BULLETIN UNIVERSITY OF FLORIDA GAINESVILLE A REVIEW OF VIRBIA (FORMERLY HOLOMELINA) OF AMERICA NORTH OF MEXICO (ARCTIIDAE: ARCTIINAE: ARCTIINI) J. M. Zaspel, S. J. Weller, and R.T. Cardé Vol. 48, No. 3, pp. 59-118 2008 ™ BULLETIN UNIVERSITY OF FLORIDA GAINESVILLE A REVIEW OF VIRBIA (FORMERLY HOLOMELINA) OF AMERICA NORTH OF MEXICO (ARCTIIDAE: ARCTIINAE: ARCTIINI) J. M. Zaspel, S. J. Weller, and R.T. Cardé Vol. 48, No. 2, pp. 59-118 2008 ™ BULLETIN ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 119 The FLORIDA MUSEUM OF NATURAL HISTORY is Florida’s state museum of natural history, dedicated to understanding, preserving, and interpreting biological diversity and cultural heritage. The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY is a peer-reviewed publication that publishes the results of original research in zoology, botany, paleontology, and archaeology. Address all inquiries to the Managing Editor of the Bulletin. Numbers of the Bulletin are published at irregular intervals. Specific volumes are not necessarily completed in any one year. The end of a volume will be noted at the foot of the first page of the last issue in that volume. Richard Franz, Managing Editor Cathleen Bester, Production Bulletin Committee Richard Franz, Chairperson Ann Cordell Sarah Fazenbaker Richard Hulbert William Marquardt Larry Page Irvy R. Quitmyer David W. Steadman, Ex officio Member ISSN: 0071-6154 Publication Date: December 30, 2008 Send communications concerning purchase or exchange of the publication and manuscript queries to: Managing Editor of the BULLETIN Florida Museum of Natural History University of Florida PO Box 117800 Gainesville, FL 32611-7800 U.S.A. Phone: 352-392-1721 Fax: 352-846-0287 e-mail: lrfranz@flmnh.ufl.edu ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 59 1 Dept. of Entomology and Nematology, P.O. BOX 110620 Natural Area Drive, University of Florida, Gainesville, FL 32611 2 Dept. of Entomology, 1980 Folwell Ave., 219 Hodson Hall, University of Minnesota, St. Paul, MN 55108 3 Bell Museum of Natural History, University of Minnesota, 1980 Upper Buford Circle, St. Paul, MN 55108 4 Dept. of Entomology, Univeristy of California-Riverside, Riverside CA 92521 Zaspel, J.M, S.J. Weller, and R.T. Cardé. 2008. A Review of Virbia (Formerly Holomelina) of America North of Mexico (Arctiidae: Arctiinae: Arctinii) Bull. Florida Museum Nat. Hist. 48(3): 59-118. A REVIEW OF VIRBIA (FORMERLY HOLOMELINA) OF AMERICA NORTH OF MEXICO (ARCTIIDAE: ARCTIINAE: ARCTIINI) J. M. Zaspel1, S. J. Weller 2,3, and R.T. Cardé4 ABSTRACT The tiger moth genus Virbia Walker (Arctiinae: Arctiini) is redescribed and its biology and taxonomic history are reviewed. The recent placement of Holomelina Herrich-Schäffer as its junior synonym is examined. North of Mexico, 14 Virbia species are recognized, including two new species, V. fergusoni Zaspel and V. rindgei Cardé. Adult habitus photographs, a checklist, diagnoses, and complete descriptions are provided for all species. Genitalia of the new species are illustrated, and diagnostic character illustrations are provided for separating all previously described species. Locality information from over 12,000 specimens representing 16 museum collections [in the United States and Canada] is compiled and available in an electronic database on the Diversity of Life Website (at www.diversityoflife.org/virbia.html). As some species appear to be habitat specialists (e.g., V. lamae in bogs), photos of selected habitats are provided. TABLE OF CONTENTS Introduction........................................................................................................60 Biology of Virbia in Canada and the United States.............................................62 Taxonomic History of Holomelina and Virbia.................................................... 68 Checklist of Virbia..............................................................................................72 Materials and Methods ..................................................................................... 73 Systematic Entomology of Genus Virbia............................................................76 V. aurantiaca................................................................................................ 77 V. costata......................................................................................................81 V. fergusoni n. sp.........................................................................................84 V. ferruginosa...............................................................................................87 V. fragilis......................................................................................................88 V. immaculata...............................................................................................89 V. laeta..........................................................................................................91 V. lamae........................................................................................................94 V. marginata.................................................................................................97 V. nigricans.................................................................................................. 98 V. opella....................................................................................................... 101 V. ostenta..................................................................................................... 102 V. rindgei n. sp............................................................................................ 104 V. rubicundaria.............................................................................................106 Acknowledgements.............................................................................................107 Literature Cited.................................................................................................. 107 Key Words: Tiger moths, Virbia, Arctiidae, morphology, lepidopteran taxonomy. 60 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) INTRODUCTION Colorful and enigmatic, the species of North American Virbia Walker, formerly Holomelina Herrich-Shäffer, have challenged lepidopterists’ species concepts and occasionally their civility towards one another, since the mid-1800s (e.g., scientific exchanges reviewed in Smith, 1889). The phenotypic diversity of adults of some spe- cies is impressive. For example, within a single reared brood of V. aurantiaca (Hübner), several phenotypes occur, and several of these were named as species (Fig. 1). Such phenotypic diversity when combined with in- adequate illustrations and descriptions of many species has resulted in complicated taxonomic history and con- fusion. Further, only subtle differences in genitalic mor- phology exist among the males of some species whose phenotypes are easily confused (e.g., V. aurantiaca and V. nigricans [Reakirt]) (Ferguson 1985; Forbes 1960; Cardé 1968, 1971). Recently, a phylogenetic review of generic limits of Virbia proposed that Virbia (Walker 1854) and Holomelina (Herrich-Schäffer 1855) were congeneric, and Holomelina was placed as the junior synonym (Zaspel 2004; Zaspel & Weller 2006). Historically, these two genera have been associated with each other (e.g., Barnes & McDunnough 1917; Forbes 1939, Forbes 1960); Ferguson (1985) suggested that the genera might be congeneric when he described the Holomelina group. Rare in Arctiidae, males possess a true scaphium, and many species also possess a subscaphium (Ferguson 1985; Zaspel 2004; Zaspel & Weller 2006). As currently defined, Virbia is comprised of small to medium sized, slender, brightly colored species (Fig. 2). The genus is distributed from south central Canada (e.g., northern Manitoba) to the North Atlantic (e.g., Nova Scotia), south to central South America (Fig. 3) (Handfield 1999; Zaspel & Weller 2006). North Ameri- can species (formerly Holomelina) are slender moths with red or pink adbasal regions in the hindwings (Figs. 2, 4). Neotropical species typically have the red pig- ment replaced with yellow hindwing patches (Fig. 5). Larvae possess homoideous crochets, a secondary loss of the smaller lateral crochets found in other arctiines (Kitching & Rawlins 1999; Jacobson & Weller 2002). Larvae are otherwise typical of Arctiini with clusters of secondary setae on raised verrucae (Kitching & Rawlins 1999; Jacobson & Weller 2002). Some North Ameri- can species have been reared on dandelion and other introduced Plantaginaceae (e.g., Forbes 1960). Typical for Arctiini, larvae are reported as feeding on unidenti- fied herbaceous plants; native hosts (pre-European settle- ment) are unknown (Ferguson 1985; Tietz 1972). Although little is known about their biology, spe- cies in this genus have figured prominently in the litera- ture on chemical ecology and pheromones (Roelofs & Figure 1. Reared series, Virbia aurantiaca (Top Row: Female parent of brood, second and third rows: Males) ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 61 Cardé 1971; Cardé & Roelofs 1973; Cardé 1974; Charlton & Roelofs 1991; Schal & Cardé 1985; Schal & Cardé 1986; Schal et al. 1987; Yin et al. 1991; Schal et al. 1998). Virbia aurantiaca was among the first non-pest lepidopterans to have its female pheromones characterized (Roelofs & Cardé 1971). Pheromone is synthesized in the fat body (Schal et al. 1998), trans- ported through the hemolymph to the pheromone glands, and then released as a liquid aerosol (Meyer 1984; Schal et al. 1987). Here, we provide a faunal treatment of the Virbia species occurring in North America north of Mexico, formerly placed in Holomelina Herrich-Schäffer. A generic diagnosis and description is provided for the ge-Figure 2. Virbia ostenta Edwards, Male. Figure 3. General range distribution of the genus Virbia. Triangles = species originally placed in Virbia; stars = species formerly placed in Holomelina. 62 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) nus. We recognize 14 species. One of these, V. marginata (Druce), is primarily distributed in Central America with disjunct populations in Colorado, New Mexico, and Arizona; it has been treated previously as a synonym of V. rubicundaria. Two new species, V. fergusoni Zaspel and V. rindgei Cardé, are described. For all species, we provide diagnoses, descriptions, and illustrations of diagnostic male and female morphology (Figs. 11-32). The adult habitus of both sexes are fig- ured (Figs. 27, Pls. 1-3), and we summarize life history information (e.g., larval hosts, voltinism, activity periods, and habitat). A database of over 12,000 specimens is available on-line through the Diversity of Life website (www.diversityoflife.org/virbia.html). BIOLOGY OF VIRBIA IN CANADA AND THE UNITED STATES Most of the information concerning the habitats and host plant use is derived from three sources: field notes of R. Cardé, unpublished sections of theses (Cardé 1971), and the Zaspel database of compiled label data. These data are supplemented with the few published rearing stud- ies (Dyar 1897; Forbes 1910, 1960; Wyatt 1939; Ferguson 1953). Data are incomplete for many species, but we note general trends when possible. Details of species life histories are provided in the Biology sections for each species. HABITAT Virbia exhibit a high degree of local habitat re- striction (Figs. 6 and 7), although most species have wide- spread distributions (e.g., V. aurantiaca, V. ferruginosa). Some phenotypically confusing species can be sorted simply by either habitat or locality infor- mation. For example, V. lamae Freeman has been col- lected only diurnally in bogs in the northeastern U.S. and Canada (Fig. 6) to the western edge of its range, Marathon County, Wisconsin (Ferguson 1954; Zaspel database). Virbia nigricans (Reakirt) has been col- lected only along a narrow corridor of secondary sucessional habitat comprised of old fields, power line cuts, and highway edges in western New Jersey (Hunterdon County) and extreme southeastern Pennsylavania (Philadelphia and Montgomery Counties; Cardé field notes, Zaspel database). Virbia nigricans has been confused historically with the dark form of V. opella, but these species are easily separated using adult habitus and locality. These species mate diurnally (Cardé 1971), and their genitalia are distinctive (see Diagnosis of V. nigricans). Another species with a highly special- ized distribution is V. rindgei Cardé (n. sp.), which is found in open Ponderosa pine forests in the Black Hills of South Dakota and Wyoming. The remaining species appear to be more general- ized in their habitat use, but their distributions are re- stricted. Virbia rubicundaria (Hübner) has been col- lected in tidal marshes on St. Catherine’s island off the coast of Georgia and also in moist, inland situations near the southeastern U.S. ocean shores such as the Florida scrub community in Highlands Co., Florida (Cardé field notes; Zaspel database). Additional species with some- what overlapping distributions can be separated based on habitat, wing pattern, or voltinism. For example, V. ferruginosa (Walker) is found in more open situations such as pitch pine - scrub oak barrens (Fig. 7), bogs, aspen parklands, and montane woods (Schmidt, personal communication), whereas V. opella appears to favor forested areas comprised of either deciduous woods or mixed hemlock and deciduous forest with a developed understory (Cardé 1965; Zaspel database). Virbia aurantiaca ranges from sand dunes and secondary suc- cession old fields (Massachusetts) to the New Jersey Pine Barrens, and from the Badlands (North Dakota) to alfalfa fields (Indiana) (Zaspel database). IMMATURE MORPHOLOGY AND BIOLOGY Most larval descriptions provide a brief account of the number of instars and general appearance. The first descriptions of immature stages were of V. aurantiaca (Saunders 1869) and V. opella (Dyar 1897). In 1960, Forbes described mature larvae for three species, V. laeta, V. aurantiaca, V. opella, and the dark form of V. opella that he referred to as nigricans. He noted that larvae of all species he examined possessed homoideous crochets, two pairs of transverse verrucae on the pro- thoracic shield, and abdominal L1 located below the spi- racle (Forbes 1960). Forbes (1960) described the larva of V. aurantiaca as dark, with black setae and verru- cae, and a contrasting conspicuous broad pale dorsal band filling the space between verrucae. All the setae were barbed and increased in length caudally. Larvae of V. laeta are lighter than V. aurantiaca, pale gray with black verrucae and a cream-colored stripe on the frons; two types of setae were present, dentate, and feathery (Forbes 1960). The third species, V. opella, was similar in body coloration to V. laeta (dull gray) but had a black head and brown frons; its black setae were described as short and stiff. Forbes (1960) described a larva that he attributed to form nigricans (1960: 23) whose body color was a light orange base heavily dusted with gray; the setae were gray, not black. This larva was likely the dark form of V. opella and not V. nigricans based on the locality information provided and the rarity of V. nigricans. Thus, two morphs of V. opella larvae ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 63 exist: a dull gray morph with black setae (typical V. opella) and a gray-orange morph with gray setae (dark form V. opella). Unlike the two previous species, V. ferruginosa larvae typically possess heteroideous crochers but have reduced lateral crochets (Schmidt, personal communication). Several species of Virbia have been reared on two introduced European plants, dande- lion (Asteraceae) or plantain (Plantaginaceae), but na- tive hosts are unknown (Forbes 1960; Tietz 1972). ADULT BIOLOGY Most of our detailed information on adult biology is limited to seven species representing the V. aurantiaca complex. This information is derived from the thesis work of Cardé (1971), supplemented by continuing field observations by Cardé. Roelofs & Cardé (1971) iso- lated and characterized female pheromone as 2- methylheptadecane (2Me-17c), the first saturated phero- mone from the Lepidoptera, for seven species in the V. aurantiaca complex: V. aurantiaca, V. ferruginosa, V. fragilis, V. immaculata, V. lamae, V. nigricans, and V. rindgei (as an undescribed species) (Roelofs & Cardé 1971). 2Me-17c was used in pheromone traps to study diel periodicities of attraction flight times. In addition, chromosome numbers were characterized for seven species. Methods for those studies are described in Cardé (1971). A summary of the data is published here for the first time (Table 1). COLLECTING ADULTS Some comments on methods for collecting adults in the V. aurantiaca complex illustrate how adults and particularly females (for initiation of laboratory colonies) were obtained. Nearly all of the males and females used in studies of the pheromone biology were collected by netting. All species can be collected by daytime flush- ing (e.g., walking through the habitat), even though spe- cies differ in whether females call and mate during the day or night. Afternoons are the most favorable time to collect; sunny and hot conditions are most conducive for flight. Flushed males fly in an erratic zigzag pattern, often for several meters or more, before landing; many males can be flushed a second or third time. Females have a reduced propensity to initiate flight, and the pro- portion of females flushed is low; this more sedentary behavior of females largely explains the great prepon- derance of males in collections. When females are flushed, their flight is readily distinguished: essentially a straight-line path with reduced velocity, usually lower altitude, and a much shorter distance than males, occa- sionally only decimeters. Upon landing in the herbage, females nearly always drop to ground level—males usu- ally land on the herbage—and “play dead” (thanotosis). These females can be difficult to locate unless their pre- cise trajectory has been tracked, and it is more efficient to net them on the wing if possible. Rarely, females can be flushed a second time. Daytime flushing of both sexes is presumably a defensive, predator avoidance reaction. Many species also are readily attracted to black light (males more strongly than females), and this collection method accounts for an abundance of museum speci- mens of V. aurantiaca, V. ferruginosa, V. immaculata, and V. rubicundaria. For day-mating species, V. lamae and V. nigricans, when black lights are placed in the precise locations where adults were collected diurnally, none are attracted to the lights. Oviposition behavior differs as well. Females of V. aurantiaca and V. rubicundaria lay eggs individually, not attached to a substrate; females of V. ferruginosa, V. fragilis, V. immaculata, V. lamae, V. nigricans, and V. rindgei lay attached, single-layer masses. PREMATING REPRODUCTIVE ISOLATION Among co-occurring species, isolation can be due to: 1) differences in pheromone, 2) rhythms of female release of pheromone, hereafter termed “calling,” and male response to pheromone, 3) exclusive seasonal flight periods, and 4) non-overlapping habitats. Figure 4. Virbia aurantiaca Hübner, Male. Figure 5. Virbia hypophaea Hampson, Male. 64 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 6. Virbia lamae, habitat, Seawall Bog, Southwest Harbor, Maine (Photograph by R.T. Cardé). Figure 7. Virbia ferruginosa, habitat, xeric outwash pitch pine - scrub oak barren in the Montague Plains Wildlife Management Area, Montague, MA. (Photograph by R.T. Cardé). ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 65 PHEROMONES Distinctive pheromone channels can be achieved in moths by a variety of strategies, including varying ra- tios of the same constituents, or altering chain length, double bond position, number and/or configuration, chirality, or functional group (Cardé & Haynes 2004). Indeed, it is typical to have closely related and sibling species differ in their pheromone blends, such that these serve as a primary means of premating reproductive isolation. The common pheromone constituent of the V. aurantiaca complex is 2-methylheptadecane (2Me-17c). Its presence in the female pheromone gland was con- firmed for V. aurantiaca, V. ferruginosa, V. fragilis, V. immaculata, V. lamae, V. nigricans, and V. rindgei (listed as an undescribed species) (Roelofs & Cardé 1971). The compound 2Me-17c evoked levels of at- traction in field trapping trials for all of the above spe- cies except V. fragilis, which was not tested. In addi- tion, V. rubicundaria and V. laeta are attracted to 2Me- 17c, although females have not yet been examined for the presence of 2Me-17c. In some species, such as V. aurantiaca and V. rubicundaria, the levels of male attraction relative to the density of field populations indi- cated that 2Me-17c serves as the primary component. Conversely, a comparatively poor level of attraction with 2Me-17c for V. lamae (Roelofs & Cardé 1971) sug- gests that additional pheromone components could be involved in communication. Indeed, in V. lamae, in addi- tion to 2Me-17c, several of its analogues are present in the gland and emitted, including n-heptadecane, n- nonadecane, 2-methyloctadane, and 2-methylnonadecane (Schal et al. 1987). Field tests of V. lamae with mix- tures of these constituents in naturally occurring ratios, however, do not have markedly improved attraction over 2Me-17c alone, leaving open the possibility that some species specificity awaits documentation of additional pheromone components. Differences in courtship be- haviors among the species, another possible factor in reproductive isolation, have not been examined. RHYTHYMS OF FEMALE CALLING Pheromone synthesized in oenocytes is transported via lipophorin to paired, tubular abdominal glands (Schal et al. 1998), each with a spine-beset opening at the ab- dominal tip (Yin et al. 1991). A calling female perches vertically and rhythmically protrudes the tip of her abdo- men at a rate of ca. once per 1.5 sec (Cardé 1971; Cardé & Roelofs 1973). Pheromone is atomized into droplets, in pulses coincident with each gland extrusion (Schal & Cardé 1985). This release mechanism is evi- dently common in arctiids but is not known among other moth lineages (Meyer 1984; Schal & Cardé 1986; Schal et al. 1987, 1998; review Hallberg & Poppy 2003). The amount of pheromone released is quite high among moths: in V. lamae up to 350 ng of 2Me17c in 10 min and 835 ng in 60 min (Schal et al. 1987). The calling rhythm of V. immaculata has been dem- onstrated to be circadian (Cardé & Roelofs 1973). Pre- sumably the rhythms of the remaining species in this complex also have a circadian basis; however, we refer to them here as diel rhythms, as their endogenous na- ture has not been confirmed formally. The calling rhythms illustrated (Figs. 8, 9), all were observed at a L:D of 16:8 at 24°C, but it is important to recognize that, like most calling and male response rhythms of moths, the expression of these rhythms is modulated by current and recent temperature regimes, with lower or decreas- ing temperatures generally advancing the rhythms of the night-calling species and delaying rhythms of the day- active species (Cardé & Roelofs 1973; Schal & Cardé 1986). Such temperature-induced modulation of calling periodicities, however, does not reduce their importance in isolation of some species pairs. FLIGHT PERIODS AND HABITAT PREFERENCES In addition to differences in diel rhythms and po- tential differences in pheromone blends, distinctive flight periods and habitat preferences isolate some co-occur- ring species. In Connecticut and Massachusetts, V. aurantiaca has two broods, with adults occurring mainly in secondary succession habitats (abandoned hay fields, power line cuts) from late May to late June and again in August. In the same areas, V. ferruginosa is single- brooded, with adult emergence in July in open scrub- oak forests. These habitat types can be contiguous, but the distinctive emergence patterns alone provide full iso- lation of the two species in southern New England. The female calling rhythms of both species from Connecti- cut overlap broadly (Fig. 8), with calling of V. aurantiaca peaking within the first hour of scotophase (dark seg- ment of the light-dark cycle) and lasting about 4 hours (Fig. 8a) and calling of V. ferruginosa commencing af- ter two hours of scotophase and also lasting for about 4 hours (Fig. 8d). Such differences in female calling rhythms would appear to provide only weak isolation for these two species, and, unlike most moths, the diel rhythm of male attraction in V. aurantiaca is considerably broader than the female’s nocturnal calling rhythm: male attraction to synthetic pheromone occurs from mid-morn- ing to evening (Cardé 1974). Although these temporal and habitat characteristics in concert provide effective isolation for V. aurantiaca and V. ferruginosa in south- 66 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 8. Calling periodicities of 2-day-old females in the Virbia aurantica complex. Observations were conducted every 30 minutes at 24° C in a 16:8 light:dark regime (see Cardé 1971 for methods). Time of scotophase indicated by the hatched bar. The sources for the lab colonies were: A-V. aurantiaca, East Haddam, CT.; B-V. immaculata, Dryden, NY; C-V. rubicundaria, Gainesville, FL; D-V. ferruginosa Branford, CT.; E-V. fragilis, Boulder, CO; and F-V. nigricans, Frenchtown, NJ. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 67 ern New England, both species are single-brooded in Maine (Brower 1974), Nova Scotia (Ferguson 1954), and generally throughout southern Canada, sharing the same flight period from July to mid August. Some par- titioning by habitat also seems important in Nova Scotia (Ferguson 1954). In southeastern Pennsylvania (Philadelphia Co.), A.M. Shapiro collected V. aurantiaca, V. nigricans, and V. immaculata in the same old field, secondary suc- cession habitat. Virbia nigricans is a day-calling spe- cies (Fig. 8f), whereas, as mentioned, V. aurantiaca calls in early evening (Cardé 1974), and V. immaculata (Cardé & Roelofs 1973) calls somewhat later in the evening (Fig. 8b). Such partitioning of calling times would afford some, but not complete isolation of V. nigricans from the other two species. The male response win- dow of V. aurantiaca includes attraction in daytime in southeastern Pennsylvania as it does in the Connecticut population (Cardé 1974). Similar levels of reproductive isolation by habitat preference and calling times should isolate V. lamae from V. aurantiaca and V. ferruginosa where they co-oc- cur. Virbia lamae is a day-calling species (Schal & Cardé 1986; Schal et al. 1987), and it is exclusively a denizen of bogs. Partial temporal (adult emergence, calling times) and spatial isolation occur for four species of the V. aurantiaca complex (Table 1). Virbia aurantiaca is bivoltine with adults emerging from May to June and again in August in southern New England and the mid- Altlantic United States (Cardé 1971); adults of V. aurantiaca emerge year-round in the south (Forbes 1960). Virbia immaculata is multivoltine in the north- central and northeastern United States, with adults emerging from late May and June and again in August and early September (Zaspel database). Two species, V. ferruginosa and V. lamae, are univoltine with a July flight period (Forbes 1960, Cardé 1971). Virbia rubicundaria and V. nigricans are multivoltine, with adults emerging year-round in the southeastern United States (Zaspel database). For those species that occasionally overlap in emer- gence and locality, female calling times may be impor- V. aurantiaca n = 25 yes yes 10am-10pm bi/multivoltine V. ferruginosa n = 32 yes yes night univoltine V. fragilis n = 25 yes yes — — V. immaculata n = 21 yes yes night multivoltine V. lamae n = 28 yes weak day univoltine V. nigricans n = 16 yes very weak day multivoltine V. rubicundaria n = 23 yes yes night multivoltine V. rindgei — — weak day — Species Chromosome # Female Male Male comes Voltinism calls attracted to lure Table 1. Biological information for nine Virbia species (Cardé 1968 & 1971). n = chromosome number. — = information not available. Figure 9. Calling periodicity of 2-day-old Virbia lamae females from Manistique, MI conducted hourly at 24° C in a 16:8 light:dark regime (see Schal and Cardé 1986 for methods). Time of scotophase indicated by the hatched bar. 68 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) tant in species discrimination. Three species are active and mate diurnally; four are active nocturnally (Table 1). Males of V. aurantiaca have the broadest known male activity period with male attraction to female phero- mone from midmorning (10:00) to early evening (20:00) (Cardé 1974) (Table 1). Virbia rubicundaria females call nocturnally (Cardé 1971). As yet unidentified com- ponents of females’ pheromone may confer some speci- ficity in the males’ response, resulting in diminished at- traction of heterospecifics (Carde 1971). Even if occasional mating of heterospecifics oc- curs, the range of chromosome numbers among the spe- cies suggests that most hybrid combinations would be at a disadvantage (Table 1). Haploid numbers range from 16 (V. nigricans) to 32 (V. ferruginosa) (Table 1, Cardé 1971). TAXONOMIC HISTORY OF HOLOMELINA AND VIRBIA Here we review the taxonomic history of Virbia, in- cluding Holomelina. We also provide a taxonomic re- view of species, but limit ourselves to those that occur in North America north of Mexico, all of which formally were placed in Holomelina. We first review the exten- sive taxonomic history of Holomelina, and then review the literature for Virbia. Last, we present a brief sum- mary of the recent phylogenetic study and its taxonomic implications (Zaspel & Weller 2006). Table 2. Taxonomic History of Virbia. B & B = Barnes & Benjamin, B & MD = Barnes & McDunnough, G-M = Guérin-Méneville, N& D = Neumögen & Dyar. Family Genus Species Synonyms Genus Lithosiidae Lithosia laeta G-M treatii Grote Crocota rubropicta Packard Lithosia Lithosina opella Grote flava B & B Crocota obscura Stretch Holomelina belmaria Ehrmann Holomelina rubricosta Ehrmann Holomelina Geometridae Crocota costata Stretch. opelloides Graef Crocota intermedia Graef Crocota parvula N. & D. Crocota cocciniceps Schaus Eubaphe pallipennis B & MD Eubaphe ferruginosa Walker quinaria Grote Crocota choriona Reakirt Holomelina trimaculosa Reakirt Crocota bucholzi Wyatt Holomelina ostenta Edwards calera Barnes Holomelina immaculata Reakirt nigricans Reakirt nigrifera Walker Cothocida rubicundaria Hübner belfragei Stretch Crocota diminutiva Graef Crocota rosa French Crocota Geometridae Eubaphe fragilis Strecker aurantiaca Hübner bimaculata Saunders Crocota brevicornis Walker Crocota lamae Freeman marginata Druce ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 69 TAXONOMIC REVIEW OF HOLOMELINA The taxonomic history of Holomelina Herrich- Schäffer is intertwined with two other genera, Eubaphe (Hübner 1823) and Crocota (Hübner 1823). These two genera originally were described among a series of lithosiine genera, and subsequent authors inferred that Hübner considered these genera as Arctiidae (Watson et al. 1980). Eubaphe and Crocota are now placed in Geometridae (Fletcher 1954; Watson et al. 1980), but the associated geometrid and arctiid species were treated as Eubaphe for nearly 120 years (Beutenmüller 1898; Druce 1885; Schaus 1901; Forbes 1910; Barnes & McDunnough 1917, 1918; Freeman 1941; Ferguson 1954). Similarly, arctiid and geometrid species were described in Crocota through the 1800s (Walker 1854; Grote 1863; Graef 1887; Smith 1889) or treated as such in checklists (Neumögen & Dyar 1893a). When Hübner described Eubaphe, he listed a single species, E. lobula (Hübner 1823), and thus E. lobula is the type species by monotypy (Fletcher 1954). Hübner subsequently described Eubaphe aurantiaca (Hübner 1827); Herrich-Schäffer used E. aurantiaca as the type spe- cies for Holomelina (Herrich-Schäffer 1855). Unlike Eubaphe and Crocota, Holomelina has always been placed in Arctiidae (e.g., Holland 1903). Hampson (1920: 316) asserted that Eubaphe had priority over Holomelina, and most taxonomists followed Hampson until Fletcher (1954) revised Eubaphe. In his revision, Fletcher (1954) noted that the type species, E. lobula, was a geometrid, and “the next valid name to replace Eubaphe in the Arctiinae appears to be Holomelina Herrich-Schäffer 1855, the type species of which is Eubaphe aurantiaca Hübner” (Fletcher 1954: 155). However, as late as 1960, some authors considered Eubaphe to be the valid generic name for these arctiid species (e.g., Forbes 1960: 21). The confusion of three generic names, Eubaphe, Crocota, and Holomelina, has resulted in congeneric species and subspecies being described in different genera (Table 2). Although Holomelina species (regardless of ge- neric name) have been treated as Arctiidae, the three genera, Holomelina, Eubaphe, and Crocota have been placed in various subfamilies, including Lithosiinae (Walker 1854), Arctiinae (Neumögen & Dyar 1893a,b), and Eubaphianae (Smith 1889). Earlier authors associ- ated Eubaphe and Crocota with Lithosiinae, listing these genera in sequence with lithosiine genera Cisthene and Delphyre in checklists (Stretch 1885), species’ descrip- tions (Walker 1854), and faunal treatments (Seitz 1914). Eubaphe (= Holomelina) was placed in Arctiini (Forbes 1960), and Holomelina subsequently has remained in Arctiini (Ferguson 1985; Franclemont 1983; Watson & Goodger 1986; Jacobson & Weller 2002). Discrimination of species was contentious in the 1800s (Smith 1889), and species taxonomy was further complicated with individual species and subspecies de- scribed in lithosiine genera (Lithosia author, Brycea Walker, Boenasa Walker, Ptychoglene Felder), arctiine genera (Cothocida Walker, Cytorus Grote), or geometrid genera (Crocota Walker, Eubaphe Walker) (Table 2). In some cases, subspecies were described in different genera than the original species (e.g. Lithosia laeta Guérin-Méneville and current syonoym Crocota treatii Grote; Table 2). Descriptions were often cursory and so general, that they applied to several species without distinguishing among them, for example, Crocota costata Stretch (Stretch 1885). In phenotypically variable spe- cies, such as V. costata and V. ferruginosa, a prolifera- tion of names resulted (Table 2). Complicating matters further, some authors refused to acknowledge the work of others. In his Preliminary Catalogue of the Arctiidae of Temperate North America with notes, Smith (1889) states: “The matter has been further confused by Mr. Grote’s persistent re- fusal to refer to Mr. Reakirt’s species.” Even when authors discussed others’ work, agreement on species names could not be reached, resulting in the application of several names to the same species (see Smith 1889). Depending on the author treating the Nearctic fauna, the number of valid species ranged from five (Walker 1854) to 17 (Smith 1889), and the number of varieties ranged from none (Walker 1854) to seven (Neumögen & Dyar 1893a,b). In the early 1900s, the species’ numbers were re- vised down to either six species (one variety) (Holland 1903) or eight species for the U.S. and Canadian fauna (Barnes & McDunnough 1917). Holland commented that “great confusion” existed concerning the number of species, and that larval rearing and mating studies were needed to resolve problematic species (Holland 1903: 115). Contemporary taxonomic treatments that included species occurring in Central and South America recognized 28 Eubaphe species and six varieties (Strand 1919) or 26 species, nine subspecies, and three varieties (Seitz 1914). Little additional taxonomic activity occurred until the 1950s and 1960s. Ferguson (1954) reviewed the Canadian fauna and renewed interest in the genus. Ferguson (1954) noted that species in Eubaphe had been difficult for taxonomists for some time; he stated, “The old question of which is a species and which is not, still remains paramount” (Ferguson 1954: 212). In treating the E. aurantiaca species complex, Ferguson discov- ered that differences in male genitalia between Eubaphe 70 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 10. Strict consensus of 779 trees: L = 1698, CI = 41, RI = 65, resulting from the analysis of all taxa AT matrix. Numbers below the branch are decays, and unlabelled branches have a decay of 3. C1 – C6 = Clades 1 through 6. Parallel lines = species formerly placed in Holomelina, Solid lines = species originally placed in Virbia, dotted line = outgroups. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 71 species were scarce, and offspring reared in the labora- tory were phenotypically diverse, i.e., they did not match the parents. (Ferguson 1954: 212). Extensive reared series at the National Museum of Natural History (NMNH) document the array of phenotypes. Additional reared series are housed at the American Museum of Natural History (AMNH, Joseph Muller), Cornell Uni- versity (CUIC, J. Franclemont material), and the Uni- versity of California, Riverside (UCR, R. Cardé mate- rial). Forbes (1960) provided a synopsis of the known biology of the genus and commented on the misuse of the generic name Eubaphe. He defended its usage as providing taxonomic stability in general usage (footnote 3a, Forbes 1960: 21). Forbes provided illustrations of diagnostic male genitalic features and a short key to the H. opella and the H. aurantiaca complex consisting of H. aurantiaca, H. laeta, and H. lamae. Other names were subsumed as varieties. In 1965, Cardé published redescriptions of H. aurantica, H. ferruginosa, and H. lamae and provided a partial key to the Nearctic Holomelina species. Cardé (1965) commented on the sibling species in the H. aurantiaca complex, listing H. aurantica, H. ferruginosa, and H. lamae as distinct species. In this paper, Cardé provided commentary on the H. aurantiaca and H. opella complexes. He was the first to realize that H. nigricans was not a synonym of H. opella based on adult habitus and male genitalia (Cardé 1965). Cardé summarized additional preliminary find- ings stating, “Areas of uncertainity are currently under investigation.” In 1968 and 1971, Cardé completed a set of taxonomic, chromosome, behavior, and pheromone studies on Holomelina for his M.S. and Ph.D. at Cornell University. His taxonomic work is not available under the International Code of Zoological Nomenclature (ICZN 1999), because unpublished theses do not con- stitute valid publications. He was unsuccessful in locat- ing Hübner types for H. aurantiaca and H. rubicundaria (Cardé 1968, 1971), which complicated his attempts to determine their species’ limits. Cardé’s work included two new species that remained as manu- script names (Cardé 1968) until now. Although Cardé’s taxonomic revisions were not published, Franclemont (1983) followed his treatment of Holomelina in the Checklist of the Moths of America north of Mexico (Hodges et al. 1983). TAXONOMIC REVIEW OF VIBRIA Virbia (Walker 1854) was described in Arctiidae and associated with lithosiine moths. Its familial place- ment has never been controversial (Nye et al. 1980; Ferguson 1985). It is a typical member of the Arctiini except some species possess homoideous crochets (Forbes 1939, 1960; Ferguson 1985; Kitching & Rawlins 1999; Jacobson & Weller 2002). Virbia was erected by Walker (1854), and he described two species, V. mentiens and V. luteilinea. Kirby (1892) subsequently designated V. mentiens as the type species. Until recently (Zaspel 2004; Zaspel & Weller 2006), the only generic synonyms were Bepara Walker (described in Melameridae) and Terna Walker (described in Callimorphidae). As of 1986, 31 of the 39 species placed in Virbia were described in Virbia (Watson & Goodger 1986). The remaining species were described in Bepara Walker, Brycea Walker, Cothocida Walker, Josioides Felder, Phaeochlaena Hübner, Ptychoglene Felder, or Terna Walker. Again, the small size of the moths led some authors to place these species in lithosiine genera (e.g., Lithosia Fabricius, Josioides Felder). A few species were described in other families: Geometridae (Crocota Hübner, Eubaphe Hübner), Dioptidae (Phaeoclaena Hübner), and Noctuidae (Phalaena L.). In contrast to the turbulent taxonomic history of Holomelina, Virbia has received little revisionary attention. Specific no- menclature has been comparatively stable; there were only four synonyms and two subspecies recognized in the most recent checklist of neotropical species (Watson & Goodger 1986). PHYLOGENETIC PLACEMENT OF VIRBIA AND HOLOMELINA There has been a long taxonomic history of asso- ciating these genera. Herrich-Schäffer (1855) was the first to do so when he described Holomelina. He in- cluded Phalaena medarda Stoll (Fig. 11) as one of two species comprising Holomelina. Later authors did not follow him and placed P. medarda in Virbia. Seitz (1914: 294) noted that Virbia species were structurally much like Eubaphe but differed by the branching of the up- permost median vein of the forewing. For over 90 years, checklists and faunal treatments have associated Eubaphe and Virbia (e.g., Seitz 1914; Forbes 1939; Hampson 1920). Forbes (1939: 220) remarked that Eubaphe and Virbia were closely related and very un- like other arctiine genera. Ferguson (1985) listed char- acters uniting Virbia and Holomelina in the Holomelina generic group, and noted that they “might not be dis- tinct” (Ferguson 1985: 199), in other words, congeneric. Recent checklists (Hodges et al. 1983; Watson & Goodger 1986) treated Virbia and Holomelina as mem- bers of the Arctiini. Phylogenetic studies have come to the same conclusion (Kitching & Rawlins 1999; Jacobson & Weller 2002). A phylogenetic analysis based on adult morphol- 72 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) ogy was conducted to determine whether Holomelina and Virbia were monophyletic with regards to each other (Zaspel 2004; Zaspel & Weller 2006). Those studies included 46 ingroup taxa (of 70 possible) and six arctiine species as outgroups. All Holomelina species, and about half the species of Virbia were included. Fifty-eight characters (195 states) were described and scored from non-genitalic and genitalic structures (Zaspel 2004; Zaspel & Weller 2006). The maximum parsimony analysis yielded 779 trees with a length (L) of 1698, a consistency index (CI) of 41 and a retention index (RI) of 65 (Zaspel & Weller 2006; Fig. 10). The monophyly of the clade comprised of Holomelina and Virbia species was well supported. The clade had a decay index greater than 10 (Bremer 1992) and was supported by five unreversed synapomorphies: 1) M2 missing in the hindwing, 2) pres- ence of a scaphium, 3) presence of a subscaphium, 4) apical hooks on the uncus, and 5) juxta ornamentation. The North American species with the Holomelina phe- notype clustered into two clades, the H. aurantiaca clade and the H. opella clade. The H. opella clade was sister to the clade consisting of Virbia, the yellow and black phenotype (Fig. 10). Multiple trees resulted from conflicting resolutions of the apical clade of neotropical species, consisting of those species traditionally assigned to Virbia. Different analyses could not recover a recip- rocally monophyletic Virbia and Holomelina (Zaspel 2004; Zaspel & Weller 2006). That is, the H. opella and H. aurantiaca clades were never grouped exclu- sive of Virbia species. Thus, the former Holomelina was an artificial assemblage of two different dispersal events from Mexico and the southwestern U.S. into Canada and the central and eastern U.S. (Zaspel 2004; Zaspel & Weller 2006). Holomelina (Herrich-Schäffer 1855) was placed as a junior synonym of Virbia (Walker 1854) (Zaspel 2004; Zaspel & Weller 2006) to reflect the results of the study. No subgenera were recognized because they would be either paraphyletic or typically treated as Holomelina (e.g., V. opella group). SUMMARY The taxonomic history of Holomelina is compli- cated by the confusion with geometrid generic names and the frequent acrimonious debates over species’ lim- its among 19th century American lepidopterists. These debates were fueled by the phenotypic diversity of the moths. As noted in the Biology Section, a single species encompasses adult morphs found in monomorphic, co- occurring species. Virbia, in comparison, has had a rela- tively quiet taxonomic history. The long-standing, phyletic association of Virbia and Holomelina was confirmed (Zaspel & Weller 2006), and in retrospect, the result was not surprising. The red and yellow hindwing coloration and species’ distributions were the principal characters formally used to distin- guish the genera. The phylogenetic study that supports this synonymy is discussed at length in Zaspel and Weller (2006). Below we provide an abbreviated checklist for the species. A checklist for the entire genus can be found in Zaspel and Weller (2006). CHECKLIST OF VIRBIA FOR NORTH AMERICA In this checklist, species are arranged alphabetically. Valid species are in bold with synonyms in italics. The genus in which a species was originally described fol- lows the author name in parentheses, and an asterisk indicates that the type was examined. When the type is not from the United States, the country is noted. Genus Virbia Walker 1854 Bepara Walker,1865a Cathocida Hampson, 1901, misspelling Cothocida Walker, 1865b Crocata authors, misspelling Crocota authors, not Hübner, 1823 Cytorus Grote, 1866 Eubaphe authors, not Hübner, 1823 Holomelina Herrich-Schäffer, 1855 Figure 11. Male genital capsule, Virbia medarda. 