eisele & MacPherson, 1992 to Nicolaea Johnson; Strymon nivnix Johnson, Eisele & MacPherson, 1990 to Calycopis Scudder; and Strymon additionalis Le Crom & Johnson, 1997 to Thecla F. The hindwings of Strymon nivnix are designated a leetotype, and Strymon anthracaetus Salazar, Velez, & Johnson, 1997 is regarded as a nomen dubium. Additional key words: Bromeliaceae, foodplants, territoriality, Heoda, Eiseliana. Strymon Hiibner is possibly the best-known New World hairstreak genus (Lycaenidae: Theclinae: Eu- maeini). R occurs from Canada to the temperate parts of Chile and Argentina. Some Strymon are common and well-known, such as S. melinus (Gray Hairstreak), and some are pests on commercial pineapple, such as S. ziha (Hewitson), S. megarus (Godart), and relatives (Harris 1927, Carter 1934, Fonseca 1934, Zikan 1956, Guagliumi 1965, 1967, D'Araujo e Silva et al. 1967-1968, Beutelspacher 1972, Otero & Marigo 1990). The name Strymon has been widely used in North America; the first extensive list of North American Strymon species (Barnes & McDunnough 1917) contained about 40 taxa and was followed by similar listings (Barnes & Benjamin 1926, McDun- nough 1938, Klots 1951, Dos Passos 1964). Although Ziegler (1960) and Clench (1961) rather drastically changed the characterization of Strymon, 14 of the 48 described Strymon species currently recognized (Appendix 1) are recorded from North America (Opler & Malikul 1992, Opler & Wright 1999). "Modern" taxonomic usage of Strymon began when the genus was distinguished primarily by genitalic structures (Ziegler 1960, Clench 1961). With in- creased knowledge of the Neotropical eumaeine fauna, however, it became clear that these characters, as originally proposed, do not delimit Strymon. For example, S. yojoa (Reakirt) has small anteriorly di- rected teeth on the dorsal valva tips?a structure mentioned in Clench's generic diagnosis?but lacks a tightly convoluted spiral of the ductus bursae (Fig. 20)?a structure noted in Ziegler's generic character- ization. Alternately, S. serapio (G. & S.) has the tightly-convoluted spiral (Fig. 25), but also has a dou- ble cornutus, not the single acuminate one described by Clench (1961) (Fig. 16). To complicate matters, the subsequently described genera Eiseliana Toledo and Heoda Johnson, Miller, & Herrera possess some genitalic structures that Ziegler and Clench used to characterize Strymon. Finally, six species described in Strymon since 1990 possess none of these characters. The purposes of this paper are to characterize Stry- mon, so that it will be clear which species belong to Strymon, and to provide an overview of the compara- tive morphology and ecology of the genus. Specifically, this paper (1) outlines the nomenclatural history of Strymon, (2) suggests that the best structure for dis- tinguishing Strymon is the unique morphology of the male genitalia valvae, which was first noted by Clench, (3) describes and illustrates morphological variation within the genus, (4) summarizes information on male behavior, larval foodplant specificity, and habitat, (5) preliminarily partitions Strymon species in nine species groups, and (6) transfers six names from Stry- mon to other genera. This work is intended to set the stage for a species revision, including the description of about five new species, mostly from the dry moun- tains of Peru and southern Ecuador. JOURNAL OF THE LEPIDOPTERISTS' SOCIETY MATERIALS AND METHODS The results in the this paper were based upon a comparison of adult morphology using the 6,000+ specimens of Strymon in the National Museum of Natural History (Smithsonian Institution, Washington, DC, USA), of which 3,972 are Neotropical, plus many specimens borrowed from other museums. This com- parison employed standard entomological techniques (Robbins 1991), including the examination of the male and female genitalia (449 dissections) of all species currently recognized in Strymon (Appendix 1), except that we relied on genitalic figures of two species. For those names that we could not identify from their original descriptions, we examined their types or pic- tures of the types. For only one species, Strymon an- thracaetus Salazar, Velez, & Johnson, could we not identify the name or find its type (explained below). In preparing a checklist of all Neotropical hairstreaks (Robbins in press), RKR examined the adult morphol- ogy of virtually all Neotropical species, although not in the same detail as with Strymon. All genitalic terms follow those in Klots (1970). All specific author names for Strymon are listed in Appendix 1 and thus are omitted from the following text. Because relationships within the Eumaeini are still poorly known, such as the genera that are most closely related to Strymon, we characterize Strymon by a complex and conspicuous trait that is unique within the Eumaeini and that is phylogenetically consistent with other traits that are unique within the Eumaeini. We tentatively divide Strymon into species group on the basis of many characters, but evidence for their monophyly awaits formal phylogenetic analysis. NOMENCLATURAL HISTORY Hiibner (1818) described Strymon and included two species, S. melinus and Hesperia acaciae Fabri- cius. (Hesperia currently belongs to the Hesperiidae.) A subsequent list of 13 Strymon species (Hiibner 1819) caused considerable confusion in the eventual selection of a type species. Scudder (1872) selected Hesperia titus F. from the 1819 list as the type (the dates of Hiibner s books were uncertain at the time), but Riley (1922) invalidated this selection and re- placed it with Strymon melinus. Finally, the Interna- tional Commission on Zoological Nomenclature (1959) placed Strymon on the Official List as Name No. 1332 with Strymon melinus as type. Hemming (1967) gives a more complete nomenclatural history. STRYMON HUBNER We characterize Strymon for the following discus- sion of comparative morphology by setae on the poste- rior dorsal surface of the valvae that are modified into anteriorly pointing "teeth" (Clench 1961) (Figs. 1-7). This structure is immediately recognizable; either a species has it or does not. The only exception is S. ziba (Fig. 8), whose valva morphology and systematic posi- tion are detailed below. Although the valva structure of Strymon is most easily illustrated using a scanning electron