8S = eighth sternite, Co = coremata (highly developed), J = juxta, V = valve (sacculus region). ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 73 Terna Walker, 1865a V. aurantiaca (Hübner, 1827) Eubaphe bimaculata (Saunders, 1869) Arctia brevicornis* (Walker, 1854) Crocota V. costata (Stretch, 1885) Crocota opelloides (Graef, 1887) Crocota intermedia* (Graef, 1887) Crocota parvula* (Neumögen & Dyar, 1893b) Crocota cocciniceps (Schaus, 1901) Eubaphe pallipennis* (Barnes & McDonnough, 1918) Eubaphe V. fergusoni Zaspel n.sp. V. ferruginosa* (Walker, 1854) Crocota quinaria (Grote, 1863) Crocota choriona (Reakirt, 1864) Crocota trimaculosa (Reakirt, 1864) Crocota buchholzi* (Wyatt, 1963a,b) Holomelina V. fragilis* (Strecker, 1878) Crocota V. immaculata* (Reakirt, 1864) Crocota V. laeta (Guérin-Méneville, 1832) Lithosia treatii (Grote, 1865) Crocota rubropicta (Packard, 1887) Lithosia V. lamae (Freeman, 1941) Eubaphe V. marginata* (Druce, 1885) Eubaphe (Guatemala) V. nigricans* (Reakirt, 1864) Crocota nigrifera* (Walker, 1865b) Cothocida V. opella (Grote, 1863) Crocota flava (Barnes & Benjamen, 1925) Eubaphe obscura (Stretch, 1885) Crocota belmaria (Ehrmann, 1895) Crocota rubricosta (Ehrmann, 1895) Crocota V. ostenta* (Edwards, 1881) Crocota calera* (Barnes, 1907) Holomelina V. rindgei Cardé n. sp. V. rubicundaria (Hübner, 1827) Crocota belfragei (Stretch, 1885) Crocota diminutiva (Graef, 1887) Crocota rosa (French, 1890) Crocota MATERIAL AND METHODS SPECIMENS AND PREPARATION Dissection methodology follows Winter (2000) and is fully described in Zaspel and Weller (2006). Varia- tions on standard techniques include using only cold 10% KOH for 18-24 hours; most wings were not cleared and slide-mounted. This was done to ensure the small, fragile structures were not ‘over cleared’ prior to dis- section. These moths have very fragile wings; there- fore, 1-2 drops of 30% ethanol were added to the wings of pinned specimens for observation of venation. Larger, more robust wings placed briefly in weak bleach (30 seconds), and then standard procedures were followed Figure 13. Virbia laeta, male, A- Forewing, R2 = R2 + R3 fused or R3 absent. B- Hindwing, M2 absent, Sc + R1 = subcosta fused with radial vein 1, Rs = radial sector vein, other abbreviations as in Fig. 12. Figure 12. Virbia aurantiaca, male forewing, all R veins present. C = costa, Sc = Subcosta, R1-5 = radial veins 1 – 5, M1- 3 = medial veins 1 –3, CuA1 - CuA2 = cubital anal veins 1 and 2, A = anal veins. 74 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) for staining with eosin Y (1% in distilled water; Fisher Scientific, Pittsburgh, PA) and slide mounting. All prepa- rations went through a dehydration series before mount- ing (Winter 2000). Balsam preparations went through two additional dehydration treatments of oil of clove (15 minutes), and then xylene (15 minutes) before slide mounting. Euparol mounts were transferred from the dehydration series into a final 15-minute treatment in Euparol essence (Bioquip, Garden City, CA) before slide mounting. Permanent slide mounts (Canada balsam [Sigma, St Louis, MO] or Euparol [Bioquip, Garden City, CA]) were made of abdominal pelts, genitalia, legs, wings, labial palps and antennae. Slides were placed on trays and cured in drying ovens for 24 – 48 hours. Terms for wing venation follow Scoble (1992). Terms for ab- dominal and genitalic morphology follow Klots (1970), Weller et al. (2000), Jacobson and Weller (2002) and Kristensen (2003). For females, when two bursae exist (double or connate condition), we could not determine which one was the appendix busa based on position or Figure 16. A. Virbia opella, male. An = antenna simple and ciliate.; B. Virbia laeta, male. An = antenna emar- ginate, asymmetrical flagellomeres. Figure 14. Virbia cyana, male forewing, R2 and R3 ab- sent, R4 + R5 = fused other abbreviations as in Fig. 12. Figure 15. A. Condition of the tarsal claw, Virbia aurantiaca, simple; B. Condition of the tarsal claw, Virbia cyana, slightly bifid. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 75 presence of a signum. Therefore, we made no distinc- tion between corpus and appendix in the descriptions. MATERIAL EXAMINED For the study of species limits, 180 specimens were dissected, representing the 14 species described below; dates from specimen labels have been standardized. In addition, a specimen-level database was compiled in Microsoft Excel that included locality data (label data), prior identifications, comments, and repository for speci- mens of Virbia species. The database includes over 12,000 entries from 16 collections (listed below) follow- ing the protocol of the NMNH (M. Pogue, personal com- munication). It is searchable and can be found of the Diversity of Life website at www.diversityoflife.org/ virbia.html. The following is a list of the institutional and pri- vate collections consulted during this study. The acro- nym of the institution or name of private collection is followed by the name of the individual that prepared the loan. Acronyms follow Heppner and Lamas (1982): AMNH - American Museum of Natural History, New York (E. Quinter); CMNH - Carnegie Museum of Natural History, Pennsylvania (J. Rawlins); CUIC -Cornell Uni- versity Insect Collection, New York (J. Liebherr); CSU - Colorado State University, Colorado (P. Opler); E. H. Strickland Entomological Museum, University of Alberta, Alberta (F. Sperling); EMEC - Essig Museum of Ento- mology, University of California, Berkeley, California (J. Powell); FMNH - Field Museum of Natural History, Il- linois (P. Goldstein); LACM - Los Angeles County Mu- seum, California (J. Donahue); MSU - Mississippi State Univeristy, Mississippi Entomological Museum, Missis- sippi (R. Brown); UNSM - University of Nebraska State Museum, Nebraska (B. Ratcliffe); NMNH - National Museum of Natural History, Washington D.C. (M. Pogue); TAMU -Texas A and M University Insect Col- lection, Texas (E. Riley); UCONN - University of Con- necticut Insect Collection; Connecticut (D. Wagner); UMSP - University of Minnesota Insect Collection, Min- nesota (P. Claussen); RTC - Personal collection of R. Cardé, University of California-Riverside. SPECIAL TERMS Adbasal-This is the space before the basal bands in most other noctoids. In arctiids it can be present, but is usually lost, or fragmented and united with the antemedial band. Basal-This represents the space between the basal and antemedial band. Antemedial-This can be complete or fragmented. Be- cause of this fragmentation, this band is almost always a composite of the basal and antemedial bands, and it is uncertain how often a true antemedial band occurs. Medial-A band near the middle of the wing between the antemedial and postmedial areas, sometimes incomplete and apparent as a spot or bar near the costa. Postmedial-This is the pale space beyond the postme- dial line. This is often present as a full band. Subterminal-This is a zigzag band running from a point on the costa about midway between the postmedial band and apex toward the tornus-the anal region or anal angle, merging with the postmedial band between M3 and Cu1, then bending sharply outward again. This band may be complete, fragmentary or absent. SPECIES DESCRIPTIONS Species were delimited primarily by differences in the male genitalia, especially characteristics of the vesica, phallus, and juxta. Female genitalia are diagnostic for some. Due to a redundant distribution of character states between species (e.g., little interspecific variation), di- agnostic morphological characters typically are illustrated once and are referred to in multiple descriptions. Some Virbia species are almost indistinguishable by their geni- talic characteristics; in addition, a variety of wing color patterns occur, rendering species determination difficult. Within the North American fauna, species range from pale pink and yellow (e.g., V. ferruginosa and V. immaculata) to deep red and brown (e.g., V. ostenta). Some species (e.g., V. aurantiaca) have wing patterns that vary within long reared series from a single female (Fig. 1). Although variable, these species do share com- mon pattern elements with other arctiines. All speci- mens were examined under a true color lamp (OTT- LITE®), and colors are described using the Naturalist’s Color Guide (Smithe 1975). In general, arctiids have greatly modified wing pat- tern systems compared to their closest relatives, the Noctuidae (Ferguson 1985, sensu Fibiger & Lafontaine 2005, not Lafontaine & Fibiger 2006). The forewing pattern typically consists of light stripes and spots against a darker background, a reverse of the classic noctuid pattern. The forewing pattern usually is based on six lightly colored, transverse bands on a dark background, these are almost always reduced to five or fewer, or lost, with many variations occurring within a genus (Ferguson 1985). Ferguson notes: Two typical pattern systems occur on the forewings of many Arctiini: a lon- 76 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) gitudinal vein pattern and a supplemen- tary longitudinal pattern consisting of paracostal, cubital, and postcubital band, also called the primary and secondary longitudinal patterns, respectively (Ferguson 1985:183). Similarly, the basic hindwing pattern of Arctiini is reversed from the noctuid one, and consists of a colored background with either dark spots or dark outer-mar- ginal borders (Ferguson 1985). A discal spot or spots may be present, but often are modified or lost (Ferguson 1985). Nomenclature for pattern elements of Virbia fol- lows Ferguson (1985). Descriptions of wing regions, patterns, and colors specific to Virbia species are de- fined in the “Specialized Terms” section. The described pattern elements are strictly for species’ identification purposes and are not intended to be interpreted as ho- mologies of pattern elements across Noctuoidea. To facilitate the construction of consistent diagnoses and descriptions, DELTA (Descriptive Language for Taxonomy) was used (Dallwitz et al. 1993). Species descriptions were first constructed using DELTA, and then minor formatting changes were made. SYSTEMATIC ENTOMOLOGY Order LEPIDOPTERA Family ARCTIIDAE Subfamily ARCTIINAE Tribe ARCTIINI Genus Virbia Walker Virbia Walker, 1854: p. 470. Type species: Virbia mentiens Walker, 1854. Bepara Walker, 1865a . Cathocida Hampson, 1901, misspelling. Cothocida Walker, 1865b. Crocata authors, misspelling. Crocota authors, not Hübner, 1823. Cytorus Grote, 1866. Eubaphe authors, not Hübner, 1823. Holomelina Herrich-Schäffer, 1855. Terna Walker, 1865a. Diagnosis (Figs. 1,2,4,5, Pl. 1-3).—- These are relatively small arctiids, with forewing length ranging from 8.2 mm to 17.1 mm in males (N = 14) and from 8.7 mm to 18.5 mm females (N = 11). Antennae are typically simple and ciliate (Fig. 16A). Forewing with two prodominant pattern types: dark forewings and hindwings with bright yellow markings (Fig. 5) or yellowish areas replaced with pink or reddish markings (e.g., V. ostenta) (Fig. 4). Male genitalia are unique in Arctiini in the pos- session of a true scaphium (except for V. cyana) (Figs. 25A, B). The corpus bursa can be single, double, or connate with the corpus (Fig. 17C). Virbia can be sepa- rated from other Arctiidae by the loss or reduction of the M2 in the hindwing (Fig. 13B) and the scaphium. Description.—- Adult habitus (Figs. 1,2,4,5, Pl. 1-3): Head color uniform, but variable, from dark black (Fig. 2) or brown (Pl. 3, Nos. 2-3) to pinkish orange (Fig. 4). Third segment of labial palpus typically long, exceeding front by at least half its length; color variable from yellow to pink, gray, or varying intensities of brown. Ocelli absent. Thorax with dorsal and ventral scale col- oration variable, from dark black (Fig. 5) or brown (Pl. 3, Nos. 2-3) to pinkish orange (Fig. 4). Forewing typi- cally with R1 - R5 all present (e.g., V. aurantiaca: Fig. 12), R3 absent or trace in V. laeta (Fig. 13A), and R2, R3 absent or trace in V. cyana (Fig. 14). Hindwing with M2 absent (Fig. 13B). Wing color pattern one of two main types (see diagnosis). Abdomen with color vari- able from dark black (Fig. 5) or brown (Pl. 3, No. 2) to pinkish orange (Pl. 2, No. 4). Sclerotization of male 8th sternite often species specific, shapes including: trian- gular (e.g., V. phalangia: Fig. 28A), oval (e.g., V. opella: Fig. 28B), pentagonal (e.g., V. marginata: Fig. 28C), pentagonal with expanded base (e.g., V. divisa: Fig. 28D), antecosta reinforced by transverse ridge, rectan- gular width greater than length (e.g., V. luteilinea: Fig. 28E), and M-shaped (e.g., V. metazonata: Fig. 28F). Coremata typically present, but variable developmed. Male genitalia: Phallus typically with coecum penis rounded, occasionally weakly notched (e.g., V. ferruginosa: Fig. 22A). Vesica directed ventrally in most species, directed posteriorly in V. latus, V. cyana, and V. polyphron. Phallic sclerite typically present, but variable: narrow, rounded triangular (e.g., V. ostenta: Fig. 22B), wide and nodular (e.g., V. aurantiaca: Fig. 22C), triangular, blunt (e.g., V. inversia: Fig. 18A), sharp and triangular (e.g., V. endophaea: Fig. 18B), and square (e.g., V. polyphron: Fig. 18C). Vesica typically rugose with endophallus ornamentation on right side, under phal- lic sclerite present, either as ventral cornutuslike patch (e.g., V. aurantiaca: Fig. 22C) or peglike patches (e.g., V. endophaea: Fig. 18B). Genital capsule (Figs. 25A, 25B) dorsad with tegumen either M-shaped or concave (e.g., V. costata: Fig. 23A), with lobes (e.g., V. aurantiaca: Fig. 23B), or entire (e.g., V. laeta: Fig. 23C). Ventrad with uncus base typically cordate (e.g., V. aurantiaca: Fig. 23B). Ventrally directed hooks present on apex of uncus with bases fused or separate; usually two hooks present (e.g., V. costata: Fig. 23D), but only one in V. semirosea and lacking in V. cyana, V. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 77 phalangia, and V. polyphron. Scaphium present (ex- cept V. cyana), conical (e.g., V. laeta: Fig. 23E), or flat- tened (e.g., V. phalangia: Fig. 23F). Costa typically elongate with apex entire (e.g., V. laeta: Fig. 25A), some- times forked (e.g., V. opella: Fig. 27D). Saccus typi- cally rounded, setose (e.g., V. marginata: Fig. 27C) in- frequently thin, square (e.g., V. feronia: Fig. 27E). Fe- male genitalia: Papillae anales convex, rounded; pseudopapillae anales typically present (e.g., V. nigricans: Fig. 24B), infrequently lacking. Posterior apo- physes entire, long and slender. 8th tergite unornamented. Anterior apophyses entire, long and slender. Eighth ster- nite round, wide and entire. Location of ostium bursa just above 8th sternite, ornamentation of ventral edge variable: central broad indentation with small lateral prominences (e.g., V. aurantiaca: Fig. 17A), straight (e.g., V. lamae: Fig. 17B), split in center with lateral promi- nences (e.g., V. opella: Fig 30A), small lateral promi- nences (e.g., V. rubicundaria: Fig. 30B), or toothed (e.g., V. satara: Fig. 30C). Shape of ductus bursa variable: rectangular (e.g., V. aurantiaca: Fig. 17A), hourglass- shaped (e.g., V. lamae: Fig. 17B), goblet-shaped (e.g., V. opella: Fig. 17C), or rectangular, tapering in middle of base (e.g., V. metazonata: Fig. 17D). Sclerotization of ductus bursa extending into corpus bursa, except in V. marginata. Corpus bursa with (e.g., V. opella: Fig. 17C) or without additional connate bursa (e.g., V. aurantiaca, V. lamae: Fig. 17A & B), typically with one to three round signa. Discussion.— Species with the Holomelina pat- tern are often polymorphic, and several phenotypes may occur in a single brood (Fig. 1). Rearing studies are needed of species exhibiting the yellow-black Virbia phenotype to determine whether polymorphism is preva- lent. In Mexico, species with Virbia and Holomelina patterns co-occur. Both male and female genitalia are useful for dis- tinguishing phenotypically confusing species. The lobing and ornamentation of the vesica (endophallus) is often species’ specific. In several species, females exhibit species’ specific ornamentation of the ostium bursa or corpus bursa, and several species have pseudopapillae anales (Fig. 24B). These moths often are found misplaced in uncurated Geometridae, Notodontidae (Josia), or lithosiine arctiids. They can be separated from Geometridae by the placement of the tympana on the metathorax (versus on the base of the abdomen in Geometridae). They can be separated from Josia based on forewing venation (M2 arising from mid discal cell in Josia and from near M3 in Virbia). The hindwing vena- tion (M2 absent in Virbia) usually will separate these species from other arctiids. Virbia aurantiaca Hübner Pl. 1 No. 5 (female), Nos. 3-4 (males) Eubaphe aurantiaca Hübner 1827: 206 [type lo- cality: Pennsylvania, USA]. Crocota bimaculata Saunders 1869: 5 [type lo- cality: London, Ontario, Canada]. Crocota brevicornis Walker 1854: 2 [type local- ity: United States]. Crocota choriona Reakirt 1864: 371 [type local- ity: Philadelphia, Pennsylvania, USA]. Holotype.—- The holotype of Eubaphe aurantiaca is lost according to Lloyd Martin, Curator of Lepidoptera at Naturhistorisches Museum, Wien (per- sonal communication 6/25/2004). A neotype is desig- nated here and labeled to ensure nomenclatural stability. The specimen was chosen from the nearest locality to the original type locality. The neotype, male [CMNH] is labeled: Virbia aurantiaca (Hbn. 1827) [RED LABEL]. Label data: At light New Lisbon, N.J. Aug. 7 1934, E.P. Darlington//Exch. A.N.S.P. GM. Acc. 20359. Material examined.— Type material: Neotype des- ignated above. All material is from the NMNH unless otherwise noted: CANADA: Nova Scotia: Lequille, 1.viii.1946, D. C. Ferguson (1 Male, JMZ100). Que- bec: Temiscouta Co., 30.vi.1985, leg. George Hensel (1 Male, JMZ94). MEXICO: Cordoba: 16.v.1908, Fred Knab (1 Female, JMZ92). Jalapa: Wm. Schaus Col- lection (1 Male, JMZ93). UNITED STATES: Arkan- sas: Washington Co., Springdale Blue Spruce Camp- ground, 28.v.1966, R. Heitzman, (1 Male, JMZ96). Connecticut: New Haven, Hamden, 3.ix.1967, D. C. Ferguson (1 Male, JMZ119). Kansas: Labette Co., Oswego, 3.vi.1966, Gary F. Hevel collector, at lights (1 Male, JMZ115). Kentucky: Oldham Co., Horner Bird Sanctuary 25.viii.1977, C. V. Covell Jr. (1 Male, JMZ113). Maine: Southwest Harbor, 21.vii.1936, near Colorado, marginata as in Barnes Collection 1937 (1 Male, JMZ183); Willimantic, 15.vii.1930, H. Wilhelm (1 Male, JMZ101). Maryland: Queen Annes Co., Gransonville Horsehead Sanctuary, 24.vii.1986, D. C. Ferguson (1 Female, JMZ109). Michigan: July Barnes Collection (1 Male, JMZ112). Montana: Sweetgrass Co., Big Tim- ber Creek 7 mi. N. Big Timber, 29.vii.1966, D. C. Ferguson (1 Male, JMZ99). New York: Long Island, Greenport, viii (1 Male, JMZ103). New Jersey: Or- ange Mts., 24.vii.1937, Otto Bucholz (1 Female, 1 Male, JMZ110; JMZ111). North Dakota: Slope Co., Burn- ing Coal Veins Pine Ridge, elev. 2500', T 136N, R 102W, Section 11, 21.vii.1982, Eric H. Metzler (2 Males, JMZ116; JMZ118). Oklahoma: Sequoia Co., Tenkiller 78 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 17. A. Bursa complex, Virbia aurantiaca, ventral view. B = bursa (membrane wrinkled), Db = ductus bursa (rectangular and narrowing toward ventor), Ob = ostium bursa (central, broad indentation and small lateral promi- nences); B. Bursa complex, Virbia lamae, ventral view. Db = ductus bursa (hourglass), Ob = ostium bursa (straight), S = signum; C. Bursa complex, Virbia opella, dorsal view. B = bursa (connate and arising from ductus, membrane of additional lobe wrinkled), Db = ductus bursa (goblet, sclerotization extending from ductus bursa into corpus bursa); D. Bursa complex, Virbia metazonata, dorsal view. B = bursa (connate, arising from ductus; membrane of additional lobe smooth), Db = ductus bursa (square). ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 79 Lake, 3 mi. W. Blackgum, 6-9.vii. 1979, D. & M. Davis (1 Male, JMZ117). South Carolina: McClellanville, Wedge Plantation, 22.iv.1974, D. C. Ferguson (1 Male, JMZ102). Tennessee: Nashville, 16-23.vi., Barnes Collection (1 Male, JMZ95). Texas: Concan, Garner Ste. Pk. 1.vi.1964, A. & M. E. Blanchard (1 Male, JMZ62), Hill Co., Lake Whitney Park, 21.iv.1985 (1 Male, JMZ98); Sinton, Welder Wildlife Foundation, 22.iv.1964, A. & M. E. Blanchard (1 Male, JMZ67); Cameron Co., Brownsville, 17.xi.1941, Collection of Fred Rindge, AMNH (1 Male, JMZ288); Santa Ana Wildlife Refuge, 21.x.1970, A. & M. E. Blanchard (2 Females, JMZ194; JMZ196). Virginia: Leesburg, 4.vii.1970, D. C. Ferguson, reared ex ovo on dandelion (1 Male, JMZ108). Wisconsin: Lindsey, 23.vii.1964, J. W. Barnes (1 Male, JMZ97). Wyoming: New Castle, 6 mi. N.W., 28.vi.1965, R. W. Hodges (1 Male, JMZ114). Diagnosis. — Virbia aurantiaca may be confused with V. ferruginosa, V. lamae, V. marginata, V. nigricans, and V. rubicundaria, but it is most com- monly confused with V. ferruginosa. The two species are separated using hindwing maculation and potentially emergence dates. The subterminal hindwing band of V. aurantiaca is solid, whereas the subterminal area of V. ferruginosa has a series of blotches and spots, not a solid band. In New York and neighboring states, V. ferruginosa emerges in July, whereas broods of V. aurantiaca emerge in June and August (Cardé1965, 1974). Flight period is less reliable in the Maritime Prov- inces, northern Michigan, and northern Maine, where there is a single brood and the adult emergence of V. aurantiaca overlaps with V. ferruginosa (Zaspel data- base). In the southern states (e.g., Georgia, Florida), V. aurantiaca may be encountered anytime in the sum- mer (Zaspel database, Adams personal communication). Virbia aurantiaca can be separated from V. nigricans and V. lamae if detailed label data are avail- able. Virbia lamae is a bog endemic and males fly (with- out flushing) diurnally. Virbia nigricans has been col- lected only in a narrow corridor of disturbed secondary forest on the border of western New Jersey (Hunterdon County) and eastern Pennsylvania (Philadelphia area). This species is much darker and smaller (wing length 8.2 mm) (Figs. Plate 2, Nos. 8,9) than V. aurantiaca (wing length 9.7 mm) (Pl. 1, Nos. 3-5). The shape of the juxta also separates the three species. In V. aurantiaca, the juxta is ovoid with the ventral margin expanding into a larger, orbicular shape (Fig. 20B), whereas in V. lamae and V. nigricans, it is cylindrical and emarginated as in V. marginata (Fig. 27C). In fe- males, V. aurantiaca has pseudopapillae anales; the other two do not. Lateral prominences surround the ostium bursa in V. aurantiaca (Fig. 26D) and V. nigricans but not V. lamae (Fig. 26B). Virbia fragilis, a western species, occasionally is misidentified as V. aurantiaca, especially specimens from the western edge of V. aurantiaca’s range. Virbia fragilis is a much larger species (14 mm), and the hindwing color is flesh ocher and lacks subterminal mark- ings. The phallic sclerite is wide and nodular in V. aurantiaca (Fig. 22C), whereas it is highly reduced in V. fragilis. In Central America, V. aurantiaca co-occurs with V. marginata (range: Guatemala, Mexico, SW USA). Diagnostic differences are provided under V. marginata. Occasionally, pale specimens of V. aurantiaca lack the subterminal hindwing band, and these can be con- fused with the boreal V. immaculata or V. rubicundaria, but V. immaculata is a larger species (forewing length 12 mm) and has a reduced phallic sclerite compared to V. aurantiaca (Fig. 22C), and V. rubicundaria is a smaller species (forewing length 8.9 mm). Females of V. aurantiaca can be confused with those of V. rubicundaria, but V. rubicundaria is restricted to the southeastern USA and is rarely sympatric with V. aurantica. Virbia rubicundaria females are also much smaller (forewing length 8.7 mm vs. 9.7 mm, respec- tively) and brighter than V. aurantiaca females (com- pare Pl. 2. No. 4, Pl. 1, No. 5). Description.—- Male habitus (Pl. 1, Nos. 3-4): Head dark clay throughout. Antennal scales dark clay. Third segment of labial palpus exceeding front of head by at least half of entire length, first and second seg- ments salmon, apex of third segment grayish horn. Tho- rax dorsal and ventral scales clay throughout. Legs clay, tarsal claw simple (Fig. 15A). Forewing length 10 mm (N = 10); R1-R5 all present, m3 present, extending to wing margin (Fig. 12). Dorsal surface ranging from clay to cinnamon throughout; ventral surface salmon. Hindwing dorsal surface peach red, typically with solid raw umber subterminal band, ventral surface salmon. Abdomen with dorsal and lateral abdominal scales clay to salmon, dorsal tergites occasionally with medium size fawn spot in center, size of spot tapering with each seg- ment towards posterior, ventral abdominal scales yellow ocher throughout. Sclerotization of 8th sternite penta- gon (Fig. 28C). Coremata reduced to small patches of scales on membranous lobes. Female habitus (Pl. 1, No. 5): Head clay. Antennal scales clay. Third seg- ment of labial palpus exceeding front of head by at least half of entire length, all segments clay. Thorax dorsal, lateral, and ventral scales salmon to clay. Legs clay with patches of salmon scales on femora and tibiae, tar- sal claw simple (Fig. 15A). Forewing length 9.7 mm (N 80 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 18. A. Phallus, Virbia inversia left, lateral view. Dps = phallic sclerite (triangular, blunt), Ve = vesica (upper medial-proximal projection on left side), Cp = patch of cornuti (on venter); B. Phallus, Virbia endophaea left, lateral view. Dps = phallic sclerite (triangular, sharp), PCrn = peglike patches of cornuti (under phallic sclerite), Pl = basiphallus (single sclerotized lobe-like projection at midpoint), Ve = vesica; C. Phallus, Virbia polyphron left, lateral view. Dps = phallic sclerite (square, rugose), R = rugose patch (under phallic sclerite), Ve = vesica (directed laterally), Ve1 = right dorsal lobe, Ve2 = left lateral lobe; D. Phallus, Virbia costata left, lateral view. CoP = coecum penis (rounded and orientation straight), Dps = dorsal phallic sclerite (narrow and rounded), Pl = basiphallus (inflected ventrally at base), Ve = directed (ventrally, membrane rugose, left side with membranous rugose projection). ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 81 = 10), venation same as in male. Forewing dorsal sur- face salmon to cinnamon with faint olive brown discal spot present, ventral surface salmon, peach red scales lining costal margin. Hindwing dorsal surface peach red with natal brown discal spot, natal brown subtermi- nal markings as spots and blotches often not reaching posterior margin, ventral surface identical. Abdomen with dorsum intermixed with salmon and clay, occasion- ally with fuscous spots on all segments, lateral scales salmon with fuscous spots on all segments, ventrally in- termixed with salmon and clay. Male genitalia: Phallus (Fig. 22C) with rounded coecum penis, straight, basiphallus inflected ventrally at midpoint, phallic scler- ite wide, nodular. Vesica directed ventrally, membrane rugose, bilobed, elongate, second lobe sclerotized, lower left side of vesica with small membranous projection, right side of vesica with patch of cornuti under phallic sclerite. Genital capsule (Fig. 25A) with dorsum of tegumen lobed, sides of tegumen divergent, conjunctiva membranous (Fig. 23B). Uncus base cordate (Fig. 23B), ventrad with uncus apex fingerlike, bifid and with two fused hooks (Fig. 27A). Scaphium rounded, triangular. Subscaphium absent. Extensions of tegumen sclero- tized. Juxta vertically ovoid with ventral margin expand- ing into larger orbicular plate (Fig. 20B). Costa of valva elongate and tip entire, 20 or more costal setae. Saccu- lus horizontally obovate, broadly hooked toward costa. Saccus rounded, concave in center. Female genitalia (Figs. 26C & D, 17A): Pseudopapillae anales present. Pheromone glands membranous, branching before tip of anterior apophyses, moderately thick, greater than 1/ 8 but less than 1/4 width of 7th abdominal sternite. Os- tium bursa slightly concave with small lateral promi- nences (Fig. 30). Ductus bursa rectangular, narrowing toward venter, sclerotization extending into corpus bursa (Fig. 17A). Corpus bursa with three signa, membrane of the corpus bursa wrinkled (Fig. 26C). Variation. —- The most common phenotype of V. aurantiaca is the ‘red form’ (Pl. 1, Nos. 3-5); however, a yellow phenotype resembling V. marginata (Pl. 3, No. 1) also occurs infrequently. Nearly all specimens of V. aurantiaca we examined have a characterstic round, un-notched coecum penis and a dorsally rounded tegumen. Rarely, red form individuals have a notched coecum penis as in V. marginata or V. ferruginosa (Fig. 22A). Some males from the South, specifically Texas, have darker raw umber forewings with a white annellar spot and a smaller forewing length than specimens from the northeast. Biology.—- This species can be reared on dande- lion and plantain (Forbes 1960, Cardé 1968), but its na- tive host is unknown. Larvae feed readily on artificial diet (Zaspel database). The chromosome number ap- pears to be 25 (Cardé 1971). In southern Connecticut, two broods can be reared, with one emerging in June (peak emergence around June 20) and the other in Au- gust (Cardé 1971). Additional generations can be reared in the southern U.S, and the species is considered facul- tatively multivoltine (Forbes 1960). Females lay indi- vidual eggs unattached, like V. rubicundaria (Cardé, rearing notes). Females have been observed releasing pheromone nocturnally (Fig. 8), but males are attracted to pheromone from 10:00 to 22:00 hours (Cardé 1974). Distribution.—- This species is among the most widely distributed in the V. aurantiaca complex. In east- ern North America, specimens have been collected from Manitoba and Nova Scotia, Canada, along the eastern seaboard, south to Cordoba, Mexico (see database for records from these localities). The western edge of its distribution is less clear. The species has been collected in Texas, Mississippi, Missouri, Tennessee, Louisiana, Oklahoma, Kansas, and North and South Dakota. We examined a single specimen from Montana that either represents a highly disjunct population or is misidentified. Virbia costata Stretch Pl. 1, No. 10 (female), Pl. 2, No. 1 (male) Crocota costata Stretch 1885: 103 [type locality: Texas, USA]. Crocota opelloides Graef 1887: 42 [type locality: Texas, USA]. Crocota intermedia Graef 1887: 42 [type locality: Texas, USA]. Crocota parvula Neumögen & Dyar 1893b: 140 [type locality: western Colorado, USA]. Eubaphe cocciniceps Schaus 1901: 269 [type lo- cality: “USA”]. Eubaphe pallipennis Barnes & McDonnough 1918: (1) 85, pl. 14, Fig. 14 [type locality: “USA”]. Material examined.—- Type material: Holotype of V. costata not examined. Holotype C. intermedia: Crocota intermedia Texas, E.L. Graef Collector, 41D1/ /Collection Brkyln. Mus.//Type No. 34162 U.S.N.M// Crocota intermedia Graef, type Tex. Coll. Edw. L. Graef, NMNH. Holotype of C. parvula: Crocota intermedia parvula Colo. Bruce//Col. B. Neumögen// Typicum Specimen//Type No. 33695 U.S.N.M//41D2// Crocota intermedia v. parvula Type Female Neumögen, NMNH. Holotype E. pallipennis: E. costata v. pallipennis Type B & McD.//Glenwood Spgs., Col. July 16-23//47 B3 [sic.], NMNH. All mate- rial deposited in NMNH unless otherwise noted: UNITED STATES: Arizona: Yavapai Co., Mayer, 23.vii.1959, R. F. Sternitzky (1 Female, JZ220). Colo- 82 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 19. A. Male genital capsule Virbia fergusoni. S = saccus (rounded, concave in center), Sa = sacculus (peanut shaped), SbSc = subscaphium (completely surrounding anal tube); B. Phallus, Virbia fergusoni, left, lateral view. CoP = coecum penis (rounded), Cp = cornutal patch, D = dorsal, Dps = dorsal phallic sclerite (reduced), Pl = basiphallus (inflected ventrally at base), Ve = vesica, (bilobed, directed ventrally with rugose membrane, right side with patch of cornuti under phallic sclerite), V = ventral; C. Male genital capsule Virbia opella. SbSc = subscaphium (lobed and incomplete, e.g., not completely surrounding anal tube), S = saccus (rounded, concave midventrally); D. Phallus, Virbia opella, left, lateral view. D = dorsal, Dps = phallic sclerite (reduced), Ve = vesica (directed ventrally, membrane rugose, bilobed, elongate, second lobe rugose), LCn = left side of vesica (with ventral patch of cornuti), RCn = right side of vesica (with patch of cornuti under phallic sclerite), V = ventral. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 83 rado: Glenwood Springs, 16-23.vii., compared with type collection in Brooklyn Museum (1 Female, JZ60); 24- 30.vii., E. costata, pallipennis paratype (1 Male, JZ65); La Plata Co., Durango, Elev. 6, 500 ft., 2.vii.1957, F. & P. Rindge, AMNH (1 Male, JZ273). Oklahoma: Comanche Co., Wichita Mts. Nat. W. Refuge, The Nar- rows, 10-18.vii.1984, D. & M. Davis (1 Male, JZ88). Texas: Alpine, 8-14.viii.1928, Poling collector, Barnes Collection (1 Male, JZ89); Brewster Co., Big Bend Nat’l Park, Green Gulch, 6.v.1972, A. & M. E. Blanchard, AMNH (1 Male, JZ256). New Mexico: Eddy Co., Big Canyon, Guadalupe Mts., 24.vii.1959, A. B. Klots, Holomelina intermedia i. intermedia det. by Graef, AMNH (1 Female, JZ257). Diagnosis.—- This species is sexually dimorphic (Plate 1, No. 10; Plate 2, No. 1). Females superficially resemble V. laeta females and males, but V. costata is restricted to southwestern U.S., whereas V. laeta oc- curs in the north central and southern U.S. Virbia costata females are larger (wing length 13.5 mm) than males of V. laeta (wing length 11 mm), and smaller than females of V. laeta (wing length 17 mm). Males of V. costata can be confused with males of V. fragilis, a western species, but the two can be separated by male genitalic characters. The genitalia of V. costata have a large subscaphium and a phallic scler- ite that is narrow and rounded (Fig. 18D), whereas V. fragilis lacks a subscaphium and has a highly reduced phallic sclerite (not shown). Description. Male habitus (Pl. 2, No. 1): Head flesh ocher throughout. Antennal scales tawny olive. Third segment of labial palpus short; first and second segments flesh ocher, third segment dark tawny olive. Thorax dorsal and ventral scales tawny olive throughout. Legs clay, tarsal claw simple (Fig. 15A). Forewing length 11.7 mm (N = 10), R3 absent, M3 present, extending to Figure 20. A. Juxta, Virbia opella: vertically ovoid with ventral margin (VM) expanding into large orbicular shape, dorsal margin (DM) concave with pointed edges; B. Juxta, Virbia (H.) aurantiaca: vertically ovoid, concave on both sides with ventral margin expanding into large orbicular shape; C. Juxta, Virbia ferruginosa: with trans- parent patches (P) present, (broadly rounded). Figure 21. Corpus bursa, Virbia ferruginosa, dorsal view. B = bursa (connate and both bursae arising from ductus bursa), Db = ductus bursa, L = lobe of second bursa, S = signum (three signa present). 84 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) wing margin (Fig. 13), dorsal surface pale pinkish buff suffused with light salmon, ventral surface clay dusted with orange rufous. Hindwing dorsal surface flesh ocher with edges fringed with pale pinkish buff scales, ventral surface yellow ocher dusted with flesh ocher. Abdo- men with dorsal, lateral, and ventral scales flesh ocher. Sclerotization of 8th sternite rectangular, length greater than width (Fig. 28E). Coremata reduced to small patches of scales on membranous lobes. Female habi- tus (Pl. 1, No. 10): Head clay, occiput ranging from salmon to peach red among specimens. Antennal scales clay. Third segment of labial palpus exceeding front of head by at least half of entire length, first and second segments salmon, third segment clay. Thorax dorsal and ventral scales clay with tufts of salmon on ventral surface near wing base. Legs tawny olive with light salmon scales on femora and tibiae, tarsal claw simple (Fig. 15A). Forewing length 13.5 mm (N = 10), vena- tion as in male. Forewing dorsal surface solid clay to solid olive brown with salmon along costal margin, ta- pering toward wing tip, ventral side salmon to peach red with pattern beginning at proximal portion of adbasal region, expanding transversely to most distal portion of postmedial region, pattern meeting inner margin, inner margin with clay to fuscous scales extending from post- medial region to wing apex, occasionally slightly sinu- ate. Hindwing dorsal surface salmon, pattern beginning at proximal portion of adbasal region, pattern expanding transversely to most distal portion of postmedial region, extending to margin of clay to fuscous scales, inner margin extending from postmedial region to wing apex, occasionally slightly sinuate, in some specimens this post- medial region is with a thin, broken subterminal line, ven- tral surface identical. Abdomen with dorsal scales fuscous, lateral scales on first three segments salmon, lateral abdominal scales tapering toward posterior in fuscous scales, ventral side fuscous throughout. Male genitalia: Phallus (Fig. 18D) with rounded coecum pe- nis, straight; basiphallus inflected ventrally at base, curved toward venter, phallic sclerite narrow and rounded. Vesica directed ventrally, membrane rugose, bilobed, elongate, second lobe membranous, left side of vesica with a rugose, lobed projection and cornutuslike patch located ventrally, right side of vesica with rugose patches under phallic sclerite. Genital capsule dorsally with tegumen m-shaped, conjunctiva weakly sclerotized (Fig. 23A). Uncus base cordate (Fig. 23B). Uncus apex triangular and with two distinct hooks. Scaphium rounded, triangular. Subscaphium completely surround- ing anal tube (Fig. 25A), extensions of tegumen sclero- tized. Juxta ovoid with ventral margin expanding into larger orbicular shape, dorsal margin concave with pointed edges (Fig. 20A). Costa of valva elongate and tip entire, 20 or more costal setae. Sacculus clawlike, extending under juxta on both sides. Saccus rounded, concave in center. Female genitalia: Opening of ovi- positor glandular, lacking pseudopapillae anales. Phero- mone glands membranous and unbranched, moderately wide, greater than 1/8 but less than 1/4 width of 7th ab- dominal sternite. Ostium bursa slightly concave with small lateral prominences (Fig. 30B). Ductus bursa rect- angular, narrowing toward venter, sclerotization extend- ing into corpus bursa (Figs. 17A, 26C). Corpus bursa double, connate, with three signa, both bursae round with wrinkled membrane (Fig. 26C). Variation.— There is almost no variation in size or color among males or females. Biology. Virbia costata has been reared on plan- tain (Zaspel Database), but its native host is not known. A single flight period occurs in July in Arizona, New Mexico, Colorado and Oklahoma. Two flight periods occur in the Big Bend region (Texas): the first in June and the second in early August (Zaspel database). Distribution. — This species occurs in the western United States; the eastern edge of its range is western Oklahoma; the northernmost record is from Colorado. Virbia fergusoni Zaspel n. sp Pl. 3 Nos. 4 (female), 5 (male) Holotype.—- GA: Clinch Co., 7 mi. NE Fargo, 13.vii.1981, D. C. Ferguson//reared on Taraxacum// Holomelina MEM n. p. det. D. Ferguson//JMZ298, (Male), NMNH. Paratypes.—- UNITED STATES: Alabama: Baldwin Co., Bon Secour, NWR 30° 15’ 46’’N, 87° 45’ 27’’W, 12.v.1994, R. E. Seymour//in oak-pine forest// William Cross Expedition//Holo. n. sp. det. D. C. Ferguson, MESSU:MEM, (1 Female, JMZ299). Florida: St. Petersburg, Fla.//Barnes Collection, NMNH, (1 Fe- male, 2 Males); Fla. Collection Wm. Schaus, NMNH, (1 Male); Putnam Co., Univ. of Florida Preserve, Welaka, 22.iii.1987//D.C. Ferguson Collector, NMNH, (1 Male); Cassia Lake Co. Fla., 8.v.1964 Taylor//Collection of Bryant Mather//16305//genitalic preparation 1865-112 Ring Cardé, NMNH (1 Male); Highlands Co., Highlands Hammock S. P., 12.vi.1987, T. L. Schiefer//Holomelina n. sp. det. D. C. Ferguson 1994, MESSU:MEM, (1 Fe- male, JMZ312); Highlands Co., Highlands Hammock S. P., 12.vi.1987, T. L. Schiefer, MESSU:MEM, (1 Male); Highlands Co., Highlands Hammock S. P., 12.6.1987, T. L. Schiefer, MESSU:MEM, (1 Male); 15 mi. W. of Co- coa Beach, 19.iii.1966, R. H. Leuschner// LACM, (1 Male, JMZ313); 15 mi. W. of Cocoa Beach, 22.iii.1966, R. H. Leuschner//Holomelina wyatti Cardé det. Ring ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 85 T. Cardé, LACM, (1 Male); 15 mi. W. of Cocoa Beach, 31.iii.1966, R. H. Leuschner//Holomelina wyatti Cardé det. Ring T. Cardé, LACM, (1 Male); 15 mi. W. of Co- coa Beach, 22.iii.1966, R. H. Leuschner//LACM, (1Male, JMZ214); 25 mi. W. of Cocoa Beach, 19.iii.1966, R. H. Leuschner//LACM, (1 Male, JMZ213); 15 mi. W. of Cocoa Beach, 19.iii.1966, R. H. Leuschner//Holomelina wyatti Cardé det. Ring T. Cardé, LACM (1 Male); 25 mi. W. of Cocoa Beach, 6.v.1966, R. H. Leuschner, LACM, (1 Male); Alachua Co., Gainesville, 15.v.1993, Jeff Slotten Collector, LACM (1 Male, JMZ215); Alachua Co., Gainesville, 7.v.1993, Jeff Slotten Collec- tor, LACM, (1 Male); Alachua Co., Gainesville, 15.ix.1995, Jeff Slotten Collector, LACM (1 Male); Alachua Co., Gainesville, 5.iv.1993, Jeff Slotten Collec- tor, 1 Male (LACM); Alachua Co., Gainesville, 3.iv.1993, Jeff Slotten Collector, LACM (1 Male); Alachua Co., Gainesville, 15.v.1993, Jeff Slotten Collector, LACM (1 Male); Alachua Co., Gainesville, 3.v.1993, Jeff Slotten Collector, LACM (1 Male); Alachua Co., Gainesville, 5.iv.1993, Jeff Slotten Collector, LACM (1 Male); Florida, Orange Co. Tosohatchee St. Preserve (off Rt. 520 Tay- lor CrRd), 12.v.2001, Jeff Slotten Collector; Deland, Fla. 16.iii.1953, H. Ramstadt, CNHM 1957 A.K. Wyatt Lepi- doptera Colln.//male genitalia on slide #938 M.O. Glenn 12.xi.1962, FMNH (1 Male); Lakeland Fla. 21.vii.1941, McElhose Chicago N.H. Mus. (Al McElhose Collec- tion)//male genitalia on slide #53 M.O. Glenn 24.v.1964. Georgia: Charlton Co., Okefenokee Natl. Wildlife Ref- uge, 19.iv.1991, D.C. Ferguson//Boardwalk Trail Chesser Island, NMNH (1 Female); Charlton Co., Okefenokee Natl. Wildlife Refuge, 1.x.1992, D.C. Ferguson//Board- walk Trail Chesser Island Homestead, NMNH (1 Male); Charlton Co., Okefenokee Refuge Hdqtrs., near Folkston 4.v.1981, D.C. Ferguson, NMNH (1 Male JMZ64); Charlton Co., Trader’s Hill, Folkston, 27.ix.1992, D.C. Ferguson, NMNH (1 Female, JMZ59); Clinch Co., 7 mi. NE Fargo, 7.v.1981, D.C. Ferguson//reared ex ovo on Taraxacum, NMNH (1 Male, JMZ210); Clinch Co., 7 mi. NE Fargo, 14.vii.1981, D. C. Ferguson//reared ex ovo on Taraxacum, NMNH (1 Male, JMZ211); Clinch Co., 7 mi. NE Fargo, 5.v.1981, D. C. Ferguson, NMNH (1 Male, JMZ212); Clinch Co., 7 mi. NE Fargo, 13.vii.1984, D.C. Ferguson, NMNH, (1 Male); Clinch Co., 7 mi. NE Fargo, 13.vii.1981, D. C. Ferguson//reared on Taraxacum//Holomelina MEM n. p. det. D. Ferguson 1994, MESSU:MEM (1 Male). Mississippi: Harrison Co., Long Beach 10.iv.1996, R. Kergosien, MESSU:MEM (1 Male). All material is from the NMNH collection unless otherwise noted: UNITED STATES: Florida: Putnam Co., Welaka, University Reserve, 17.iv.1973, D. C. Ferguson (1 Male, JMZ86). Geor- gia: Atlanta, 17.v.1946, P. W. Fattig (1 Male, JMZ127); Clinch Co., 7 mi. NE Fargo, 5.v.1981, D. C. Ferguson// parent of brood (1 Female, JMZ185), Clinch Co., 7 mi. NE Fargo, 13.vii.1981, D. C. Ferguson//reared ex ovo on Taraxacum (1 Male, JMZ298). South Carolina: McClellanville, Wedge Plantation, 24.iv.1974, D. C. Ferguson (1 Male, JMZ85). Etymology.