microscope (SEM), Clench discovered it us- ing a light microscope. Almost all species that have been placed in Strymon since 1960 (e.g., Ziegler 1960, Clench 1961,1964, Johnson et al. 1990, Johnson et al. 1992, Johnson & Kroenlein 1993, Johnson & Salazar 1993, Austin & Johnson 1997) have this valva struc- ture. The only superficially similar valvae in the Eu- maeini are those of Allosmaitia Clench, whose valvae are needle-like posteriorly, unlike Strymon, and cov- ered with teeth that are not anteriorly directed (Clench 1964). COMPARATIVE MORPHOLOGY Male genitalia. The setae on the valvae of eu- maeine hairstreaks are indistinguishable when viewed with an SEM from those specialized setae that were termed B-type trichoid sensilla (Ma & Schoonhoven 1973). In Strymon, these setae are modified. The an- teriorly directed teeth on the posterior dorsal surface of the valvae appear to be modifications of the sockets of these setae. Supporting this interpretation, the setal flagellum is still present, but is short (usually less than 1 u.m long) (Figs. 1-7). In S. ziba, some of the setal sockets on the posterior dorsal valva tips are slightly modified into anteriorly pointing teeth with a flagellum that is not reduced in length (Fig. 8). As discussed be- low, some evidence suggests that this structure is tran- sitional between that in other eumaeines and in Stry- mon while other evidence suggests that it is a further modification of that which occurs in other Strymon. We illustrate the male genitalia of nine Strymon species to show the range of morphological variation (Figs. 9-17). The tips of the gnathos in all Strymon species are subterminally expanded and then sharply tapered to a down-curved point (Figs. 9-17), but this structure does not appear to be sufficiently distinct from other eumaeines to distinguish the genus un- equivocally. At least one cornutus is present within the shaft of the penis unless the vesica is partially everted (Figs. 12, 14, 15). If there is one cornutus, it is long and slender (slightly wider in S. maritalis and S. christophei) and sometimes barely sclerotized (S. yo- joa, Fig. 11). If there are two cornuti, they are usually paired and about the same size (Fig. 16) (first noted and illustrated by Schwartz & Miller 1985). The pri- mary exception is S. ziba, which has one cornutus con- VOLUME 55, NUMBER 3 87 FIGS. 1-8. Scannh S. fflelinus at lower magj itron microen phs (SEMs ?garus; 8, and S. ziba (Arrow B ? slightly modified setal socket). Italia valva tips in lateral and dorsal aspects. 1, S. melinus in lateral aspect; 2, sal aspect (Arrow A ? setal flagellum); 5, S. bazochii; 6, S. gabatha; 7, S. JOURNAL OF THE LEPIDOPTERISTS' SOCIETY siderably larger than the other (Fig. 17). Strymon male genitalia are asymmetrical. For example, the penis is twisted and down turned (Clench 1961) except for S. ziba and S. sylea (Figs. 10, 17). The saccus is generally asymmetrical to the right (Figs. 9b, 12, 14, in ventral aspect, they are on the left side). Strymon species have paired brush organs (sensu Eliot 1973) that lie on the dorsal vinculum, but we omit them in the figures for clarity. The vinculum is not modified in structure, as it is in some eumaeines (Robbins 1991), except for slight projections of the vinculum in some species, such as S. tyleri and S. crambusa. We are unable to distinguish the structure of die brash organs in Strymon from those that occur in Lamprospilus Hiibner, Electrostrymon Clench, Ziegleria Johnson, and Calycopis Scudder. Males of S. istapa in one part of its range (Florida to Puerto Rico) may have or lack brush organs (Robbins & Nicolay 1999), and a similar dimorphism also appears to occur in S. bicolor. Other than the variation outlined above, the male genitalia of Strymon seem to have few good structures for distinguishing species. For example, the male gen- italia of S. istapa, S. columella, S. limenia, and S. tous- sainti are essentially indistinguishable except for small differences in size (Robbins & Nicolay 1999) ? these species are distinguished by their wing pattern and fe- male genitalia. Except for S. ziba and S. sylea, few Stry- mon species appear to be authoritatively identifiable solely on die basis of their male genitalic structures. Female genitalia. The bursa copulatrix of Strymon is asymmetrical, and we illustrate the range of varia- tion (Figs. 18-26). All eumaeines with a sclerotized, looped ductus bursae belong to Strymon, as first noted by Ziegler (1960), but the exact shape of the loop varies greatly interspecifically (Figs. 18-26). It is also highly variable intraspecifically, as in S. cestri, where some individuals lack the sclerotized loop of the duc- tus bursae, some have it, and others are intermediate between these extremes (Figs. 27-29). A few Strymon species appear to always lack the loop (S. yojoa, S. tegea, S. ohausi, S. sylea, and S. ziba), but the ductus bursae is twisted at the point where the loop would otherwise occur except in S. sylea (Figs. 19, 20, 23). The signa, which are "boat"-shaped, occur in all Stry- mon species, but may be small in some, such as S. li- menia (Hewitson) (Fig. 24), and may lack the anterior pointing spine (S. sylea, Fig. 19). Similar signa are found on occasion in other eumaeines, such as Tricho- nis Hewitson (Robbins 1987). There are two small teeth inside the anterior ductus bursae (Fig. 18). Sim- ilar teeth occur in other eumaeines, such as those il- lustrated for Rekoa palegon (Cramer) (Robbins 1991). The female 8th abdominal tergum of Eumaeini is sclerotized, the sternum is membranous, and two small circles lacking setae are also membranous (e.g., illustrated by Field 1941). Since spiracles on the 8th abdominal segment of the endoporian Ditrysia, in- cluding the butterflies, are absent or vestigial (Dug- dale 1974), we presume that these membranous cir- cles are vestigial spiracles. In female Strymon, there are two lands of 8th abdominal segments. The first, of which S. melinus is an example (Fig. 30), is like many eumaeines with presumed vestigial spiracles at the juncture between the tergum and sternum. The sec- ond, of which S. bicolor is an example (Fig. 31), has the presumed vestigial spiracles located within the ter- gum, whose latero-posterior part has more furrows than the first kind. Johnson et al. (1992) first described the furrowed tergum as "rough" and used it as a distin- guishing character of their new genus Heoda. How- ever, this structure is difficult to discern and actually occurs in all members of the S. istapa group (except for S. acis, Appendix 1), including species that Johnson et al. (1992) placed in Eiseliana and Strymon. When the ductus seminalis does not arise from the posterior tip of the corpus bursae in Strymon, such as S. limenia, the tip of the corpus bursae, posterior of the ductus seminalis, is lightly sclerotized (Fig. 24). All Strymon species with this structure are a subset of those whose females have an 8th tergum with the lat- ero-posterior surface furrowed and with imbedded presumed vestigial spiracles. The posterior end of the corpus bursae of a few other species, such as S. ziba, S. martialis, and S. christophei, is also sclerotized (Figs. 23, 26), but this structure appears to be analogous with the structure in S. limenia and relatives because the ductus seminalis arises from the posterior tip of the corpus bursae in these species. Wings. The wing venation of Strymon is typical of the Eumaeini with 10 forewing veins (Eliot 1973). A male scent patch (sensu Robbins 1991) occurs on the dorsal surface of the forewing in the distal part of the discal cell of most species, but is lacking in S. melinus, S. avalona, S. sabinus, S. tyleri, S. rufofusca, S. cyano- fusca, S. ohausi, S. christophei, S. oribata, and S. legota. Because these species appear to belong to a number of different species groups, as determined be- low on the basis of many characters, we suspect multi- ple losses of the forewing scent patch in Strymon. Wing pattern, shape, and size vary greatly among Strymon species (Figs. 34-53), and it is difficult to characterize Strymon on the basis of these traits. Gen- erally, the pattern on the ventral wing surface is not sexually dimorphic, but males have more sharply pro- duced forewing apices. Seasonal variation of the veil- VOLUME 55, NUMBER 3 89 FIGS 9-17 Male genitalia of Strymon; teeth on valvae are inconspicuous at this magnification. 9, S. melinus in lateral (a) and ventral (b) as- pects (Arrow D ? acuminate cornutus); 10, S. sylea (Arrow C ? tip of gnathos); 11, S. yojoa; 12, S. mulucha- 13, S. albata- 14, S. martialis; 15, S. limenia; 16, S. serapio (Arrow E ? paired cornuti); and 17, S. zifca. tral wing surface can be marked in some species, such as S. melinus, but virtually absent in others, such as S. bazochii. For example, individuals of S. melinus are smaller and darker on average in early spring in North America than they are in the middle of summer. The dorsal surface of the forewing of most female Strymon has a patch of black scales centered at the end of the discal cell, which is sometimes mistaken for androco- nia (Figs. 35, 40). Other eumaeines sometimes also have a similar appearing patch of dark scales, such as females oiTmolus venustus (Druce). Forewing length varies from more than 2 cm (S. sylea, S. oreala, and S. gabatha) to less than 0.8 cm (S. ohausi, S. ochraceus). And some species, such as S. gabatha and S. serapio, vary greatly in size intraspecifically, which is perhaps related to their bromeliad larval foodplants (flowers of Aechmea and Tillandsia, respectively). Head. The antennal club of Theclinae, including the Eumaeini, is generally cylindrical and incrassate, but those of most Strymon are abrupt and flattened, resembling those of Polyommatinae and Lycaeninae (Eliot 1973). An antenna with an abrupt and flattened club also occurs in a few other eumaeines, such as Penaincisalia Johnson. The frons of S. melinus is cov- ered with white and orange scales, and there are also orange scales near the base of the antennae. Most other Strymon share this head coloration, but a few lack white or orange scales, such as S. bazochii (Go- dart). However, a hairstreak with orange scales near the base of the antennae, an orange-white frons, and an abrupt, flattened antennal club is almost definitely a Strymon, which allows field recognition of this genus in most cases. BEHAVIOR AND ECOLOGY Male behavior. Male Strymon occupy mating ter- ritories, usually on hilltops and along forest edges. The males perch in the "territory," fly at other butterflies that enter this area, and then return to a perch very close to the original one unless courtship has ensued (e.g., Powell 1968, Robbins 1978, Alcock 1983, Alcock & O'Neill 1986, 1987, Cordero & Soberon 1990). We 90 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY 24a 24b 26a 26b FIGS 18-26. Female genitalia bursa copulatrix of Strymon. 18, S. vielmus in ventral (a) and lateral (b) aspects (Arrow F - signum); 19 S. salea- 20 S yoioa in ventral (a) and lateral (b) aspects; 21, S. mulucha (Arrow G - teeth inside ductus bursae); 22, S. abata; 23, S. iimrtial^ 24, st'Jl in ventral (a) and lateral (b) aspects; 25 S. serapio in ventral (a) and lateral (b) aspects; and 26, S. ziba in ventral (a) and lateral (b) aspects. VOLUME 55, NUMBER 3 91 FIGS. 27-29. Female genitalia variation of the bursa copulatrix of S. cestri. 27, Peru; 28, Costa Rica; 29, Mexico. have observed this behavior in the United States, Panama, Venezuela, Ecuador, Peru, and Brazil for 15 Strymon species (RKR unpubl). The males of most species occupy mating territories in the afternoon. A few species do so whenever the weather is favorable (i.e., S. tyleri, S. davara), and one species only in the early morning (males of S. serapio on hilltops in Panama and southern Brazil between 0730 and 1000 hours). The following discussion is based on our un- published observations. When male Strymon perch in a mating territory, they often open their wings 15-180 degrees, in addi- tion to moving their hindwings back and forth as in all Theclinae (Robbins 1980). We have observed the un- usual wing opening behavior (Figs. 54, 55) in S. meli- nus, S. mulucha, S. yojoa, S. davara, S. cestri, S. bubastus, S. dindus, S. ochraceus, S. ziba, S. megarus, S. azuba, and S. gabatha. Males of other hairstreak genera generally do not open their wings when perch- ing, but we have observed this behavior in some species of Tmolus and Chlorostrymon. Another un- usual male Strymon behavior is perching head down- wards on tree-trunks, in addition to perching on leaves. This behavior has been recorded in S. ziba, S. gabatha, S. ochraceus, and