—- This species is named in honor of Dr. Douglas Ferguson. His reared series have been in- valuable in establishing phenotypic diversity within and among species. Dr. Ferguson encouraged JMZ’s inter- est in the revision of Holomelina and Virbia. His assis- tance and informed discussion of these moths is grate- fully acknowledged. Diagnosis.—- The coloration is distinctive. This species is bright, geranium pink with varying amounts of dark brown scales on the hindwing adbasal and subter- minal areas (Pl. 3, Nos. 4,5). This species can be con- fused with lighter V. opella males, but V. fergusoni has much darker forewings and more pronounced contrast in the hindwings. Paler forms of V. fergusoni lack the discal spot in the hindwing that occurs in V. opella. Two male and one female genitalic character can be used to separate this species from V. opella. The vesica of V. fergusoni lacks a cornutuslike patch located ventrally on the left side of the vesica (Fig. 19B) that occurs in V. opella (Fig. 19D). The subscaphium is not lobed in V. fergusoni (Fig. 19A) and is lobed in V. opella (Fig. 19C). Females of V. fergusoni have a signum, while V. opella females lack signa (Fig. 17C). This species may be confused with the much smaller V. rubicundaria, but it can be separated from the latter as determined in the diagnosis of V. rubricundaria. Description.—- Male habitus (Plate 3, No. 5): Head raw umber, occiput peach red. Antennal scales grayish horn color. Third segment of labial palpus ex- ceeding front of head by at least half of entire length, first and second segments peach red, third segment raw umber. Thorax dorsal and ventral scales raw umber. Legs cinnamon, with a mixture of peach red and clay on femora and tibiae, tarsal claw slightly bifid (Fig. 15B). Forewing length 11 mm (N = 10), R3 absent (Fig. 13), M3 present, extending to wing margin. Dorsal surface raw umber with dark fuscous on crossveins of discal spot forming a small transverse band, ventral surface geranium pink, terminal margins a fringe of raw umber. Hindwing dorsal surface peach red, subterminal region raw umber, occasionally meeting postmedial region in sinuate pattern, most specimens with raw umber scales shading in region from subterminal band along fold from discal cell to 3A, fuscous discal spot present or absent, ventral surface identical. Abdomen with dorsum gera- 86 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) nium pink with patches of fuscous on most segments, lateral and ventral scales buff throughout. Sclerotiza- tion of 8th sternite rectangular, length greater than width (Fig. 28E). Coremata absent. Female habitus (Pl. 3, No. 4): Head cinnamon, occiput salmon. Antennal scales fuscous. Third segment of labial palpus short, color same as in male. Thorax dorsal and ventral scales cinnamon. Legs raw umber with patches of geranium pink on femora, tarsal claw slightly bifid (Fig. 15B). Forewing length 12 mm (N = 10), venation as in male. Forewing dorsal surface cinnamon, occasionally with salmon cos- tal margin, ventral surface peach red throughout. Hindwing dorsal surface peach red, terminal margins fringed with tawny scales, ventral surface identical. Abdomen with color same as in male. Male genitalia: Phallus (Fig. 19B) with straight rounded coecum penis, basiphallus inflected ventrally at base, curved toward venter, phallic sclerite reduced. Vesica directed ven- trally when everted, membrane rugose, bilobed, elongate, second lobe of vesica rugose, right side of vesica with patch of cornuti under phallic sclerite (Fig. 19B). Geni- tal capsule (Fig. 19A) dorsally with tegumen entire, con- junctiva sclerotized (Fig. 23C). Uncus base cordate (Fig. 23B), ventrad with uncus apex fingerlike, bifid, with two fused hooks (Fig. 27A). Scaphium conical, pointed. Subscaphium completely surrounding anal tube (Fig. 19A), extensions of tegumen sclerotized. Juxta ovoid with ventral margin expanding into larger orbicular shape, dorsal margin concave with pointed edges (Fig. 20A). Costa of valva elongate and tip forked; 20 or more cos- tal setae (Fig. 19A). Sacculus peanut-shaped (Fig. 19A). Saccus rounded, concave in center. Female genitalia: Pseudopapillae anales present (Fig. 24B). Pheromone glands membranous, branching after tip of anterior apo- physes, moderately thick, greater than 1/8 but less than 1/4 width of 7th abdominal sternite. Ostium bursa straight, without lateral prominences (Fig. 17B). Duc- tus bursa hour-glass shaped (Fig. 17B), sclerotization ex- tending form ductus bursa into corpus. Corpus bursa bilobed, typically with one signum, membrane smooth (Fig. 21). Variation.—- Some male specimens of this spe- cies are significantly lighter in color with a raw umber subterminus extending toward the wing base, forming a blotchy line where the vannus meets the remigium. The male genitalia of some individuals had costa apices that were entire (Fig. 19A) rather than bifed. No color varia- tion in females was observed. Biology.— Virbia fergusoni was reared on dendelion (Taraxacum spp.) by Ferguson. Its native host is not known. This species appears to have mul- tiple broods with adult flight periods from mid-March Figure 22. A. Phallus, Virbia ferruginosa, left, lateral view. CoP = coecum penis (slightly knotched, orienta- tion straight), Pl = basiphallus (inflected ventrally at mid- point), Dps = phallic sclerite (lateral view, wide and nodu- lar), Ve = vesica (directed ventrally, membrane bilobed, lobe membranous), Cp = patch of cornuti (under phallic sclerite); B. Phallus, Virbia ostenta left, lateral view. Dps = phallic sclerite (narrow and rounded), Ve = vesica (directed ventrally, membrane smooth, entire), Cp = patch of cornuti (located ventrally), Ve1 = right dorsal lobe, Ve2 = left lateral lobe; C. Phallus, Virbia aurantiaca left, lateral view. CoP = coecum penis (rounded, orientation straight), Pl = basiphallus (inflected ventrally at midpoint), Dps = phallic sclerite (wide and nodular), Ve = vesica (directed ventrally, with membra- nous projection = Pr), Cp = patch of cornuti (under phal- lic sclerite), Ve1 = right dorsal lobe, Ve2 = left lateral lobe. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 87 through mid-July. A single specimen was taken in Sep- tember (Georgia). One label indicates the habitat was a mixed oak-pine forest. Distribution. —- This species is restricted to the southeastern United States; the most northern specimen was collected in McClellanville, South Carolina, from Georgia and northern Florida west to Baldwin Co., Ala- bama. Virbia ferruginosa Walker Pl. 1, Nos. 6 (female), 7(male) Crocota ferruginosa Walker 1854: 535 [type lo- cality, St, Martin’s Falls, Albany River, Hudson’s Bay, Ontario, Canada]. Crocota quinaria Grote 1863:30 [type locality, “Canada”]. Crocota trimaculosa Reakirt 1864: 371 [type lo- cality, Philadelphia, Pennsylvania, USA]. Holomelina buchholzi Wyatt 1963: 100-102 [type locality, Suffolk, Virginia, USA]. Material examined. — Type material: Syntypes of V. ferruginosa: Hudson Bay, G. Barnston 11.1917// Arctiidae slide #907//photographed BM negative 43750, Hudson Bay, G. Barnston 11.1917, also “caterpillar with fine long brown hairs”//Arctiidae slide #43757//BM nega- tive 43757; Holomelina ferruginosa data: label 284 Hudson Bay G. Barnston 11.1917, “Crocota ferruginosa”, NHM. Holotype (synonym of V. ferruginosa): VA: Suffolk, 4.viii.1941, Otto Buchholz, C.N.H.M. Type Male Holomelina aurantiaca Wyatt, Buchholz Collection, ANMH. Paratypes of V. ferruginosa: VA: Suffolk, 19.vii.1941, Otto Bucholz, Reared Series//CNHM Paratype//Holomelina, AMNH (1 Female, 1 Male, JMZ279; JMZ278); Suffolk, Otto Bucholz, 24.vii.1941//CNHM Paratype//Holomelina, AMNH (1 Female, JMZ280). All material is from the NMNH collection unless otherwise noted: CANADA: British Columbia: Vancouver, 3.vii.1921, T. A. Moilliet/ /Blackmore Collection (1 Male, JMZ57). Quebec: Lac à la Tortue, 17.vii.1947, Eugene Munro (1 Male, JMZ90); Temiscouata Co., 30.vi.1985, leg. George Hensel (1 Male, JMZ94). UNITED STATES: Connecticut: New Haven Co., Guilford, 24.vii.1966, leg. Ring T. Cardé, RTC, (1 Male, JMZ315). Illinois: Edgebrook, 9.x.1947 A. K. Wyatt (1 Female, JMZ162); 11.x.1947, A. K. Wyatt, Bred (1 Male, JMZ161). Indiana: Tremont, 27.vii.1952, A. K. Wyatt, ex ova (1 Female, JMZ70); Tremont, 3.vii.1944, A. K. Wyatt (1 Female, JMZ316); 9.vii.1954, A. K. Wyatt//CNHM 1957 A. K. Wyatt Lepi- doptera Collection, FMNH (1 Male, JMZ314). Minne- sota: Olmstead Co. (1 Male, JMZ121). Virginia: Mont- gomery Co. (1 Female, JMZ195). Wisconsin: Oneida Co., Lake Katherine, 10.vii.1960, M. M. Bower (1 Male, JMZ91). Diagnosis.—- Virbia ferruginosa can be confused with V. aurantiaca, V. fragilis, and V. immaculata. It can be separated from V. aurantiaca based on colora- tion and size as described in the diagnosis for V. aurantiaca. This species usually can be separated from V. fragilis by locality. Virbia fragilis is a western spe- cies found in the Rocky Mountain region, whereas V. ferruginosa typically occurs in the boreal north and eastern U.S. deciduous forests. The following male genitalic features can also be used to distinguish the two species: males of V. ferruginosa have a bilobed vesica and a knotched coecum penis (Fig. 22A), whereas those of V. fragilis have a single-lobed vesica and a rounded coecum penis (Fig. 22B). Females of V. ferruginosa have a connate, double bursae (Fig. 17C), whereas those of V. fragilis lack an additional bursa (Fig. 17A). In the Eastern U.S., the geographic distribution of V. ferruginosa overlaps that of V. immaculata, but flight period is a reliable means of separating the two. The flight period of V. immaculata extends from late spring to early summer – but rarely July, and its second brood emerges in August; V. ferruginosa’s flight period is in July. Abdominal coloration often can be used to dis- criminate the two. The abdominal tergites of V. ferruginosa often have a central cinnamon brown spot, whereas V. immaculata typically does not. In V. ferruginosa males, the second lobe of the vesica bears a rugose or sclerotized patch (Fig. 22A) whereas in V. immaculata the second lobe is sclerotized and bears cornuti (Fig. 22C). Unlike V. ferruginosa, females of V. immaculata lack an additional bursa. Description. —-Male habitus (Pl. 1, No. 7): Head cinnamon brown. Antennal scales cinnamon brown. Third segment of labial palpus exceeding front of head by at least half of entire length, all segments salmon. Thorax dorsal and ventral scales cinnamon brown throughout. Legs clay throughout, tarsal claw simple (Fig. 15A). Forewing length 13.4 mm (N = 10), R1-R5 all present, M3 present, extending to wing margin (Fig. 12). Dorsal surface cinnamon brown, with a faint olive brown discal spot present, ventral surface salmon with peach red scales lining costal margin. Hindwing dorsal surface pinkish warm buff, cinnamon brown subtermi- nal markings as spots and blotches often not reaching posterior margin, exterior posterior margin occasionally fringed with cinnamon brown scales, discal spot cinna- mon brown, ventral surfaces salmon. Abdomen light buff, with first segment on dorsal side a tuft of cinna- mon brown, tergites often with a medium size cinnamon brown spot in center, size tapering with each segment 88 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) towards posterior. Sclerotization of 8th sternite pen- tagonal (Fig. 28C). Coremata reduced to small patches of scales on membranous lobes. Female habitus (Pl. 1, No. 6): Head clay. Antennal scales clay. Third seg- ment of labial palpus exceeding front of head by at least half of entire length, all segments clay. Thorax clay. Legs cinnamon brown with patches of scales on femora and tibiae from salmon to buff, tarsal claw simple (Fig. 15A). Forewing length 15 mm (N = 10), venation same as in male. Dorsal surface cinnamon extending from wingbase through postmedial region, subterminal region clay color, with a faint natal brown discal spot present, although rare, some specimens with small white spot or several large spots in annelar region, ventral surface dark salmon suffused with peach red scales. Hindwing dorsal surface peach red, with a natal brown discal spot, natal brown subterminal markings as spots and blotches often not reaching posterior margin, ventral suface iden- tical. Abdomen with dorsum intermixed salmon and clay, occasionally with fuscous spots on all segments, lateral sides salmon color with fuscous spots on all segments, ventral sides a blend of salmon and clay. Male genita- lia: Phallus (Fig. 22A) with coecum penis weakly notched, straight, basiphallus inflected ventrally at midpoint, curved toward venter, phallic wide, nodular. Vesica directed ventrally when everted, membrane rugose, bilobed, elon- gate, second lobe membranous; right side of vesica with patch of cornuti under phallic sclerite. Genital capsule with dorsum of tegumen lobed, sides of tegumen diver- gent (Fig. 23B), conjunctiva strongly sclerotized. Un- cus base cordate (Fig. 23B), ventrad with uncus apex fingerlike, bifid, with two fused hooks (Fig. 27A). Scaphium rounded, triangular. Subscaphium absent, extensions of tegumen sclerotized. Juxta rectangular, tapering in center, expanding toward ventral base with backward directed barbs on dorsal margin, broadly rounded ornamentation present (Fig. 20C). Costa of valva elongate and apex entire, 20 or more costal setae. Sacculus horizontally obovate, broadly hooked toward costa. Saccus rounded, concave in center. Female geni- talia (Fig. 21): Pseudopapillae anales present (Fig. 24B). Pheromone glands membranous, branching after tip of anterior apophyses, moderately thick, greater than 1/8 but less than 1/4 width of 7th abdominal sternite. Os- tium bursa slightly concave with small lateral promi- nences (Fig. 30B). Ductus bursa rectangular, narrow- ing toward venter, sclerotization extending into corpus bursa (Fig. 21). Corpus bursa double, connate, with one signum, both bursae round with wrinkled membrane (Fig. 21). Variation.—- In some males, the apex of the third labial palp segment is cinnamon brown or has a patch of cinnamon brown scales. In others, the cinnamon brown subterminal markings form a large longitudinal band ex- tending through the posterior margin. Some males from reared series were curated un- der the name V. buchholzi (AMNH and FMNH). These males have bright yellow ocher scales through- out. Cardé (1968) proposed that V. buchholzi was a color variant of V. ferruginosa. Franclemont (1983) placed it as a junior synonym of V. ferruginosa. We have examined several V. ‘bucholzi’ specimens and agree that it is a color variant of V. ferruginosa. Biology.—- Virbia ferruginosa is univoltine with adults emerging during July. Larvae have been reared on dandelion (Taraxacum spp.; Cardé, Rearing Notes). Females release pheromone (exhibit calling behavior) at night (Fig. 8D). Distribution.— This species occurs in Canada from Nova Scotia to British Columbia (Zaspel database). In the United States, it is sympatric with V. aurantiaca in the Northeast and upper Midwest. Its range extends south to Virginia, Mississippi, Missouri, and Lousiana. A few specimens superficially resembling V. ferruginosa have been collected in Colorado; however, it is likely that these specimens are V. fragilis. Virbia fragilis Strecker Pl. 2, Nos. 6 (female), 7 (male) Crocota fragilis Strecker 1878: app: 1859 [type locality, Pagosa Springs, Colorado, USA]. Material examined.—- Type material: Holotype of V. fragilis: Crocota fragilis Streck. Pagosa Springs Col. Orig. Type McCanley//Crocota fragilis Streck. Colorado, Pagosa Springs McCanley Orig. Type Strecker Colln. 27530 Field Museum Nat. Hist.//Holotype det. A. Watson 1986//Genitalic preparation 201 Ring Cardé Genitalic preparation 201 Ring Cardé//Lepidoptera type photograph No. 184 Field Museum, FMNH. All mate- rial is from the NMNH collection unless otherwise noted: CANADA: Alberta: Lloydminster, 5.vii.1942, P. F. Bruggeman (1 Male, JMZ55). British Columbia: Ar- rowhead Lake//Barnes Collection (1 Male, JMZ226). UNITED STATES: Colorado: Denver, Chimney Gulch, 16-23.ix (1 Female, JMZ73). New Mexico: Las Vegas (1 Male, JMZ56). South Dakota: Pennington Co., Spring Creek Camp, 11 mi. NE Hill City, 31.vii.1955, F. & P. Rindge, AMNH (1 Male, JMZ274); Slate Creek, Black Hills, 9 mi. NW of Hill City, 6000', 20.vii.1964, D. C. Ferguson (1 Female, JMZ72); 9 mi. NW of Hill City, 6000’ July 15, 1964 D. C. Ferguson (1 Male, JMZ193). Utah: Ephraim Sanpete Co., Major’s Flat, 10.vii.1981, D. C. Ferguson (1 Male, JMZ192). ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 89 Diagnosis. — Virbia fragilis is easily confused with V. immaculata, V. costata males, and it is some- times confused with V. ferruginosa. Characters sepa- rating V. fragilis and V. ferruginosa are given above. Virbia fragilis and V. immaculata are geo- graphically isolated from each other. Virbia fragilis oc- curs in western Canada and the western and southwest- ern U.S., whereas V. immaculata occurs in north cen- tral and eastern U.S. Males of V. fragilis tend to be larger (wing length 14 mm) than those of V. immaculata (wing length 12 mm). Male genitalia further distinguish them: the vesica of V. fragilis has a single lobe (Fig. 22B), whereas the vesica of V. immaculata is bilobed with the second lobe sclerotized. Males of V. fragilis can be separated from V. costata based on genitalia. In V. fragilis, the phallic sclerite is reduced (Fig. 22B), the uncus apex is finger- like and bifid with two hooks (Fig. 27A). In V. costata, the phallic sclerite is narrow and rounded (Fig. 18D), the uncus apex is triangular with two hooks (Fig. 23D). In females, V. fragilis possess pseudopapillae anales (Fig. 24B), and lack an additional bursa (Figs. 24A, 26C), whereas females of V. costata lack pseudopapillae anales and have a connate, double bursa (Fig. 17D). Description. —-Male habitus (Plate 2, No. 7): Head clay throughout. Antennal scales clay. Third seg- ment of labial palpus exceeding front of head by at least half of entire length; all segments clay. Thorax dorsal and ventral scales clay throughout. Legs clay, with patches of flesh ocher scales on femora and tibiae; tar- sal claw simple (Fig. 15A). Forewing length 14 mm (N = 10), R1-R5 all present, M3 present, extending to wing margin (Fig. 12). Dorsal surface clay with cinnamon scales from wingbase through postmedial region, ven- tral surface flesh ocher. Hindwing dorsal surface flesh ocher throughout, ventral surface identical. Abdomen with dorsal, lateral and ventral scales warm buff. Scle- rotization of 8th sternite rectangular, length greater than width (Fig. 28E). Coremata reduced to small patches of scales on membranous lobes. Female habitus (Plate 2, No. 6): Head color same as in male. Antennal color same as in male. Third segment of labial palpus ex- ceeding front of head by at least half of entire length, color same as in male. Thorax color same as in male. Leg color same as in male, tarsal claw simple (Fig. 15A). Forewing length 12 mm (N = 2), venation as in male. Dorsal surface cinnamon extending from wingbase through postmedial region, costal margin peach red, ter- minal wing margins fringed with dark salmon scales, oc- casionally with a faint natal brown spot between medial and postmedial regions present. Although rare, some specimens with a small white spot in annellar region, ventral surface dark salmon suffused with peach red scales. Hindwing dorsal surface peach red, with a natal brown discal spot, natal brown subterminal markings as spots and blotches often not reaching posterior margin of wing, ventral surface identical. Abdomen with color same as in male, but occassionally with patches of fuscous scales on all dorsal segments tapering towards posterior. Male Genitialia: Phallus with rounded coecum penis, straight, basiphallus inflected ventrally at midpoint, curved toward venter (Fig. 25C), phallic sclerite highly reduced (Fig. 22B). Vesica directed ventrally when everted, membrane rugose, bilobed, second lobe sclero- tized, left side of vesica with a small membranous pro- jection located ventrally, right side of vesica with a patch of cornuti located under phallic sclerite (Fig. 22C). Geni- tal capsule dorsum of tegumen concave, bent inward, conjunctiva weakly sclerotized (Fig. 23A). Uncus base cordate (Fig. 23B), ventrad with uncus apex fingerlike, bifid, and with two hooks (Fig. 27A. Scaphium rounded, triangular. Subscaphium absent, extensions of tegumen sclerotized. Juxta cylindrical, emarginated, with ends barbed toward dorsum with broadly rounded ornamen- tation present (Fig. 20C). Costa of valva elongate and tip entire, 20 or more costal setae. Sacculus horizontally obovate, broadly hooked toward costa. Saccus rounded, concave in center. Female genitalia: Pseudopapillae anales present (Fig. 24B). Pheromone glands membra- nous, branching after tip of anterior apophyses, moder- ately thick, greater than 1/8 but less than 1/4 width of 7th abdominal sternite. Ostium bursa slightly concave with small lateral prominences (Fig. 30B). Ductus bursa rectangular, narrowing toward venter, sclerotization ex- tending into corpus bursa (Fig. 17A). Corpus bursa with three signa, membrane wrinkled (Fig. 26C). Variation.— One V. fragilis male with white discal spots on the forewings was found from Alberta, Canada. Few females were available for examination, but no size or color variation was observed. Biology.