S. megarus. Only once have we seen a male hairstreak of another genus perch on a tree-trunk {Chalybs janias [Cramer]). Larval foodplants. The larvae of Strymon eat plants in more than 30 families, ranging from gym- nosperms to monocots and dicots, including Alstroe- meriaceae, Amaranthaceae, Begoniaceae, Boragi- naceae, Bromeliaceae, Cactaceae, Cannabidaceae, Chenopodiaceae, Compositae, Convolvulaceae, Crassu- laceae, Euphorbiaceae, Flacourtiaceae, Gesneriaceae, Gramineae, Guttiferae, Haemodoraceae, Juglandaceae, Labiatae, Leguminosae, Malvaceae, Melastomataceae, Musaceae, Orchidaceae, Pinaceae, Polygonaceae, Por- tulacaceae, Rosaceae, Sapindaceae, Sterculiaceae, Strelitziaceae, Surianaceae, Ulmaceae, and Verbe- naceae (RKR unpubl, plant family names follow Willis 1973). Some individual Strymon species, such as S. melinus, are exceedingly polyphagous (Ehrlich & Raven 1965, Tietz 1972), eating plant reproductive structures in most of these families. Many of these Strymon species, though, most frequently feed on plants in the Leguminosae and Malvaceae, and some are recorded as pests of beans and cotton (Ehrlich & Raven 1965). The only Eumaeini with larvae that eat plants in the Bromeliaceae belong to Strymon (RKR unpubl.). Some are serious pests of commercial pineapple (Har- ris 1927, Carter 1934, Fonseca 1934, Zikan 1956, Gua- gliumi 1965, 1967, DAraujo e Silva et al. 1967-1968, Beutelspacher 1972, Otero & Marigo 1990), but the agricultural literature refers to lycaenid pineapple pests as either Thecla echion L. (a misidentification of S. megarus or S. ziba, cf. Honey & Scoble 2001) or Thecla basalides (Geyer), a misspelling of Strymon basilides (Geyer), which, in turn, is a junior synonym of S. megarus (Appendix 1). Consequently, it is unclear exactly how many Strymon species feed on pineapples, but there are records for at least four species. The lar- vae of S. ziba, unlike those of the others, eat plants in a number of monocot families in addition to those of the Bromeliaceae (e.g., Harris 1927, Robbins & Aiello 1982). Habitat and range. Although Strymon occur in habitats ranging from tropical wet lowland rainforest to temperate climates, they are most diverse in xeric and seasonally very dry tropical areas, which includes most of the Pacific Coast of Mexico and Central America, northern Colombia and northern Vene- zuela, the mountains of Peru and southern Ecuador (where there are a number of undescribed species), the llanos of central Venezuela and surrounding countries, and eastern Brazil in the cerrado and caatinga life zones west to the Bolivian chaco. Stry- mon melinus is the only widespread Strymon in tem- perate North America while S. bicolor ranges from Peru in the Andes to Chile's temperate central valley (Santiago and surroundings), and S. eurytulus occurs from Bolivia and southern Brazil south into Patagonia (Argentina). 92 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY FIGS. 30-33. SEMs of female 8th abdominal tergum in dorso-lateral aspect. 30, S. melinus- 31, S. bicolor- 32, S. eurytulus (Arrow I - row in tergum); 33, S. bazochii. Arrows H ? presumed vestigial spiracles. -fur- STRYMON AND ITS SPECIES GROUPS The monophyly of Strymon, as we have character- ized it, is supported by a complex and conspicuous valva structure that is unique within the Eumaeini and whose presence or absence is unambiguous except for S. ziba. Other characters that are unique, or nearly so, within the Eumaeini are restricted to subsets of Stry- mon, adding further support to our characterization of Strymon as a monophyletic genus. All species with a scleortized looped ductus bursae belong to Strymon, as do all eumaeines with a ventro-lateral surface of the female 8th abdominal tergum that is furrowed with imbedded presumed vestigial spiracles. Behaviorally and ecologically, almost all eumaeine males that perch with their wings open 15-180? belong to Strymon (a behavior that may occur in all Strymon), as do all eu- maeine males that perch on tree-trunks (one exception noted above), and all species whose larvae eat plants in the Bromeliaceae. We have divided Strymon into nine species groups (listed in Appendix 1) on the basis of many characters, most of which were discussed above. Some of these groups, such as the S. istapa and S. serapio groups, are reasonably well-characterized while others, such as the S. mulucha and S. alhata groups, lack clear-cut derived distinguishing traits. For example, some characters suggest that S. cestri belongs to the S. yojoa species group, and others that it belongs to the S. mulucha species group. In the following paragraphs we charac- terize the S. istapa and S. serapio species groups and discuss the systematic positions of S. ziba and S. sylea. VOLUME 55, NUMBER 3 93 39 ' 'y> 35 37 BRilll V m ?&&&* Mill 1cm 41 ??? ?%?>.? 43 FIGS. 34^3. Dorsal and ventral wing surfaces. 34, male S. melinus (New Jersey, USA) (Arrow J ? patch of black scales on female dorsal forewing); 35, female S. melinus (New Jersey, USA); 36, male S. sylea (Loreto, Peru); 37, male S. yojoa (Panama Prov., Panama); 38, male S. mulucha (Canal Area, Panama); 39, male S. albata (Canal Area, Panama); 40, female S. martialis (Florida, USA); 41, male S. limenia (Santiago, Cuba); 42, male S. serapto (Santa Catarina, Brazil); and 43, male S. zifoa (Canal Area, Panama). We characterize those species with paired cornuti that are about the same size as the Strymon serapio group (Appendix 1). All recorded larval foodplants in this group are plants in the Bromeliaceae. In fact, ex- cept for S. ziba, all neotropical lycaenid larvae that use Bromeliaceae, including all pests of commercial pineap- ple, belong to this group. Except for S. ziba and one ob- servation of a male Chalybs, all males that are known to perch on tree-trunks belong to the S. serapio group. We characterize the Strymon istapa group as those species with a female 8th abdominal tergum whose ventro-lateral surface is furrowed with imbedded pre- sumed vestigial spiracles (Appendix 1). This tergum structure is otherwise unreported in the Eumaeini. Those species with a lightly sclerotized corpus bursae between the origin of the ductus seminalis and the posterior tip of the corpus bursae are a subset of this group. In all other Strymon, the ductus seminalis arises at the posterior tip of the corpus bursae. The in- clusion of S. acis in this species group is provisional, as explained in Appendix 1. The systematic position of S. ziba is unresolved be- cause of conflicting evidence. On the one hand, S. ziba appears to belong to the S. serapio group. Both have two cornuti, larvae that eat plants in the Bromeliaceae, and males that perch on tree trunks. The ventral wing pattern and male behavior of S. ziba and S. megarus are nearly identical, suggesting that they are sisters. If this systematic position is correct, then the unique valva structure of S. ziba is a modification of the Stry- mon valva. On the other hand, S. ziba appears to be the sister to the remainder of Strymon. Evidence sup- porting this systematic position is that the valva struc- ture of S. ziba appears to be intermediate between that of Strymon and other eumaeines. And unlike the S. serapio group, S. ziba has paired cornuti of unequal size, and its larvae eat plants in various monocot fami- lies, not just Bromeliaceae. This evidence suggests that S. ziba does not belong to the S. serapio group. Han- ner and Bobbins (in prep.) are trying to resolve this conflicting evidence using mitochondrial DNA se- quences. 94 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY FIGS. 44-45. Males perching with their wings open. 54, S. melinus (California, USA); 55, S. megarus (Veracruz, Mexico). The systematic position of S. sylea is tentative be- cause its penis is upturned and its ductus bursae is straight, unlike virtually all other Strymon species. The male genitalia valvae of S. sylea possess the unusual basally directed teeth of Strymon when viewed with a light microscope. The tips of the gnathos, scent patch, and antennal club of S. sylea are consistent with Stry- mon, and we cannot place S. sylea in any other eu- maeine genus. Because S. sylea is an exceedingly rare species in collections, we have not had the opportunity to examine its valvae using an SEM, which would de- finitively confirm that the valva teeth are indeed the Strymon type. The placement of S. sylea in Strymon appears to be the best option for now. NOMENCLATURE Generic synonymy. Using our characterization of Strymon, based largely on that of Clench (1961), there are five generic synonyms of Strymon, listed with their type species in parentheses. Citations to all original de- scriptions for this and the following list can be found in Lamas et al. (1995). Strymon Hubner 1818 (Rusticus melinus Hiibner) Callipareus Scudder 1872 (Strymon melinus Hubner) Callicista Grote 1873 {Callicista ocellifera Grote) Uranotes Scudder 1876 (Strymon melinus Hubner) Eiseliana Toledo 1978 (Eiseliana koehleri Toledo) Heoda Johnson, L. Miller & Herrera 1992 (Thecla heodes Druce) Our characterization of Strymon appears to be reasonable despite the uncertain systematic position of S. ziba. If S. ziba belongs to the S. serapio group, which is supported by some evidence, then S. ziba clearly belongs to Strymon. If, on the other hand, S. ziba is the sister to the remainder of Strymon, which is supported by other evidence, then placing S. ziba in Sti~ymon avoids naming a monotypic genus for S. ziba. The generic names Callicista, Eiseliana, and Heoda could be applied to the S. istapa species group, but we believe that our characterization of Strymon is more reasonable and stable. Recognizing Callicista, Eise- liana, or Heoda would leave the name Strymon for the remainder of species with anteriorly directed teeth on the dorsal valvae, but there is no evidence that this re- maining group of species is monophyletic. And if a generic name were also proposed for the S. serapio group, there would still be no evidence for the mono- phyly of Strymon. Lastly, our characterization of Stry- mon is consistent with the way that Strymon has been used in North America for the last 40 years (Ziegler 1960, Clench 1961). Species groups. We list the 183 names that belong to Strymon as we have characterized it, partitioned into species groups (Appendix 1). We list the charac- ters for each species group, but many characters are homoplastic, and formal phylogenetic analysis is nec- essary to establish the monophyly of these groups. The original description of Strymon anthracaetus Salazar, Velez, & Johnson, 1997 was too poor to identify this name or to determine whether it belongs to Strymon. In August 2000 G. Lamas could not find the type at the museum in Manizales, Colombia, where it was supposed to be deposited. Consequently, we regard S. anthracaetus as a nomen dubium. Names removed from Strymon. Many names that were described in Strymon do not belong to Stry- mon as we have characterized it. We confirm the pre- vious transfer to other genera (Bridges 1988) of 23 VOLUME 55, NUMBER 3 95 names that were originally described in Strymon (Ap- pendix 2). We transfer the following six species, which were re- cently described in Strymon, to other genera and give brief reasons for the new generic placement. One is transferred to "Thecla" because it belongs to a genus that is yet undescribed. 1. Strymon angulus (Le Crom & Johnson, 1997) is transferred to Thereus Hiibner, new combination. Robbins (1991, 2000) characterized Thereus by a pair of sclerotized invaginations on the membrane attached to the ventro-lateral sides of the papillae anales, a pair of ventro-lateral brush organs in addition to the pair of dorsal ones, and sexual dimorphism in the antennal club (>4 more nudum segments in the female than the male). Although these characters are not mentioned in the original description of S. angulus, the illustrated holotype is a male of Thereus endera Hewitson, which possesses the three synapomorphies of Thereus listed above (Robbins in press). 2. Strymon daplissus Johnson & Salazar, 1993 is transferred to Ministrymon Clench, new combina- tion. The illustrated holotype of S. daplissus is a male of Thecla clytie Edwards, which Clench (1961) placed in Ministrymon. 3. and 4. Strymon carmencitae (Le Crom & Johnson, 1997) and Strymon cryptogramus (Johnson, Eisele & MacPherson, 1992) are transferred to Nicolaea John- son, new combinations. We characterize Nicolaea by its male genitalia vinculum, which is strongly curved an- teriorly in lateral aspect and which is flattened dorsally in anterior aspect. These characters cannot be seen in the original descriptions, but die illustrated holotypes of Strymon carmencitae and Strymon cryptogramus are specimens of Nicolaea fabulla (Hewitson, 1868) andN. torris (Druce, 1907), respectively, which possess the vinculum of Nicolaea as described (Robbins in press). 