—- Little is known about the natural his- tory of Virbia fragilis. It appears to have a single brood, emerging in July. Peak female pheromone release and calling behavior occurs in scotophase (e.g., between 20 and 23 hours; Fig 8E), similar to most species in the complex. Distribution.—- This species has been collected in open fields in the Black Hills, South Dakota and in Boul- der, Colorado. The range extends north to Alberta and British Columbia, south to New Mexico. Virbia immaculata Reakirt Pl. 2, Nos. 2 (female), 3 (male) Crocota immaculata Reakirt 1864: 372 [type lo- 90 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 23. A. Dorsal view of uncus-tegumen complex, Virbia costata. Cv = conjunctiva (sclerotization weak), Dt = tegumen (concave, bent inward); B. Dorsal view of uncus-tegumen complex, Virbia aurantiaca. Cv = conjunctiva (sclerotization strong), Dt = tegumen (rounded with three-dimensional shapes on both sides with risen node in center), U = base of uncus (cordate); C. Tegumen, Virbia laeta. Dt = tegumen, entire); D. Uncus, Virbia costata, ¾ view. H = hooks of the uncus (two hooks present), U = uncus (apex an inverted heart-shape); E. Uncus complex, Virbia laeta, ¾ view. H = hooks of the uncus, Sc = scaphium (rounded, triangular); F. Uncus complex, Virbia phalangia, ventral view. U = uncus, Sc = scaphium (flattened). ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 91 cality, Philadelphia, Pennsylvania, USA] Material examined. — Type material: Holotype of V. immaculata: Philadelphia, Pennsylvania//immaculata Reakirt//C. immaculata Reakirt Coll. Reakirt//Original Type, [sic.] FMNH. All material is from the NMNH collection unless otherwise noted: UNITED STATES: Illinois: Edgebrook, 10.x.1947, A. K. Wyatt, Bred// 1019a (1 Male, JMZ105); 7.x.1947, A. K. Wyatt//CNHM 1957 Alex K. Wyatt Lepidoptera Collection (1 Female, 1 Male, JMZ309; JMZ310); Chicago, 8.x.1948, A. K. Wyatt//CNHM 1957 Alex K. Wyatt Lepidoptera Col- lection, FMNH (1 Male, JMZ308). Indiana: Tremont, 25.viii.1952, A. K. Wyatt, ex ova (1 Male, JMZ106). Michigan: Alpena, 10.vii.1931, Dr. G. W. Rawson//G. W. Rawson Collection 1962 (1 Male, JMZ126). New York: Boreas River, 8.vii.1937, L. R. Rupert (1 Male, JMZ104). Pennsylvania: New Brighton, 18.vi.1907// Merrick Museum//Barnes Collection (1 Male, JMZ125); Warrendale, 21.viii.1946, LACM (1 Male, JMZ191). Wisconsin: Racine, O. Meske Collection (1 Male, JMZ107). Diagnosis.—- Virbia immaculata can be confused with V. fragilis, V. aurantiaca, and V. ferruginosa. The characters for separating these are given above under the relevant species’ diagnosis. Description. — Male habitus (Pl. 2, No. 3): Head clay throughout. Antennal scales scales clay. Third segment of labial palpus exceeding front of head by at least half of entire length, all segments clay. Thorax dorsal and ventral scales clay. Legs clay, with patches of flesh ocher scales on femora and tibiae, tarsal claw simple (Fig. 15A). Forewing length 12 mm (N = 10), R1-R5 all present, M3 present, extending to wing margin (Fig. 12). Dorsal surface clay with cinnamon extending from wingbase through postmedial region, ventral sur- face flesh ocher. Hindwing dorsal surface ocher, ven- tral surface identical. Abdomen with dorsal, lateral, and ventral abdominal scales warm buff. Sclerotization of 8th sternite rectangular, length greater than width (Fig. 28E). Coremata reduced to small patches of scales on membranous lobes. Female habitus (Plate 2, No. 2): Head color same as in male. Antennal color same as in male. Third segment of labial palpus exceeding front of head by at least half of entire length, color same as in male. Thorax color same as in male. Leg color same as in male, tarsal claw simple (Fig. 15A). Forewing length 12 mm (N = 10), venation as in male. Dorsal surface cinnamon extending from wingbase through post- medial region, posterior margins fringed with salmon scales, with a faint natal brown spot between medial and postmedial regions present, although rare, some specimens with a small white spot in annellar region, ventral surface dark salmon suffused with peach red. Hindwing dorsal surface peach red, with a natal brown discal spot, subterminal markings as spots and blotches not reaching posterior margin, ventral surface identical. Abdomen with color same as in male. Male genitalia: Phallus with rounded coecum penis, straight, basiphallus inflected ventrally at midpoint (Fig. 22C), phallic sclerite reduced (Fig. 22B). Vesica directed ventrally when everted, membrane rugose, bilobed, second lobe sclero- tized, left side with small membranous projection located ventrally (Fig. 22C), right side of vesica with patch of cornuti under phallic sclerite. Genital capsule with dor- sum concave, bent inward (Fig. 23A), conjunctiva scle- rotized. Uncus base cordate (Fig. 23B). ventrad with uncus apex fingerlike, bifid, with two hooks (Fig. 27A). Scaphium rounded, triangular. Subscaphium absent. Juxta cylindrical, emarginated, with ends barbed toward dorsum, broadly rounded (Fig. 20C). Costa of valva elongate and tip entire, 20 or more costal setae. Saccu- lus horizontally obovate, broadly hooked toward costa. Saccus rounded, concave in center. Female genitalia: Pseudopapillae anales present (Fig. 24B). Pheromone glands membranous, branching before tip of anterior apophyses, moderately thick, greater than 1/8 but less than 1/4 width of 7th abdominal sternite. Ostium bursa slightly concave with small lateral prominences (Fig. 30B). Ductus bursa rectangular, narrowing toward ven- ter, sclerotization extending into corpus bursa (Fig. 17A). Corpus bursa with three signa, membrane wrinkled (Fig. 26C). Variation. — Females of V. immaculata from some localities (e.g., Indiana) are typically smaller (wing length = 11 mm) than in others (e.g., Illinois; wing length = 13 mm). Biology.—- Virbia immaculata has been reared on both dandelion (Taraxacum) and plantain (Plantago) (Zaspel database). This species is bivoltine with adults emerging in June and again in August. Calling occurs in scotophase or the dark period of the light-dark cycle (Fig. 8B) and is advanced by decreasing temperatures (Cardé et al. 1973). Six matings have been observed in an open field in Dryden, New York, all within 30 minutes of sunset (Cardé 1971). Distribution.—- This species occurs north of lati- tude 40 degrees, from the Eastern Coast of North America west to Manitoba. This species can also be found in Iowa, Illinois, and Indiana. Virbia laeta Guerin-Meneville Pl. 3, Nos. 6 (female), 7 (male) Lithosia laeta Guerin-Meneville [1832] 1829- 1844: 322 [type locality: “l’Amérique du Nord”, USA]. 92 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 24. A. Female genitalia, Virbia laeta, dorsal view. Ap = anterior apophysis, B = bursa (smooth), Db = ductus bursa, Pa = papillae anales, Pg = pheromone glands, S = signum; B. Ovipositor, Virbia nigricans, dorsal view. Pg = pheromone glands, PsPa = pseudopapillae anales, internal view. Crocota treatii Grote 1865: 322 [type localty, Massachusetts, USA]. Lithosia rubropicta Packard 1887: 52 [type lo- cality, Rangley Lakes, Maine USA]. Material examined.—- Type material: Holotype of V. laeta: Ex Mus AEO Dris Boisduval//Oberthur Col- lection//Type TF Höfer//Bd//47Bl NMNH. All mate- rial is from the NMNH unless otherwise noted: CANADA: New Brunswick: Canequet, 11.vi.1950, D. C. Ferguson (1 Male, JMZ78). UNITED STATES: Arkansas: Washington Co., Devil’s Den St. Park, 21.vii.1966, R. L. Brown Collector (1 Male, JMZ124). New Hampshire: Claremont, 20.vi.1908, Collection Brklyn Museum (1 Male, JMZ80). Florida: Highlands Co., Archbold Biological Sta. 5.iv.1962, D. C. Ferguson (1 Male, JMZ79). Maine: Lincoln, 8.vii (1 Male, JMZ123). Minnesota: St. Paul, U. Farm, 22.vii.1935, A. A. Granovsky//light trap, UMSP (1 Male, JMZ122). Missouri: Willard, 19.vi., A. E. Brower// Eubaphe laeta treatii (1 Female, JMZ160). Texas: Jasper Co., Town Bluff, 6.vi.1967, A. & M. E. Blanchard (1 Male, JMZ159); Montgomery Co., Conroe, 12.iii.1967, A. & M. E. Blanchard (1 Male, JMZ77). Wisconsin: Marathon Co., Township of Stettin, 6.vii.1978, Russell A. Rahn (1 Male, JMZ81). Diagnosis.—- Both sexes of this species superfi- cially resemble V. costata females; however, V. laeta is restricted to northcentral and southern U.S. and V. costata is restricted to southwestern US. See the diag- nosis of V. costata for additional diagnostic differences. Description.—- Male habitus (Pl. 3, No. 7): Head fuscous, occiput geranium pink. Antennal scales fuscous. Third segment of labial palpus short, first and second segments geranium pink, third segment fuscous. Tho- rax dorsal and ventral scales fuscous with small tufts of geranium pink at wing base. Legs fuscous to dark gray- ish brown, tarsal claw simple (Fig. 15A). Forewing length 11 mm (N = 10), R3 absent (Fig. 13), M3 present, extending wing margin. Dorsal surface fuscous to dark grayish brown with light salmon band running full length of costal margin, ventral surface dusted in light salmon. Hindwing dorsal surface geranium pink, pattern begin- ning at proximal portion of adbasal region, expanding transversely to distal portion of postmedial region, pat- tern extending down wing and meets inner margin of fuscous to dark grayish brown scales extending from ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 93 Figure 25. Male genitalic capsule of Virbia laeta. A- Ventral view, B- Lateral view; C- Left, lateral view of phallus. C = costa, CoP = coecum penis, Dps = dorsal phallic sclerite, Dt = dorsal tegumen, H = hook, J = juxta, Pl = basiphallus, Sa = sacculus, S = saccus, Sb = subscaphium, Sc = scaphium, U = uncus, V = vinculum, Ve = vesica. 94 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) postmedial region to wing tip, occasionally with a slightly sinuate pattern, ventral surface identical. Abdomen ge- ranium pink with small dark grayish brown spots in cen- ter of each abdominal tergite, lateral abdominal scales geranium pink, ventral abdominal scales fuscous to dark grayish brown. Sclerotization of 8th sternite rectangu- lar shape, length greater than width (Fig. 20E). Coremata reduced to small patches of scales on membranous lobes. Female habitus (Pl. 3, No. 6): Head color same as in male. Antennal color same as in male. Third segment of labial palpus exceeding front of head by at least half of entire length, color same as in male. Thorax color same as in male. Leg color same as in male, tarsal claw simple (Fig. 15A). Forewing length 17 mm (N = 10), venation as in male. Dorsal surface color same as in male, ventral surface color same as in male. Hindwing dorsal surface color same as in male, ventral surface color same as in male. Abdomen with color same as in male. Male genitalia: Phallus (Fig. 25C) with rounded coecum penis, inflected ventrally, basiphallus straight, phallic sclerite absent. Vesica directed ventrally when everted, membrane smooth, entire, left side with small membranous projection located ventrally, right side of vesica with patch of cornuti (Fig. 22A). Genital capsule with dorsum entire (Fig. 23C), conjunctiva sclerotized. Uncus base cordate (Fig. 23B), ventrad with uncus apex slightly bifurcate, toothed, with two hooks (Fig. 25A). Scaphium rounded, triangular. Subscaphium completely surrounding anal tube, extensions of tegumen sclerotized. Juxta ovoid with ventral margin expanding into large or- bicular shape (Fig. 25A). Costa of valva elongate and tip entire, 20 or more costal setae. Sacculus blunt and extending under juxta, clawlike. Saccus rounded, con- cave in center. Female genitalia (Fig. 24A): Pseudopapillae anales present (Fig. 24B). Pheromone glands membranous, branching after tip of anterior apo- physes, moderately thick, greater than 1/8 but less than 1/4 width of 7th abdominal sternite. Ostium bursa straight, without prominences (Fig. 17B). Ductus bursa rectangular, narrowing toward venter, sclerotization ex- tending into corpus bursa (Fig. 17B). Corpus bursa with three signa, membrane smooth (Fig. 24A). Biology.—- Larvae have been reared on dande- lion (Taraxacum) and plantain (Plantago) (Forbes 1960). Adults are typically found in pine woodlands (Heppner 2003). In Louisiana, V. laeta can be collected every month except December (Brou 2003). In Texas, two flight periods (March, June) appear to occur, but in more northern parts of the range, collection dates sug- gest a univoltine brood emerging in June and July. Distribution.—- New Brunswick south to Florida, west to Minnesota south to Texas. Virbia lamae Freeman Pl. 1, Nos. 8 (female), 9 (male) Holomelina lamae Freeman 1941: 123 [type lo- cality, Baddeck, Nova Scotia, Canada]. Material examined.— Type material: Holotype not examined. All material is from the NMNH collection unless otherwise noted: CANADA: Nova Scotia: Halifax Co., Bog Prospect Road, 21.vii.1852, D. C. Ferguson (1 Female, JMZ58); Mount Uniake, 19.vii.1946, D. C. Ferguson//Eubaphe lamae Freeman//Otto Bucholz Collection, AMNH (1 Male, JMZ277). UNITED STATES: Maine: Seawall Bog, Acadia Pk. Mt. Desert Is., 29.vii.1966, leg. Ring Cardé//Female Parent of Brood (1 Female, 1 Male, JMZ296; JMZ295); Acadia Pk. Mt. Desert Is, 29.vii.1966, leg. Ring Cardé/ /Female Parent of Brood, RTC (1 Female, JMZ297). Wisconsin: Marathon Co., Township of Norrie, Russella (1 Male, JMZ63). Diagnosis.—- This species can be confused with V. aurantiaca and is occasionally misidentified as V. marginata. Virbia lamae and V. marginata have dis- junct distributions; V. lamae occurs in the Northeastern U.S. and Canada, and V. marginata occurs in northern Central America with disjunct populations in the south- western U.S. Genitalic differences in both sexes separate V. lamae and V. aurantiaca. In male V. lamae, the juxta is cylindrical and emarginated as in V. marginata (Fig. 27C) whereas in V. aurantiaca, the juxta is ovoid with the ventral margin expanding into a larger, orbicular shape (Fig. 20B). In females, V. aurantiaca has pseudopapillae anales (Fig. 24B); V. lamae does not. The ostium bursa is unornamented in V. lamae (Fig. 26B); in contrast, lat- eral prominences surround the ostium bursa in V. aurantiaca (Fig. 26D). The dark coloration in the anal fold of the hindwing can also be used to separate V. lamae from V. aurantica and V. marginata. Description. —-Male habitus (Pl. 1, No. 9): Head ranging from clay color to fawn color among specimens. Antennal scales ranging from clay to fawn among speci- mens. Third segment of labial palpus exceeding front of head by at least half of entire length, all segments clay to fawn. Thorax dorsal and ventral scales from clay to fawn. Legs clay, tarsal claw simple (Fig. 15A). Forew- ing length 10 mm (N = 9); R1-R5 all present, M3 present, extending to wing margin (Fig. 13). Dorsal surface clay with fawn extending from wingbase through postmedial region, ventral surface salmon color extending from wingbase through postmedial region, subterminal region natal brown often not reaching posterior margin, scales over veins retaining salmon scales. Hindwing dorsal surface from yellow ocher to salmon with a natal brown ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 95 Figure 26. A. Female Virbia lamae, dorsal view. Db = ductus bursa (hourglass), Pg = pheromone glands (membra- nous, branching after tip of anterior apophysis, moderately thick); B. Female Virbia lamae, ventral view. Ob = ostium bursa (lacking lateral prominences); C. Female Virbia aurantiaca, dorsal view. B = bursa (with wrinkled mem- brane), Db = ductus bursa (rectangular, narrowing toward ventor, sclerotization extending into corpus bursa), Pg = pheromone glands (membranous, branching before tip of anterior apophysis, moderately thick), PsPa = pseudopapillae anales, internal view, S = signum (three present); D. Female Virbia aurantiaca, ventral view. Ob = ostium bursa (slightly concave with small lateral prominences); E. Female Virbia opella, dorsal view. B = bursa (connate and membrane smooth), Db = ductus bursa (goblet shape, sclerotization extending into corpus bursa), Pg = pheromone glands (membranous, branching after tip of anterior apophysis, thin), PsPa = pseudopapillae anales present; F. Fe- male Virbia opella, ventral view. Ob = ostium bursa (split midventrally with two broad, lateral prominences). 96 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Figure 27. A. Uncus complex, Virbia aurantiaca, ventral view. U = uncus (fingerlike, with bifed apex), H = hooks; B. Uncus in lateral view, Virbia marginata. N = notch (rounded); C. Male genital capsule Virbia marginata. C = costa (distal tip entire), J = juxta (cylindrical, emarginated with ends barbed toward dorsum), Sa = sacculus (horizon- tally obovate, broadly hooked toward costa, with setae present), S = saccus (rounded, concave in center); D. Male genital capsule, Virbia opella. C = costa, (distal tip forked), Sa = sacculus (peanut-shaped), Se = setae; E. Male genital capsule, Virbia feronia. C = costa, (distal tip entire), Sa = sacculus (thin, square-shaped), Se = setae. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 97 discal spot, natal brown subterminal markings a thick band often not reaching posterior margin, ventral sur- face identical. Abdomen with dorsal, lateral and ventral scales light buff with first segment on dorsal side a tuft of fawn scales, additional tergites occasionally with a medium size fawn spot in center, size of spot tapering with each segment towards posterior. Coremata ab- sent. Female habitus (Pl. 1, No. 8): Head color same as in male. Antennal color same as in male. Third seg- ment of labial palpus exceeding front of head by at least half of entire length, color same as in male. Thorax color same as in male. Legs clay, with patches of buff scales on femora and tibiae, tarsal claw simple (Fig. 15A). Forewing length 9.8 mm (N = 4), venation same as in male. Dorsal clay extending from wingbase through postmedial region, posterior margin cinnamon brown, occassionally faint natal brown spot between between medial and postmedial regions present, large white anellar spot present, ventral surface clay extending from wingbase through postmedial region, ocassionally suf- fused with salmon scales, especially along costal mar- gin, natal brown subterminal markings as spots and blotches often not reaching posterior margin, posterior margin clay with scales over veins retaining salmon scales. Hindwing dorsal surface salmon, with a large natal brown discal spot, natal brown subterminal mark- ings as thick spots and blotches often not reaching pos- terior margin, ventral surface identical. Abdomen color same as in male. Male genitalia: Phallus with rounded coecum penis rounded and weakly notched (Fig. 22A), inflected ventrally, basiphallus straight, phallic sclerite wide and nodular (Fig. 22C). Vesica directed ventrally when everted, membrane rugose, bilobed, elongate, sec- ond lobe sclerotized, right side of vesica with patch of cornuti under phallic sclerite (Fig. 22A). Genital cap- sule with dorsum concave, bent inward, conjunctiva weakly sclerotized (Fig. 23A). Uncus base cordate (Fig. 23B), ventrad with uncus apex fingerlike, bifid, with two hooks (Fig. 27A). Scaphium rounded, triangular. Subscaphium absent, extensions of tegumen sclerotized. Juxta cylindrical, emarginated, with ends barbed toward dorsum (Fig. 27C). Costa of valva elongate and tip en- tire, 20 or more costal setae. Sacculus horizontally obo- vate, broadly hooked toward costa. Saccus rounded, concave in center. Female genitalia: Lacking pseudopapillae anales. Pheromone glands membranous, branching after tip of anterior apophyses, moderately thick greater than 1/8 but less than 1/4 width of 7th ab- dominal sternite. Ostium bursa without lateral promi- nences (Figs. 17B, 26A, 26B). Ductus bursa hourglass shaped, sclerotization extending into corpus bursa (Fig. 17B). Corpus bursa with three signa, membrane wrinkled (Fig. 17B). Variation.—- Of the material examined, V. lamae males rarely with white annellar spots on the forewings; no additional variation for males or females was observed. Biology.— Larvae have been reared on plantain (Plantago) and dandelion (Taraxacum) (Zaspel data- base). This diurnal species can only be collected with nets in bogs during July and early August (Cardé, Field notes). Blacklighting in Seawall Bog, Southwest Har- bor, Maine failed to attract any V. lamae, although these were abundant in daytime (Cardé personal observation). Distribution.— This species can be collected in Nova Scotia, New Brunswick, Maine, Wisconsin, and Michigan. Locally, it is restricted to open peat bogs, usually near large bodies of water (maritime or Great Lakes). Virbia marginata Druce Pl. 2, No. 10 (female), Pl. 3, No. 1 (male) Eubaphe marginata Druce 1885: 136, t. 13, f. 8 [type locality, Dueñas, Guatemala] Material examined.—- Type material: Holotype of V. marginata: Type Goodman-Salvin Collection, 97 53’ Dueñas Guatemala, G.C. Champion//Type sp. figured E.C.A. Lep. Het. Eubaphe marginata Druce (NHM). All material is from the NMNH collection unless other- wise noted: GUATEMALA: Guatemala City, viii/ Barnes and Schaus Collection (2 Males, JMZ222; JMZ224). MEXICO: Jalapa, Schaus//ETOwen Do- nor (1 Female, JMZ69). UNITED STATES: Arizona: White Mts., Barnes Collection (1 Male, JMZ221). Colo- rado: Col. remmingtoni det. Ring T. Cardé, LACM (1 Male, JMZ182); Denver, Barnes Collection (1 Male, JMZ184). New Mexico: Jemez Springs, Barnes Col- lection (1 Male, JMZ223); Sandoval Co., 1 mi. S. Sulfer Springs, 7800 ft., 31.vii.1961, F., P. & M. Rindge//Col- lected on NSF grant 69037, AMNH (1 Female, JMZ305). Diagnosis.