5. Strymon nivnix (Johnson, Eisele & MacPherson, 1990) is transferred to Calycopis Scudder, new com- bination. The forewings of the holotype of S. nivnix belong to a different genus than the hindwings (Rob- bins in press), which appear to be the slightly aberrant hindwings of Calcyopis cecrops F. We designate the hindwings as the lectotype, which is placed in Calycopis as characterized by Clench (1961) and Field (1967). 6. Strymon additionalis (Le Crom & Johnson, 1997) is transferred to Thecla F, new combination. The holotype is a male of "Thecla" emessa Hewitson, 1867, which is characterized by the form of its scent patch on the dorsal surface of the forewings, a white medial stripe on the frons, and a penis tip with its ventral sur- face flattened. Because the latter two characters were not illustrated in the original description, our identifi- cation is based upon the illustrated wing pattern and scent patch. ACKNOWLEDGMENTS We dedicate this paper to the memories of J. B. Ziegler and H. K. Clench, who together modernized the generic taxonomy of North American hairstreaks. For the loan or gift of Strymon specimens over the years, we are grateful to P. Ackery, C. Brevignon, C. Callaghan, M. Casagrande, J.-Y. Gallard, J. Glassberg, G. Lamas, J. MacDonald, L. Miller, Jacqueline Miller, James Miller, O. Mielke, J. Rawlins, B. Sullivan, R. Vane-Wright, and J. Weintraub. We thank J. Glassberg for allowing us to use his photographs of perching Strymon males. We are grateful to V. Malikul for help with photography, S. Braden with the SEM, and G. Venable with digitization and layout of the fig- ures. For detailed comments on the manuscript, we thank J. Brown, J. Burns, P. Gentili-Poole, J. Glassberg, J. Hall, D. Harvey, G. Lamas, C. Penz, B. Sullivan, and the late B. Ziegler. LITERATURE CITED ALCOCK, J. 1983. Territoriality by hilltopping males of the great purple hairstreak, Atlides halesus (Lepidoptera, Lycaenidae): convergent evolution with a pompilid wasp. Behav. Ecol. Socio- biol. 13:57-62. ALCOCK, J. & K. M. O'NEILL. 1986. Density-dependent mating tactics in the grey hairstreak, Strymon melinus (Lepidoptera: Lycaenidae). J. Zool. Series A 209:105-113. . 1987. Territory preferences and intensity of competition in the grey hairstreak Strymon melinus (Lepidoptera: Lycaenidae) and the tarantula hawk wasp Hemipepsis ustulata (Hy- menoptera, Pompilidae). Am. Midi. Nat. 118:128-138. AUSTIN, G. T. & K. JOHNSON. 1997. 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Characters: Male lacking a scent patch on the dorsal surface of the FW, no basal patch of white scales on the ventral surface of the HW (except S. tyleri), penis tip down-turned with a single slender cornutus, ductus bursae with simple sclerotized loop, ductus seminalis arises from the un- sclerotized posterior end of the corpus bursae, female with 8th tergum unfurrowed and lacking imbedded vestigial spiracles. Remarks: Some members, such as S. melinus, S. tyleri, and S. rufofusca, can be exceed- ingly common where they occur. 1. Strymon melinus (Hiibner, 1813) SYNONYMS AND SUBSPECIES: Thecla hyper- ici (Boisduval & Leconte, 1835), Thecla humuli (Harris, 1841), Thecla pudica (H. Edwards, 1877), Strymon atrofasciata McDunnough, 1921, Strymon setonia McDunnough, 1927, Strymon meinersi Gunder, 1927, Thecla clarionensis (Heid, 1933), Strymon youngi Field, 1936, Stry- mon franki Field, 1938, Strymon caldasensis Salazar, Velez & K. Johnson, 1997 2. Strymon avalona (W.G. Wright, 1906) 3. Strymon tyleri (Dyar, 1913) 4. Strymon sabinus (C. Felder & R. Felder, 1865) SYNONYM: Thecla promissa (Moschler, 1883) 5. Strymon rufofusca (Hewitson, 1877) SYNONYMS AND SUBSPECIES: Thecla (Uran- otes) valentina (Berg, 1896), Thecla lucaris (A.G. Weeks, 1901), Thecla grisea (Dufrane, 1939), Thecla nigriplaga (Dufrane, 1939), Strymon gua- nensis Le Crom & K. Johnson, 1997 6. Strymon cyanofusca K. Johnson, Eisele & MacPherson, 1990 STRYMON ALBATA GROUP. Characters: Male with a scent patch on the dorsal surface of the FW, no basal patch of white scales on the ventral surface of the HW (except S. alhata), penis tip down-turned with a single slender cornutus, ductus bursae with simple sclerotized loop, ductus seminalis arises from the unsclerotized posterior end of the corpus bursae, female with 8th ter- gum unfurrowed and lacking imbedded vestigial spira- cles. Remarks: The only consistent difference with the previous group is that the males have a scent patch. 7. Strymon alhata (C. Felder & R. Felder, 1865) SYNONYM: Thecla sedecia (Hewitson, 1874) 8. Strymon alea (Godman & Salvin, 1887) SYNONYM: Callicista laceyi (Barnes & McDun- nough, 1910) 9. Strymon bebrycia (Hewitson, 1868) SYNONYMS: Thecla chonida (Hewitson, 1874), Strymon buchholzi H.A. Freeman, 1950 STRYMON YOJOA GROUP. Characters: Male with a scent patch on the dorsal surface of the FW (except S. ohausi), no basal patch of white scales on the ventral surface of the HW (except S. yojoa), penis tip down- turned with a single slender cornutus that is barely sclerotized, ductus bursae with a twist, but lacking a sclerotized loop, ductus seminalis arises from the unsclerotized posterior end of the corpus bursae, fe- male with 8th tergum unfurrowed and lacking imbed- ded vestigial spiracles. Remarks: Johnson et al. (1992) noted that Thecla tegea Hewitson does not belong to Strymon, but its male genitalia have anterior pointing teeth on the valvae, and its male and female genitalia are exceedingly similar to those of S. yojoa. 10. Strymon yojoa (Reakirt, 1867) SYNONYMS AND SUBSPECIES: Thecla claraba (Hewitson, 1867), Thecla heroea (Hewitson, 1868) 11. Strymon tegaea (Hewitson, 1868) SYNONYM: Thecla seitzi (Spitz, 1931) 12. Strymon ohausi (Spitz, 1933) STRYMON MULUCHA GROUP. Characters: Male with a scent patch on the dorsal surface of the FW, with basal patch of white scales on the ventral surface of the HW, penis tip down-turned with a single slender cornutus, ductus bursae with simple sclerotized loop (except for some individuals of S. cestri), ductus semi- nalis arises from the unsclerotized posterior end of the corpus bursae, female with 8th tergum unfurrowed and lacking imbedded vestigial spiracles. Remarks: This species group is probably paraphyletic with re- spect to the previous group. 13. Strymon mulucha (Hewitson, 1867) SYNONYMS: Tmolus invisus (Butler & H. Druce, 1872), Strymon necjebus Le Crom & K. Johnson, 1997, Strymon necjabus Le Crom & K. Johnson (missp.), 1997, Strymon novasignum Austin & K. Johnson, 1997, Strymon clavus Austin & K. John- son, 1997, Strymon implexus Austin & K. Johnson, 1997, Strymon inmirum Austin & K. Johnson, 1997, Strymon incanus Austin & K. Johnson, 1997 14. Strymon cestri (Reakirt, 1867) SYNONYMS AND SUBSPECIES: Thecla cydia (Hewitson, 1874), Thecla crossoea (Hewitson, 1874), Strymon chamiensis Salazar, Velez & K. 98 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY Johnson, 1997, Strymon germana Austin & K. Johnson, 1997 15. Strymon davara (Hewitson, 1868) SYNONYMS: Thecla joannisi (Dufrane, 1939), Thecla pallida (Dufrane, 1939) 16. Strymon crambusa (Hewitson, 1874) 17. Strymon astiocha (Prittwitz, 1865) SYNONYMS: Thecla faunalia (Hewitson, 1868), Thecla deborrei (Capronnier, 1874), Strymon ha- los Austin & K. Johnson, 1997, Strymon consper- gus Austin & K. Johnson, 1997 STRYMON MARTIALIS GROUP. Characters: Male with or without a scent patch on the dorsal surface of the FW, no basal patch of white scales on the ventral surface of the, penis tip down-turned with a single "wide" cornutus, ductus bursae with sclerotized loop complex, especially in S. martalis (Fig. 23), ductus seminalis arises from the posterior end of the corpus bursae which is sclerotized dorsally, fe- male with 8th tergum unfurrowed and lacking imbed- ded vestigial spiracles. Remarks: A sister species rela- tionship between the two included species is a new hypothesis, but one that appears to be reasonably strongly supported by the shape of the cornutus and by the sclerotized patch on the corpus bursae. This species group may be more closely related to the S. melinus species group than its position here indicates. 18. Strymon martialis (Herrich-Schaffer, 1865) 19. Strymon christophei (W.P. Comstock & Hunting- ton, 1943) STRYMON 1STAPA GROUP. Characters: Male with a scent patch on the dorsal surface of the FW (except for S. oribata), no basal patch of white scales on the ven- tral surface of the HW, penis tip down-turned with a single slender cornutus, ductus bursae with sclerotized loop simple except in S. acts, ductus seminalis arises either from the unsclerotized posterior end of the cor- pus bursae or from a point anterior of a sclerotized patch on the dorsal surface of the corpus bursae, fe- male with 8th tergum furrowed (except in S. acts) and with imbedded presumed vestigial spir- acles. Remarks: The inclusion of S. bazochii and S. acis in this group are new hypotheses. The latter species is included because it has imbedded presumed vestigial spiracles and basal spots on the ventral surface of the HW, which most of the species in this group share. Its inclusion is highly tentative because its fe- male 8th abdominal tergum lacks furrows and the sclerotized loop of its ductus bursae is exceedingly complex, unlike any otfier Strymon species. 20. Strymon istapa (Reakirt, 1867) SYNONYMS AND SUBSPECIES: Lycaena mod- esta (Maynard, 1873), Callicista ocellifera (Grote, 1873), Thecla cybira (Hewitson, 1874), Thecla arecibo (W.P. Comstock & Huntington, 1943), Thecla clarionica (Vazquez, 1958), Thecla socor- roica (Vazquez, 1958), Strymon clenchi Austin & J.F Emmel, 1998 21. Strymon bazochii (Godart, 1824) SYNONYMS: Hyreus thius (Geyer, 1832), Thecla agra (Hewitson, 1868), hecla infrequens (A.G. Weeks, 1901), Strymon gundlachianus M. Bates, 1935, Strymon diagonalis Austin & K. Johnson, 1997 22. Strymon acis (Drury, 1773) SYNONYMS AND SUBSPECIES: Papilio mars (Fabricius, 1776), Thecla gossei (W.P. Comstock & Huntington, 1943), Thecla bartrami (W.P. Comstock & Huntington, 1943), Thecla casasi (W.P. Comstock & Huntington, 1943), Thecla petioni (W.P. Comstock & Huntington, 1943), Strymon armouri Clench, 1943, Strymon leu- costicha Clench, 1992 23. Strymon columella (Fabricius, 1793) SYNONYMS: Papilio dion (Schaller, 1788), Tmo- lus erytalus (Butler, 1870), Thecla antigua (W.P. Comstock & Huntington, 1943) 24. Strymon limenia (Hewitson, 1868) 25. Strymon toussainti (W.P. Comstock & Huntington, 1943) SYNONYMS: Strymon andrewi K. Johnson & Ma- tusik, 1988, Strymon rhaptos K. Johnson, Eisele & MacPherson, 1990, Strymon amonensis D.S. Smith, K. Johnson, J.Y. Miller & McKenzie, 1991 26. Strymon bubastus (Stoll, 1780) SYNONYMS AND SUBSPECIES: Papilio minereus (Fabricius, 1787), Thecla salona (He- witson, 1868), Thecla sapota (Hewitson, 1877), Thecla peruensis (Dufrane, 1939), Thecla ponce (W.P. Comstock & Huntington, 1943), Strymon vividus Le Crom & K. Johnson, 1997 27. Strymon eurytulus (Hiibner, 1819) SYNONYMS AND SUBSPECIES: Thecla ameri- censis (Blanchard, 1852), Thecla argona (Hewit- son, 1874), Thecla rana (Schaus, 1902, Thecla tu- cumana (H.H. Drace, 1907), Theclanigra (Lathy, 1926), Strymon peristictos K. Johnson, Eisele & MacPherson, 1990, Heoda nivea (K. Johnson, L.D. Miller & Herrera, 1992) 28. Strymonflavaria (Ureta, 1956) SYNONYM: Heoda erani (Benyamini & K. John- son, 1996) 29. Strymon ollantaitamba (K. Johnson, L.D. Miller & Herrera, 1992) VOLUME 55, NUMBER 3 99 30. Strymon colombiana (K. Johnson, L.D. Miller & Herrera, 1992) SYNONYM: Heoda lecromi (K. Johnson & Lugo, 1997) 31. Strymon bicolor (Philippi, 1859) SYNONYMS AND SUBSPECIES: Thecla quadri- maculata (Hewitson, 1874), Thecla heodes (H. H. Druce, 1909), Thecla leptocosma (Hayward, 1949), Thecla tricolor (Ureta, 1949), Heoda ata- cama (K. Johnson & L.D. Miller, 1992), Eiseliana probabila (K. Johnson, L.D. Miller & Herrera, 1992), Heoda suprema (K. Johnson, L.D. Miller & Herrera, 1992), Heoda shapiroi (K. Johnson, L.D. Miller & Herrera, 1992) 32. Strymon wagenknechti (Ureta, 1947) 33. Strymon oribata (Weymer, 1890) SYNONYMS AND SUBSPECIES: Thecla areni- cola (Jorgensen, 1934), Thecla punona (Clench, 1944), Thecla rojasi (Ureta, 1956), Eiseliana koehleri (Toledo, 1978), Eiseliana patagoniensis (K Johnson, L.D. Miller & Herrera, 1992) STRYMON SYLEA GROUP. Characters: Male with a scent patch on the dorsal surface of the FW, no basal markings on the ventral surface of the HW, penis tip up-turned with a single slender cornutus, ductus bur- sae straight without a loop or twist, ductus seminalis arises either from the unsclerotized posterior end of the corpus bursae, female with 8th tergum unfur- rowed and lacking imbedded vestigial spiracles. Re- marks: The upturned penis tip and straight ductus bursae, which is like most Eumaeini other than Stry- mon, suggest that the single species in this group may be sister to the remainder of the genus. 