—- Virbia marginata (Plate 2, No. 10; Plate 3, No. 1) has been treated as a synonym of V. rubicundaria (Plate 2, Nos. 4,5); however, these two species have very different phenotypes. Further, the distributions of the two species do not overlap. Virbia marginata occurs in Mexico, Guatemala and the South- western U.S., whereas V. rubicundaria occurs in the southeastern U.S. Both sexes are morphologically dis- tinct. Males of V. marginata have a tegumen that is concave (Fig. 23A), whereas those of V. rubicundaria have a lobed tegumen (Fig. 23B). Females of both spe- cies have an ostium bursa with lateral prominences (Fig. 30), however, V. marginata lack pseudopapillae anales and typically have one signum (Fig. 17B). Females of V. rubicundaria have pseudopapillae anales and possess 98 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) three signa (Fig. 24B). Occasionally, V. marginata is confused with V. lamae (Plate 1, Nos. 8,9); however, V. lamae is restricted to the northern U.S. and Canada, and the species are not sympatric. Description.—- Male habitus (Plate 3, No. 1): Head yellow ocher. Antennal scales yellow ocher. Third seg- ment of labial palpus exceeding front of head by at least half of entire length, all segments yellow ocher. Thorax dorsal and ventral scales yellow ocher. Legs clay with patches of salmon scales on femora and tibiae, tarsal claw simple (Fig. 15A). Forewing length 10 mm (N = 10), R3 absent, M3 present, extending to edge (Fig. 13). Dorsal surface clay, ventral surface identical, but occa- sionally suffused with salmon scales. Hindwing dorsal surface yellow ocher, pattern beginning at proximal por- tion of adbasal region, expanding transversely to most distal portion of postmedial region, meeting inner margin of raw umber scales, extending from postmedial region to wing apex, occasionally with a slightly sinuate pattern, small raw umber discal spot present, ventral surface iden- tical. Abdomen yellow ocher, dorsal tergites occasion- ally with a medium size fawn color spot in center, size tapering with each segment towards posterior. Scleroti- zation of 8th sternite pentagonal (Fig. 28C). Coremata reduced to small patches of scales on membranous lobes. Female habitus (Pl. 2, No. 10): Head cinnamon. Anten- nal scales cinnamon. Third segment of labial palpus short, cinnamon. Thorax cinnamon. Legs yellow ocher, tarsal claw simple (Fig. 15A). Forewing length 11 mm (N = 1); venation same as in male. Dorsal surface cinnamon, with a faint olive brown discal spot, ventral surface iden- tical. Hindwing dorsal surface flesh ochre, pattern be- ginning at proximal portion of adbasal region, expanding transversely to most distal portion of postmedial region, meeting inner margin of dark raw umber, extending from postmedial region to wing apex, occasionally with a sinu- ate pattern, large dark raw umber discal spot present; ventral surface identical. Abdomen flesh ochre, dorsal tergites occasionally with a medium size dark raw um- ber spot in center, size tapering with each segment to- wards posterior. Male genitalia: Phallus rounded coecum penis rounded weakly notched (Fig. 22A), inflected ven- trally, basiphallus straight, phallic sclerite wide and nodu- lar (Fig. 22C). Vesica directed ventrally when everted, membrane rugose, bilobed, elongate, second lobe scle- rotized, right side of vesica with patch of cornuti under phallic sclerite (Fig. 22C). Genital capsule (Figs. 27C) with dorsum of tegumen concave, bent inward (Fig. 23A), conjunctiva membranous. Uncus base cordate (Fig. 23B), ventrad with uncus rounded at sides with notch at cen- ter, with two hooks (Fig. 27B). Scaphium rounded, pointed. Subscaphium absent, extensions of tegumen sclerotized. Juxta cylindrical, emarginated, with ends barbed toward dorsum. Costa of valva elongate and tip entire, 20 or more costal setae. Sacculus horizontally obovate, broadly hooked toward costa (Fig. 27C). Sac- cus rounded, concave in center. Female genitalia: Lack- ing pseudopapillae anales. Pheromone glands membra- nous, branching after tip of anterior apophyses, thin, less than 1/8 width of 7th abdominal sternite. Ostium bursa slighty concave with small lateral prominences (Fig. 30B). Ductus rectangular, narrowing toward venter (Fig. 17A). Corpus bursa with one signum, membrane smooth (Fig. 17B). Variation.—- Some specimens collected in Colo- rado, New Mexico and the White mountains of Arizona are phenotypically identical to V. marginata and share the diagnostic conditions of the coecum penis and tegumen. Additional V. marginata specimens have been collected from Texas and Mexico, but are rare, making it difficult to examine population variation. The U.S. populations appear to have disjunct distributions from the main species’ range. Potentially, these population isolates are genetically distinct species; however, adult morphology does not distinguish them. Biology.—- Unknown. Distribution.— This species primarily occurs in Mexico, Guatemala, and Honduras. Specimens from New Mexico and Arizona are tentatively identified as V. marginata for reasons provided in Variation. Virbia nigricans Reakirt Pl. 2, nos. 8 (female), 9 (male) Crocota nigricans Reakirt 1864: 371 [type local- ity, Philadelphia, Pennsylvania, USA]. Cothocida nigrifera Walker 1865b: 499 [type lo- cality, “USA”]. Material examined.— Type material: Holotype of V. nigricans: Crocota nigricans Philadelphia, Pa. Reak. Type Coll. Reak.//Crocota nigricans Reak. Pennsylvania Philadelphia Type Coll. Reak. Strecker Colln. Field Museum of Nat. Hist.//genitalic preparation 172 Ring Cardé//Lepidoptera Photograph No. 186 Field Museum [sic.] FMNH. Holotype (synonym of V. nigricans): Holomelina nigricans “D.T.” (green la- bel) nigrifera Walker “Cathocida nigrifera” N. America, 62 85’//Arctiidae slide 969, BM negative 43760, NHM. All material is from the NMNH collection un- less otherwise noted: UNITED STATES: New Jer- sey: Hunterdon Co., 10.vii.1964, Joseph Muller, ex ovo (1 Female, JMZ269); 16.vii.1964, Joseph Muller, ex ovo (1 Female, JMZ270); 11.vii.1964, Joseph Muller, ex ovo (1 Male, JMZ271); 25.vii.1964, Joseph Muller, ex ovo, AMNH (1 Male, JMZ272); Hunterdon Co. (1 Female, JMZ187). ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 99 Figure 28. A. Male 8th sternite, Virbia phalangia. A = antecosta triangular, a = anterior, p = posterior; B. Male 8th sternite, Virbia opella. A= antecosta oval, egg-shaped, a = anterior, p = posterior; C. Male 8th sternite, Virbia marginata. A = antecosta pentagonal, a = anterior, p = posterior; D. Male 8th sternite, Virbia divisa. A = antecosta pentagonal with expanded anterior edge, a = anterior, p = posterior; E. Male 8th sternite, Virbia lutelinea. A = antecosta reinforced by transverse ridge, rectangular, width greater than length, a = anterior, p = posterior; F. Male 8th sternite, Virbia metazonata. A = antecosta M-shaped, a = anterior, p = posterior. 100 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) gion flesh ocher extending from wingbase along ante- rior margin to postmedial region, with a natal brown discal spot, fringes buff to clay color, ventral surface identical. Abdomen buff to clay. Sclerotization of 8th sternite pen- tagonal (Fig. 28C). Coremata reduced to small patches of scales on membranous lobes. Female habitus (Plate 2, No. 8): Head same as in male, but in some specimens infused with orange-rufous. Antennal color same as in male. Third segment of labial palpus short, color usually same as in male or infused with orange-rufous. Thorax buff to clay. Legs buff or clay, tarsal claw simple (Fig. 15A). Forewing length 9.3 mm (N = 3), venation same as in male. Dorsal surface same as in male, occassionally suffused with orange rufous scale, ventral surface same as in male. Hindwing dorsal surface usu- ally same as in male, some specimens with large discon- tinuous terminal bands dark sepia, ventral surface same as in male. Abdomen usually same as in males, some specimens suffused with orange rufous. Male genitalia: Phallus with rounded coecum penis, straight, basiphallus inflected ventrally at midpoint, phallic sclerite wide and nodular (Fig. 22C). Vesica directed ventrally when everted, membrane rugose, bilobed, elongate, second lobe sclerotized, right side of vesica with patch of cornuti under phallic sclerite (Fig. 22C). Genital capsule dor- sally with tegumen concave, bent inward (Fig.23A), con- junctiva membranous. Uncus base cordate (Fig. 23B). Ventrad with uncus apex fingerlike, bifid, with two fused hooks (Fig. 27A). Scaphium conical, pointed. Subscaphium absent. Juxta cylindrical, emarginated, with ends barbed toward dorsum (Fig. 27C). Costa elongate and tip entire, 20 or more costal setae. Sacculus hori- zontally obovate, broadly hooked toward costa. Saccus rounded, concave in center. Female genitalia: Lacking pseudopapillae anales. Pheromone glands membranous, unbranched, thick, greater than 1/4 width 7th abdominal sternite (Fig. 24B). Ostium bursa slighty concave with small lateral prominences (Fig. 30B). Ductus rectangu- lar, narrowing toward venter; sclerotization extending into corpus bursa (Fig. 17A). Corpus bursa with three signa, membrane wrinkled (Fig. 26C). Variation.—- Little variation exists among V. nigricans males; one specimen examined from Penn- sylvania has slightly more pinkish coloration in the hindwing. Similarly, V. nigricans females can vary slightly in size and color. Biology.—- Larvae have been reared on artificial diet; the species is bivoltine with adults emerging in late May to June and again in early to mid August. Males are diurnal and weakly attracted to 2Me-17c (Cardé 1971). Females lay unattached eggs individually like the species V. aurantiaca (Cardé 1971). Diagnosis.—- Virbia nigricans has been confused with dark V. opella specimens. Virbia nigricans is smaller (forewing length 9.3 mm), and the species re- stricted to western New Jersey and eastern Pennsylva- nia. Virbia opella tends to be larger (forewing length 11 mm) and has a wide geographic range. Virbia nigricans is a daymating species (Fig. 8F, Cardé 1971) and its habitat is secondary old fields. Virbia opella is active at night. Additionally, dark coloration in the anal fold of the hindwing can also be used to separate V. nigricans from V. opella. Genitalia of both sexes are diagnostic. Males of V. nigricans lack a subscaphium and the costal tip is en- tire (Fig. 27C); males of V. opella possess a well-devel- oped subscaphium and the costa tip is forked (Fig. 27D). Female V. nigricans have a corpus bursa with 3 signa (Fig. 26C), whereas V. opella females have a connate, double corpus bursa (Fig. 17C) and lack signa. Description.—- Male habitus (Plate 2, No. 9): Head buff to clay. Antennal scales raw umber. Third segment of labial palpus exceeding front of head by at least half of entire length, all segments buff to clay. Thorax dorsal and ventral scales buff to clay. Legs legs buff to clay, tarsal claw simple (Fig. 15A). Forewing length 8.2 mm (N = 10), R1-R5 all present, M3 present, extending to edge (Fig. 12). Dorsal surface tawny buff from base, extending toward postmedial region, fading into clay color through subterminal region to wing apex, with a faint raw umber discal spot present, ventral sur- face light to dark flesh ocher, occurring in varibale amounts. Hindwing dorsal surface natal to army brown extending from inner margin to outer edge, medial re- Figure 29. Adult habitus: Virbia opella Grote, Male: 15 mi W of Cocoa, Orange County, Florida 19.iii.1966, R.H. Leuschner Coll., AMNH. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 101 Distribution.— This species has a highly restricted distribution, occuring only in western New Jersey (Hunterdon Co.) and Pennsylvania (Philadelphia and Montgomery Cos.) in secondary secession habitats. Virbia opella Grote Pl. 3, Nos. 2 (female), 3 (male) Crocota opella Grote 1863: 345 [type locality, Pennsylvania, USA] Crocota flava Barnes & Benjamen 1925: 127- 129 [type locality, Quincy, USA]. Crocota obscura Stretch 1885: 103 [type locality, Pennsylvania, USA] Crocota belmaria Ehrmann 1895: 345 [type lo- cality, Pittsburg, Pennsylvania, USA]. Crocota rubricosta Ehrmann 1895: 345 [type lo- cality Jeanette, USA]. Material examined.—- Type material: Holotype not examined. All material is from the NMNH collection unless otherwise noted: UNITED STATES: Illinois: Quincey, Poling //Barnes Collection (1 Female, JMZ66); 1900.v., Poling //Barnes Collection (1 Female, JMZ158); 1900.vi., Poling //Barnes Collection (1 Male, JMZ157); Magnolia, 1951.ii., A. K. Wyatt, ex ova (1 Female, JMZ61). Massachusetts: Plymouth Co., E. Wareham Agr. Exp. Sta., 10.vii.1976, W. E. Tomlinson at black- light trap (1 Female, JMZ258); E. Wareham Agr. Exp. Sta., 27.vii.1977, W. E. Tomlinson at blacklight trap, AMNH (1 Male, JMZ261); Martha’s Vineyard, F. M. Jones (1 Male, JMZ128). New Jersey: Cape May Co., Eldora, J. Muller //Offspring No. 5, 17.viii. (1 Fe- male, JMZ268); Eldora, 1977.vi.1923, J. Muller (1 Fe- male, JZ290); Eldora, Jos. Muller //offspring No. 5, 17.viii. (1 Male, JMZ267); Orange Mts., 6.ix., Otto Bucholz Collection, AMNH (1 Male, JMZ289); Lakehurst, nigricans (1 Male, JMZ83). New York: Horseheads, 30.viii.1943, L. R. Rupert //Bred ex ova Food: Dande- lion//1022 nigricans (1 Male, JMZ84). Pennsylvania: New Brighton, 7.vii.1975 //Eubaphe opella (1 Male, JMZ82). Diagnosis.—- Virbia opella is a highly phenotypi- cally variable species ranging from a dark charcoal form (Pl. 3, Nos. 2,3) to a pinkish form (Fig. 29). Nearly all have a brown discal spot in the hindwing, although the spot is obscured in the dark form (Plate 3, Nos. 2,3), and paler forms may lack the discal spot. This species is routinely confused with V. nigricans, and occasionally with V. aurantiaca and V. ferruginosa. Virbia nigricans is restricted to western New Jersey and east- ern Pennsylvania, and is a much smaller, darker species (forewing length 9.3 mm) compared to the larger V. opella (forewing length 11 mm). For genitalic differ- Figure 30. A. Ostium bursa, Virbia opella. Ob = ven- tral edge of ostium (split), P = lateral prominence; B. Ostium bursa, Virbia rubicundaria. Ob = ventral edge of ostium bursa (with small, lateral prominences); C. Ostium bursa, Virbia satara. Ob = ventral edge of os- tium (toothed, with lateral prominences). 102 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) ences, see the Diagnosis of V. nigricans. Virbia opella is easily separated from V. aurantiaca and V. ferruginosa based on forewing venation; R3 is absent or just a trace in V. opella and present in the other two species. The pinkish form of V. opella (Fig. 29) can be confused with V. fergusoni (Fig. 91), but V. fergusoni has darker forewings and more pronounced color con- trast in the hindwings. For genitalic differences, see the Diagnosis of V. fergusoni. Description. —-Male habitus (Fig. 28, Pl. 3, No. 3): Head olive brown. Antennal scales olive brown. Third segment of labial palpus exceeding front of head by at least half of entire length, first and second segments orange rufous, third segment olive brown. Thorax dor- sal and ventral scales olive brown. Legs cinnamon, oc- casionally suffused with orange-rufous scales, tarsal claw slightly bifid (Fig. 15B). Forewing length 11 mm (N = 10), R3 absent, M3 present, extending to edge (Fig. 13). Dorsal surface olive brown to dark drab with a sepia discal spot present, ventral surface clay suffused with salmon extending from adbasal region through medial region. Hindwing dorsal surface identical to forewing, ventral surface identical. Abdomen olive brown, edges of tergites, pleurites and sternites layered with scales ranging from clay to orange rufous among specimens. Sclerotization of 8th sternite oval, egg shape (Fig. 28B). Coremata absent. Female habitus (Pl. 3, No. 2): Head cinnamon. Antennal scales fuscous. Third segment of labial palpus short; first and second segments range from clay to orange-rufous, third segment grayish horn. Tho- rax dorsal and ventral scales cinnamon. Legs intermixed with olive brown and clay, tarsal claw slightly bifid (Fig. 15B). Forewing length 12 mm (N = 10), venation as in male.Forewing dorsal surface cinnamon, faint fuscous discal spot present, ventral surface peach red, terminal margins a fringe of fuscous scales. Hindwing dorsal surface peach red, tufts of faint cinnamon scales ex- tending out from wingbase tapering towards antemedial region with a dark fuscous discal spot present, ventral surface identical. Abdomen dorsal abdominal scales peach red, occassionally with fuscous spots on all seg- ments, ventral and lateral scales peach red to clay, pat- tern of fuscous same as dorsal side. Male genitalia: Phallus (Fig. 19D) with rounded coecum penis, straight, basiphallus inflected ventrally at base, curved toward venter, phallic sclerite reduced. Vesica directed ven- trally when everted, membrane rugose, bilobed, elon- gate, second lobe of vesica rugose, left side of vesica with patch of cornuti located ventrally, right side of vesica with patch of cornuti under phallic sclerite. Genital cap- sule (Fig. 19C) dorsum with tegumen entire, conjunctiva membranous (Fig. 23C). Uncus base cordate, (Fig. 23B), ventrad with uncus apex fingerlike, bifid and with two hooks (Fig. 27A). Scaphium rounded, triangular. Subscaphium lobed and incomplete, extensions of tegumen sclerotized (Fig. 19C). Juxta ovoid with ven- tral margin expanding into larger orbicular plate, dorsal margin concave with pointed edges (Figs. 19C, 20A). Costa of valva elongate and tip forked, 20 or more cos- tal setae. Sacculus peanut shaped. Saccus rounded, concave in center. Female genitalia: Pseudopapillae anales present. Pheromone glands membranous, branch- ing after tip of anterior apophyses, thin, less than 1/8 width of 7th abdominal sternite. Ostium bursa split with two, large, lateral prominences (Fig. 30A). Ductus bursa goblet shaped, sclerotization extending into corpus bursa (Fig. 17C). Corpus bursa double, connate, signa absent, both with smooth membrane (Figs. 17C, 26E). Variation.—- Some females of V. opella deviate greatly from the colors discussed in the description. Many females have olive brown to dark drab scales as in the males. Some females have these scales, but the pattern and color discussed in the description can be seen through these dark scales. Some females display the wing pat- tern and color discussed in the description, but are much brighter; these differences in wing color do not seem to be correlated with distribution (e.g., no pattern of geo- graphic variation). This species is most closely related to and may even be confused with V. fergusoni. The most disctinctive male genitalic characters that can be used to separate this species from V. fergusoni species are the presence of a cornutuslike patch located ven- trally on left side of vesica and the lobing of the subscaphium. In addition, V. fergusoni females typi- cally have a signum, while V. opella females are lacking signa. In addition to the morphological differences, wing color and pattern can be used to separate V. opella spe- cies from V. fergusoni. Biology.—- Larvae have been reared on dande- lion (Taraxacum) (Forbes 1960; Heppner 2003; Zaspel database). Virbia opella is multivoltine, and adult males and females fly at night and can be collected at a black light in habitats ranging from oak forest to scrub oak (Martha’s Vineyard, Massachusetts). In Louisiana, V. opella have three broods from November-February (Brou, personal communication). Distribution.— This species occurs from Maine west to Illinois and south to Texas. Virbia ostenta Edwards Pl. 1, Nos. 1 (female), 2 (males) Crocota ostenta Edwards 1881: 12 [type locality, Prescott, Arizona, USA]. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 103 Holomelina calera Barnes 1907: 10 [type local- ity, Huachuca Mts., Arizona, USA]. Material examined. — Type material: Holotype of V. ostenta: Crocota ostenta Type Hy. Ed.//No 8195 Collection Hy. Edwards//Prescott Arizona//7600//Type No. A.M.N.H., AMNH. Holotype (synonym of V. ostenta): July 16-23//Huachuca Mts. Ariz.//Holomelina calera Type Barnes//Photograph Pl. 12 No. 3// PhotographPl. 26 No. 4//47/BL, NMNH. All material is from the NMNH collection unless otherwise noted: UNITED STATES: Arizona: Santa Rita Mts., S. S. Nicolat Madera Canyon, 20.iii.1946 (1 Male, JMZ87); Huachuca Mts., 24-30.vii//Barnes Collection (1 Female, JMZ225). Diagnosis.—- Virbia ostenta is clay colored forew- ings accented with a costal streak of light salmon scales and with geranium pink extending from the proximal por- tion of the adbasal region and meeting with the dark brownish olive ground color (Pl. 1, Nos. 1, 2). Large for Virbia, the forewing length averages 17 mm. This spe- cies resembles V. costata, but V. costata is smaller (Forewing length 11 -12 mm) and has a darker forewing that lacks a costal streak. This species can also be mis- taken for the Mexican V. semirosea Druce, but the forewing of V. semirosea is darker, closer in hue to V. costata females than V. ostenta and the hindwing of V. semirosea is more red than pink like V. ostenta. Description.—- Male habitus (Pl. 1, No. 2): Head yellow ocher. Antennal scales yellow ocher. Third seg- ment of labial palpus exceeding front of head by at least half of entire length, first and second segments a blend of yellow ocher and light salmon scales, third segment dark clay infused with cinnamon. Thorax a blend of yellow ocher and clay. Legs dark brown, sepia, tarsal claw simple (Fig. 15A). Forewing length 17.1 mm (N = 10); R1-R5 all present, M3 present, extends to edge (Fig. 12). Dorsal surface clay with a thin light salmon band running full length of costal margin, ventral surface dusted in light salmon. Hindwing dorsal surface dark brownish olive, pattern with geranium pink beginning at proximal portion of adbasal region, expanding trans- versely to most distal portion of postmedial region, ex- tending halfway down wing and meets ground color with a sinuate margin, ventral surface identical. Abdomen dorsal side dark brownish olive, lateral geranium pink, ventral yellow ocher. Sclerotization of 8th sternite rect- angular, length greater than width (Fig. 28E). Coremata absent. Female habitus (Pl. 1, No. 1): Head sepia. An- tennal scales dark brown, sepia. Third segment of labial palpus exceeding front of head by at least half of entire length, all segments dark brown, sepia. Thorax dark brown, sepia. Legs yellow ochre with sparse patches of light salmon on femora and tibiae, tarsal claw simple (Fig. 15A). Forewing length 18.5 mm (N = 4), venation same as in male. Dorsal surface antique brown with thin peach red band running full length of costal margin, ventral surface dusted in geranium pink. Hindwing dor- sal surface fuscous, pattern with geranium pink begin- ning at proximal portion of adbasal region, expanding Figure 31. A. Male genital capsule Virbia rindgei. J = juxta (cylindrical, emarginated, with ends barbed toward dorsum), Sa = sacculus (horizontally obovate, broadly hooked toward costa), S = saccus (rounded, concave midventrally); B. Phallus, Virbia rindgei, left, lateral view. Dps = phallic sclerite (wide, nodular), Ve = vesica (bilobed, primary lobe elongate with smooth membrane, second lobe rugose), Cpn = patch of cornuti (right side of vesica under phallic sclerite). 