34. Strymon sylea (Hewitson, 1867) STRYMON SERAPIO GROUP. Characters: Male with a scent patch on the dorsal surface of the FW, no basal patch of white scales on the ventral surface of the HW, penis tip down-turned with paired cornuti, ductus bursae with simple sclerotized loop, ductus seminalis arises from the unsclerotized posterior end of the cor- pus bursae, female with 8th tergum unfurrowed and lacking imbedded vestigial spiracles. Remarks: All lar- val foodplant records in this groups are Bromeliaceae, as noted. 35. Strymon serapio (Godman & Salvin, 1887) SYNONYMS AND SUBSPECIES: Thecla lemnos (H.H. Druce, 1890), Thecla mesca (Dyar, 1914), Thecla inconspicua (Lathy, 1930), Strymon gol- bachi K. Johnson, Eisele & MacPherson, 1990, Strymon trunctogen K. Johnson & Salazar, 1993, Strymon altamiraensis K. Johnson & Kroenlein, 1993, Strymon henaoi Salazar, Velez & K. Johnson, 1997, Strymon hurtadoi K. Johnson, 1997, Stry- mon rosari Torres & K. Johnson, 1997, Strymon originatus K. Johnson, Hernandez & Cock, 1997 36. Strymon glorissima K. Johnson & Salazar, 1993 SYNONYM: Strymon campbelli K. Johnson & Salazar, 1993 37. Strymon gabatha (Hewitson, 1870) SYNONYMS: Thecla balius (Godman & Salvin, 1887), Strymon alexandra K. Johnson & Kroen- lein, 1993, Strymon alicia Salazar, Velez & K. Johnson, 1997 38. Strymon monopeteinus Schwartz & J.Y Miller, 1985 39. Strymon veterator (H.H. Druce, 1907) SYNONYMS: Strymon lorrainea K. Johnson, Eisele & MacPherson, 1990, Strymon coronos K. Johnson, Eisele & MacPherson, 1990 40. Strymon oreala (Hewitson, 1868) 41. Strymon dindus (Fabricius, 1793) 42. Strymon ochraceus K. Johnson & Salazar, 1993 SYNONYM: Strymon baricharensis Le Crom & K. Johnson, 1997 43. Strymon lucena (Hewitson, 1868) SYNONYMS: Thecla cardus (Hewitson, 1874), Thecla canitus (H.H. Druce, 1907), Strymon spe- cials K. Johnson, Eisele & MacPherson, 1997 44. Strymon legota (Hewitson, 1877) 45. Strymon azuba (Hewitson, 1874) SYNONYMS: Strymon montevagus K. Johnson, Eisele & MacPherson, 1990, Strymon rojos K Johnson & Kroenlein, 1993 46. Strymon eremica (Hayward, 1949) SYNONYMS AND SUBSPECIES: Strymon lariy- ojoa K. Johnson, Eisele & MacPherson, 1990, Strymon barbara K. Johnson, Eisele & MacPher- son, 1990, Strymon nicolayi K. Johnson, Eisele & MacPherson, 1990 47. Strymon megarus (Godart, 1824) SYNONYMS AND SUBSPECIES: Tmolus basilides (Geyer, 1837), Thecla basalides (W.F. Kirby, 1871) (missp.), Thecla tigonia (Schaus, 1902), Strymon amphyporphijra K. Johnson, Eisele & MacPherson, 1990, Strymon rotundum Austin & K. Johnson, 1997, Strymon gallardi Faynel & K. Johnson, 2000 STRYMON ZIRA GROUP. Characters: Male with a scent patch on the dorsal surface of the FW, no basal patch of white scales on the ventral surface of the HW, penis tip up-turned with two large unpaired cornuti, ductus bursae slightly twisted without a sclerotized loop, ductus seminalis arises from the sclerotized pos- terior end of the corpus bursae, female with 8th ter- 100 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY gum unfurrowed and lacking imbedded vestigial spir- acles. Remarks: The single included species is wide- spread and common. It is the only Strymon lacking the clear-cut anterior pointing teeth on the valvae. 48. Strymon ziba (Hewitson, 1868) SYNONYMS AND SUBSPECIES: Thecla thulia (Hewitson, 1868), Thecla diagutta (Hayward, 1949), Strymon baptistorum K. Johnson, Eisele & MacPherson, 1990, Strymon dondiego K. Johnson & Adams, 1997, Strymon profusorubrus Le Crom & K. Johnson, 1997, Strymon lecromi K. Johnson, 1997, Strymon spinatus Austin & K. Johnson, 1997, Strymon latamaculus Austin & K. Johnson, 1997, Strymon pallidulus Austin & K. Johnson, 1997, Stry- mon tholus Austin & K. Johnson, 1997 APPENDIX 2. Alphabetical list of specific names that were originally described in Strymon and that have been transferred to other genera (Bridges 1988). For those names that are junior synonyms, we note its se- nior synonym. 1. Cyanophrys agricolor (Butler & Druce, 1872) 2. Satyrium aliparops (Michener & dos Passos, 1942), a junior synonym of Satyrium liparops (Leconte) 3. Satyrium borealis (Lafontaine, 1969), a junior syn- onym of Satyrium calanus (Hiibner) 4. Satyrium caryaevorus (McDunnough, 1942) 5. Satyrium chlorophora (Watson & Comstock, 1920), a junior synonym of Satyrium saepium (Boisduval) 6. Lamprospilus coelicolor (Butler & Druce, 1872) 7. Satyrium coolinensis (Watson & Comstock, 1920), a junior synonym of Satyrium acadica (Edwards) 8. Satyrium desertorum (Grinnell, 1917), a junior synonym of Satyrium sylvinus (Boisduval) 9. Electrostrymon dowi (Clench, 1941), a junior syn- onym of Electrostrymon angelia (Hewitson) 10. Satyriumfletcheri (Michener & dos Passos, 1942), a junior synonym of Satyrium liparops (Leconte) 11. Satyrium godarti (Field, 1938), a junior synonym of Satyrium calanus (Hiibner) 12. Calycopis gottschalh (Clark & Clark, 1938), a ju- nior synonym of Calycopis cecrops (Fabricius) 13. Satyrium immaculosus (Comstock, 1913), a junior synonym of Satyrium titus (Fabricius) 14. Satyrium kingi (Klots & Clench, 1952) 15. Satyrium montanensis (Watson & Comstock, 1920), a junior synonym of Satyrium acadica (Ed- wards) 16. Satyrium muskoka (Watson & Comstock, 1920), a junior synonym of Satyrium acadica (Edwards) 17. Satyrium okanagana (McDunnough, 1944), a ju- nior synonym of Satyrium saepium (Boisduval) 18. Cyanophrys pastor (Butler & Druce, 1872), a ju- nior synonym of Cyanophrys longula (Hewitson) 19. Satyrium polingi (Barnes & Benjamin, 1926) 20. Satyrium provo (Watson & Comstock, 1920), a ju- nior synonym of Satyrium saepium (Boisduval) 21. Satyrium swetti (Watson & Comstock, 1920), a ju- nior synonym of Satyrium acadica (Edwards) 22. Satyrium violae (Stallings & Turner, 1947), a junior synonym of Satyrium favonius (Smith) 23. Satyrium watsoni (Barnes & Benjamin, 1926), a junior synonym of Satyrium titus (Fabricius)