104 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) transversely to most distal portion of postmedial region, extending halfway down wing and meets ground color with a sinuate margin, ventral surface identical. Abdo- men fuscous. Male genitalia: Phallus (Fig. 22B) with rounded coecum penis, straight, basiphallus inflected ventrally at base, phallic sclerite present, narrow and rounded. Vesica directed ventrally when everted, mem- brane smooth, entire, left side with patch of cornuti lo- cated ventrally. Genital capsule with dorsum of tegumen concave, bent inward, conjunctiva weakly sclerotized (Fig. 23A). Uncus base cordate (Fig. 23B), ventrad with uncus apex slightly bifurcate, toothed with one hook (Fig. 23E). Scaphium rounded, triangular. Subscaphium completely surrounding anal tube (Fig. 19C), extensions of tegumen sclerotized. Juxta vertically ovoid with ven- tral margin expanding into larger orbicular shape, dorsal margin concave with pointed edges (Fig. 20A). Costa of valva elongate and tip entire, 20 or more costal setae. Sacculus claw-like, extending under juxta on both sides. Saccus rounded, concave in center. Female genitalia: Lacking pseudopapillae anales. Pheromone glands mem- branous, branching before tip of anterior apophyses, thin, less than 1/8 width of 7th abdominal sternite. Ostium bursa slighty concave with small lateral prominences (Fig. 30B). Ductus rectangular, narrowing toward venter (Fig. 17A). Corpus bursa double, connate, three signa present in corpus, both bursae round with smooth membrane (Fig. 17D). Variation.— Female V. ostenta specimens are rare in collections. Almost no variation in color or size was observed in either sex examined. Biology.— Unknown. Distribution.— Virbia ostenta is restricted to the southwestern U.S. (New Mexico, Arizona) and Mexico, occurring in mountain ranges. Virbia rindgei Cardé n. sp. Pl. 3, Nos. 8 (female), 9 (male) Holotype.— Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., WYO., 5900 ft , 8.vii.1962., F., P. & M. Rindge//Collected on NSF Grant G-25314, Male, AMNH. Paratypes.—- UNITED STATES: South Da- kota: Custer Pk.: S. Dak. Lawrence Co., 6794 ft., 1.vii.1962, F., P. & M. Rindge// JMZ307, Male, AMNH; Lead, Lawrence Co., S. Dak., 5200 ft., 25.vii.1953, F. & P. Rindge, Male, AMNH; Spring Creek Camp 11 mi. NE Hill City S. Dak., 30.vii.1955, F. & P. Rindge, Male, AMNH. Wyoming: Reuter Canyon Camp 5 mi. N. Sundance Crook Co., 5,900 ft., 10.vii.1962, F. P. & M. Rindge//Collected on NSF Grant G 25134//Genitalic preparations 1963-41 Ring Cardé, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 6100 ft., 13.vii.1959, F., P. & B. Rindge//JMZ276, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 5900 ft., 8.vii.1962, F., P. & M. Rindge, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 5900 ft., 8.vii.1962, F., P. & M. Rindge, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 5900 ft., 9.vii.1962, F., P. & M. Rindge, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 5900 ft., 9.vii.1962, F., P. & M. Rindge//Collected on NSF Grant G-25314//Genitalic Preparation 163-42 Ring Cardé, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 5900 ft., 9.vii.1962, F., P. & M. Rindge//Collected on NSF Grant G-25314//Genitalic Preparation 164-65 Ring Cardé, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 5900 ft., 3.vii.1962, F., P. & M. Rindge//Collected on NSF Grant G-25314//Genitalic Preparation 164-72 Ring Cardé, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 5900 ft., 9.vii.1962, F., P. & M. Rindge//Collected on NSF Grant G-25314//Genitalic Preparation 164-82 Ring Cardé, Male, AMNH; 5-6 mi. N. of Sundance Crook Co., 5900- 6300 ft., 4.vii.1962, F., P. & M. Rindge, Male, AMNH; Reuter Canyon Camp 5 mi. N. of Sundance Crook Co., 5900 ft., 8.vii.1962, F., P. & M. Rindge//166//Collected on NSF Grant G-25314, Female, AMNH. All material is from the NMNH collection unless otherwise noted: UNITED STATES: South Dakota: Gunnison Co., 6 mi. E. of Somerset, 29.vi.1959, leg. C. L. Remington (1 Female, JMZ306); Custer Park, 6500-6794 ft., 30.vii.1955, F. & P. Rindge (1 Male, JMZ311). Etymology. —-This species is named in honor of Dr. Fred Rindge. He is among the first to collect this species and recognize it as distinct from other species in the V. aurantiaca complex. Dr. Rindge has had an illustrious career as a Lepidopterist and his collections have greatly increased our knowledge of North Ameri- can lepidopteran fauna. Diagnosis.—- Virbia rindgei might be confused with V. aurantiaca. Wing pattern, coloration and geni- talia readily separate these species. Virbia rindgei males have much darker forewings with rusty undertones (Fig. Pl. 3, No. 8), while V. aurantiaca usually have solid brown forewings (Pl. 1, No. 4). The hindwings of V. rindgei have dark brown shading from the subterminal band along the fold from discal cell to 3A (often cover- ing most if not all of the dark brown discal spot), whereas the subterminal area of V. aurantiaca is a solid band. Virbia rindgei females (Pl. 3, No. 9) have much brighter forewings than V. aurantiaca females (Pl. 1, No. 5). In males, the dorsal margin of the juxta of V. rindgei is ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 105 concave with pointed edges (Fig. 31A) whereas the juxta of V. aurantiaca lacks this elaborate dorsal margin (Fig. 20B). Female genitalia are not diagnostic. Although V. aurantiaca is widespread, it has not been collected in the Black Hills of South Dakota (Lawrence Co.) or Wyoming (Crook Co.). Description. — Male habitus (Pl. 3, No. 8): Head mikado brown. Antennal scales mikado brown. Third segment of labial palpus exceeding front of head by at least half of entire length, first and second segments peach red, third segment mikado brown. Thorax mi- kado brown. Legs cinnamon, with patches of salmon color scales on femora and tibiae, tarsal claw simple (Fig. 15A). Forewing length 12 mm (N = 10); R1-R5 all present, M3 present, extending to edge (Fig. 12). Dorsal surface mikado brown, faint sepia discal spot present, ventral surface a mixture of salmon and peach red scales throughout, posterior margin a fringe of sepia. Hindwing dorsal surface peach red, subterminal region a thick band of sepia meeting postmedial region in sinuate pattern, in most specimens sepia scales shading in region from sub- terminal band along fold from discal cell to 3A, large sepia discal spot present, ventral surface identical. Ab- domen clay, dorsal tergites occasionally with medium size sepia spot in center, size tapering with each seg- ment towards posterior. Sclerotization of 8th sternite pentagonal (Fig. 28C). Coremata reduced to small patches of scales on membranous lobes. Female habi- tus (Pl. 3, No. 9): Head dark cinnamon. Antennal scales dark cinnamon. Third segment of labial palpus exceed- ing front of head by at least half of entire length, all segments dark cinnamon. Thorax mikado brown. Legs clay, with patches of salmon on femora and tibiae, tarsal claw simple (Fig. 15A). Forewing length 12 mm (N = 3), venation same as in male. Forewing dorsal surface mikado brown throughout, faint natal brown discal spot present, occassionally small white spot in annelar region present, ventral surface identical, dusted with peach red. Hindwing dorsal surface peach red, subterminal region a thick band of sepia scales, meeting postmedial region in sinuate pattern, in most specimens, sepia scales shad- ing in region from subterminal band along fold from discal cell to 3A with a large sepia discal spot, ventral surface identical. Abdomen dorsal, lateral, and ventral abdomi- nal scales scales clay to peach red, dorsal tergites with large sepia spot in center, size tapering with each seg- ment towards posterior. Male genitalia: Phallus with rounded coecum penis, inflected ventrally, straight, phallic sclerite wide and nodular (Fig. 31B). Vesica directed ventrally when everted, membrane smooth, bilobed, sec- ond lobe sclerotized (Fig. 22C), right side of vesica with patch of cornuti under phallic sclerite (Fig. 31B). Geni- tal capsule (Fig. 31A) with dorsum of tegumen lobed, sides divergent, conjunctiva membranous. Uncus base cordate (Fig. 23B), ventrad with uncus apex fingerlike, bifid, and with two fused hooks (Fig. 27A). Scaphium rounded, triangular. Subscaphium absent. Juxta cylin- drical, emarginated, with ends barbed toward dorsum. Costa of valva elongate and tip entire, 20 or more costal setae. Sacculus horizontally obovate, broadly hooked toward costa. Saccus rounded, concave in center. Fe- male genitalia: Pseudopapillae anales present. Phero- mone glands membranous, branching after tip of ante- rior apophypses, moderately thick, greater than 1/8 but less than 1/4 width of 7th abdominal sternite. Ostium bursa slightly concave with small lateral prominences (Fig. 30B). Ductus bursa rectangular, narrowing toward venter, sclerotization extending into corpus bursa (Fig. 17B). Corpus bursa with three signa, membrane of the corpus bursa wrinkled (Fig. 26C). Variation.—- Virbia rindgei has a highly restricted distribution and is only known from specific localities in Colorado, South Dakota and Wyoming. This species is rare in collections and only represented by small series of mostly males in some of the larger collections (e.g., AMNH & NMNH), rendering further study difficult. Recent collecting attempts in South Dakota and Wyo- ming were unsuccessful (R.T. Cardé, D. Ferguson, J. Zaspel). This species has not been collected since the late 1970’s. Biology.—- The larvae are not known. The moths were collected in Ponderosa Pine forest with an open Figure 32. Male genital capsule Virbia rubicundaria. J = juxta (cylindrical and emarginated with ends barbed toward dorsum), Sa = sacculus (horizontally obovate and broadly hooked toward costa), S = saccus (rounded and concave midventrally). 106 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) understory (Rindge unpublished field notes). From the label data, this species appears to be single brooded, with a flight in mid – late July, possibly early August. This species has been collected at elevations near or above 6000 feet. Distribution.—- This species has been collected at high elevations in Colorado, South Dakota and Wyoming. Virbia rubicundaria Hübner Pl. 2, Nos. 4 (female), 5 (male) Crocota rubicundaria Hübner 1827: 256 [type locality, Georgia, USA]. Crocota belfragei Stretch 1885: 103 [type local- ity, Texas, USA]. Crocota brevicornis Walker 1854b: 536 [type lo- cality, “USA”]. Crocota diminutiva Graef 1887: 42 [type locality, Texas, USA]. Crocota rosa French 1890: 133 [type locality, Texas, USA]. The holotype of Crocota rubicundaria is lost ac- cording to Dr. Lloyd Martin, Curator of Lepidoptera at Naturhistorisches Museum, Wien (personal communi- cation 6/25/2004). A neotype is designated here and labeled to ensure nomenclatural stability. The specimen was chosen from the nearest locality to the original type locality. Crocota rubicundaria Hübner, neotype Male [NMNH] is labeled: Virbia rubicundaria (Hbn. 1827) [RED LABEL]. Label data: GA: Trader’s Hill, Folkston, Charlton County 27.ix.1992, D.C. Ferguson//JMZ59, Female, NMNH. Material examined.—- Type Material. Holotype lost. Syntype (synonym V. rubicundaria): Holomelina brevicornis syntype “140” Arctiidae slide 905 BM nega- tive 43756, NHM. All material is from the NMNH col- lection unless otherwise noted: UNITED STATES: Florida: Gainesville, iv.1969, Dale Hubeck (1 Female, 1 Male; JMZ294, JMZ293); Highlands Co., Archibold Biological Station, 23.vi.1966, Ring T. Cardé (1 Male, JMZ291); Archibold Biological Station, 16.vi.1966, Ring T. Cardé, RTC (1 Male, JMZ292); Florida City, 20.iv.1947, Otto Bucholz //Otto Bucholz Collection (1 Male, JMZ252); Lakeland, 5.v.1912 (1 Male, JMZ253); Monticello, 4-8.x.1914, AMNH (1 Male, JMZ254); Lutz, F. W., 1.iv.1916, LACM (1 Male, JMZ190); Florida City, 23.iv.1947, Otto Bucholz //Otto Bucholz Collection (2 Females, JMZ248; JZ249); 15.iv.1947, Otto Bucholz // Otto Bucholz Collection, AMNH (1 Female, JMZ250). Georgia: Liberty Co., St. Catherine’s Island, 7- 12.viii.1991, E. Quinter & A. Sharkov, AMNH (1 Fe- male, JMZ251). Texas: Anderson Co., Tennessee Colony, 21.vii.1965, A. & M. E. Blanchard (1 Male, JMZ188); Bosque Co., Laguna Park, 24.ix.1970, A. & M. E. Blanchard (1 Male, JMZ189). Diagnosis.—- Virbia rubicundaria females can be confused with V. aurantiaca, however, these spe- cies are rarely sympatric. See diagnosis of V. aurantiaca. Virbia rubicundaria (Pl. 2, Nos. 4,5) can be con- fused with V. fergusoni (Pl. 3, Nos. 4,5), but these spe- cies can be separated based on size, coloration, and geni- talia of both sexes. Virbia rubicundaria is typically smaller (wing length 9 mm) with a lighter forewing (clay with light salmon highlights) compared to V. fergusoni (wing length 11 mm) with darker raw umber coloration. The male genitalia of V. rubricundaria lack a subscaphium (Fig. 32), the coecum penis is notched and the phallic sclerite is wide and nodular (Fig. 22C) whereas V. fergusoni males have a large subscaphium (Fig. 19A), a rounded coecum penis and reduced phallic sclerite (Fig. 19B). Females of V. rubricundaria have a single bursa (no additional bursae) ornamented with three signa (Fig. 26C). In contrast, females of V. fergusoni have a bi- lobed corpus bursa with a single signum (Fig. 26E). Description.—- Male habitus (Pl. 2, No. 5): Head clay with light orange hues. Antennal scales clay with light orange hues. Third segment of labial palpus ex- ceeding front of head by at least half of entire length, all segments a blend of clay and flesh ocher. Thorax clay with light orange hues. Legs a blend of clay and flesh ocher, tarsal claw simple (Fig. 15A). Forewing length 8.9 mm (N = 10); R1-R5 all present, M3 present, extend- ing to edge (Fig. 12). Dorsal surface clay with light orange hues, ventral surface dark salmon. Hindwing dorsal surface flesh ocher, ventral surface ranging from identical to light salmon. Abdomen flesh ocher to dark. Sclerotization of 8th sternite pentagonal (Fig. 28C). Coremata reduced to small patches of scales on mem- branous lobes. Female habitus (Pl. 2, No. 4): Head color same as in male. Antennal color same as in male. Third segment of labial palpus exceeding front of head by at least half of entire length, color same as in male. Thorax same as in male. Legs same as in male, tarsal claw simple (Fig. 15A). Forewing length 8.7 mm (N = 10); venation same as in male. Forewing dorsal surface clay with light orange hues, ventral surface identical, dark salmon with a faint natal brown spot between me- dial and postmedial regions present. Although rare, some specimens with small white spot in annelar region, ven- tral surface identical. Hindwing dorsal surface flesh ocher with a natal brown discal spot present, natal brown subterminal markings as spots and blotches often not reaching posterior margin, ventral surface identical. Abdomen same as in male. Male genitalia: Phallus with ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 107 rounded coecum penis and weakly knotched (Fig. 22A), inflected ventrally, basiphallus straight, phallic sclerite wide and nodular (Fig. 22C). Vesica directed ventrally when everted, membrane rugose, bilobed, elongate, sec- ond lobe sclerotized, right side of vesica with patch of cornuti under phallic sclerite (Fig. 22C). Genital cap- sule (Fig. 31) dorsum with tegumen lobed, sides diver- gent, conjunctiva membranous. Uncus base cordate (Fig. 23B), ventrad with uncus apex fingerlike, bifid, with two fused hooks (Fig. 27A). Scaphium rounded, triangular. Subscaphium absent. Juxta cylindrical, emarginated, with ends barbed toward dorsum. Costa of valva elongate and tip entire, 20 or more costal setae. Sacculus hori- zontally obovate, broadly hooked toward costa. Saccus rounded, concave in center. Female genitalia: Pseudopapillae anales present (Fig. 24B). Pheromone glands membranous, branching before tip of anterior apophypses, moderately thick, greater than 1/8 but less than 1/4 width of 7th abdominal sternite. Ostium bursa with small prominences on both sides (Fig. 30B). Duc- tus bursa rectangular, narrowing toward venter, sclero- tization extending into corpus bursa (Fig. 26C). Corpus bursa with three signa, membrane of the corpus bursa wrinkled (Fig. 26C). Variation.—- Little variation in color pattern exists among male V. rubicundaria specimens. Although rare, some males have a faint trace of a subterminal band on the hindwings. Some V. rubicundaria males from the southwest were slightly larger than those from Florida and Georgia. Of sixteen males, three had a rounded coecum penis, not gently notched. Some females have a brownish discal spot in the forewing and one speci- men had a faint white spot in the annelar region of the forewing. Female hindwing patterns were consistent. Biology.—- Larvae have been reared on dande- lion (Taraxacum) and wild lettuce (Lactuca floridana Gaertner) (Zaspel Database). A multivoltine species, moths are collected year-round throughout its range. Distribution.— This species is found in Georgia and Florida, and distributed along the Gulf Coastal States to east Texas. ACKNOWLEDGMENTS We thank Michelle DaCosta, Howard Frank, Ralph Holzenthal, William Miller, and three anonymous reviwers for helpful comments. We also thank Demetrios Vital for assistance with illustrations and Kevin Nixon for assistance with the website. This work was sup- ported SJW funding sources: NSF-DEB-9981416, NSF- REU supplement, and a Minnesota Experiment Station Project, MN-17-022. 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Weller, 2006. Review of generic limits of the tiger moth genera Virbia Walker and Holomelina Herrich-Schäffer (Lepidoptera: Arctiidae: Arctiinae) and their biogeography. Zootaxa. 1159: 1-68. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 111 112 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Plate 1: 1-10 1. Adult habitus: Virbia ostenta Edwards, Female: Huachuca Mountains., Arizona//Barnes Collection//24-30.vii., USNM. 2. Adult habitus: Virbia ostenta Edwards, Male: Williams, Coco-Nino Co., Arizona, 10.viii.1955, N. McFarland at light, LACM. 3. Adult habitus: Virbia aurantiaca Hübner, Male: Hartford Co., Farmington, Connecticut 9.vi.1961, leg. Ring T. Cardé, RTC. 4. Adult habitus: Virbia aurantiaca, Hübner, Male: Guilford Co., New Haven, Connecticut, 17.viii.1966, leg. Ring T. Cardé, RTC. 5. Adult habitus: Virbia aurantiaca Hübner, Male: Guilford Co., New Haven, Connecticut, 19.viii.1966, leg. Ring T. Cardé, RTC. 6. Adult habitus: Virbia ferruginosa Walker, Female: Edgebrook, Illinois 1-11.ix., A. Kwiat Bred//CNHM 1957 Alex. K. Wyatt Lepidoptera Colln, FMNH. 7. Adult habitus: Virbia ferruginosa Walker, Male: Guilford, New Haven CONN. 3.vii.1965 leg. Ring T. Cardé, RTC. 8. Adult habitus: Virbia lamae Freeman, Female: Seawall Bog, Acadia Pk., Mt. Desert Is., Maine 25.vii.1964, leg. Ring T. Cardé//Female parent of Brood, RTC. 9. Adult habitus: Virbia lamae Freeman, Male: Seawall Bog, Acadia Pk., Mt. Desert Is., Maine 29.vii.1966, leg. Ring T. Cardé, RTC. 10. Adult habitus: Virbia costata Stretch, Female: Prescott, Yavapai County Arizona 28.vii.1970, Collected by Lloyd Martin//Holomelina intermedia (Graef) det. Ring T. Cardé, RTC. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 113 114 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Plate 2: 1-10 1. Adult habitus: Virbia costata Stretch, Male: Otero Co. 11.v.1926, N. Mexico//Harold M. Bower Coll. Access’d C.A.C.M. 1964, FMNH. 2. Adult habitus: Virbia immaculata Reakirt, Female: Edgebrook Illinois, Bred 7.x.1947, A.K. Wyatt Coll.//CNHM 1957 Alex K. Wyatt Lepidoptera Colln., FMNH. 3. Adult habitus: Virbia immaculata Reakirt, Male: Edgebrook Illinois, Bred 7.x.1947, A.K. Wyatt Coll.//CNHM 1957 Alex K. Wyatt Lepidoptera Colln., FMNH. 4. Adult habitus: Virbia rubicundaria Hübner, Female: FLA., Highlands Co. Archbold Biol. Sta. 20.vi.1966, Ring T. Cardé, RTC. 5. Adult habitus: Virbia rubicundaria Hübner, Male: FLA., Highlands Co. Archbold Biol. Sta. 20.vi.1966, Ring T. Cardé, RTC. 6. Adult habitus: Virbia fragilis Strecker, Female: Spring Creek Camp 11 mi. NE Hill City Pennington County S.Dak., 1.viii.1955, F. and P. Rindge, AMNH. 7. Adult habitus: Virbia fragilis Strecker, Male: Custer Park Lawrence County, S. Dakota, 6500-6794 feet, 30.vii.1955, F. and P. Rindge, AMNH. 8. Adult habitus: Virbia nigricans Reakirt, Female: Hunterdon Co., New Jersey 11-16.v.1964, Joseph Muller//ex ovo, AMNH. 9. Adult habitus: Virbia nigricans Reakirt, Male: Frenchtown, New Jersey, Hunterdon Co. Rte.29/10 mi. N Rte. 519, 4.viii.1967, R. Cardé, RTC. 10. Figure 86. Adult habitus: Virbia marginata Druce, Female: 1 mile S. Sulfer Springs, Sandoval Co., New Mexico, 7800 ft., 31.vii.1961, F. and P. Rindge.//Collected on NSF grant G 9037, AMNH. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 115 116 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3) Plate 3: 1-9 1. Adult habitus: Virbia marginata Druce, Male: Guatemala City Guatemala//Schaus and Barnes coll//Dognin Col- lection//July//Holomelina marginata, USNM. 2. Adult habitus: Virbia opella Grote, Female: Magnolia Ill. ex ova ii.1951, A.K. Wyatt, USNM. 3. Adult habitus: Virbia opella Grote, Male: Magnolia Ill. ex ova 9.vii.1950, A.K. Wyatt//Otto Buchholz Collection, AMNH. 4. Adult habitus: Virbia fergusoni Zaspel, Female: FLA, Highlands Co., Highlands Hammock State Park, 12.vi.1987, T.L. Schiefer//Holomelina n. sp. det. D.C. Ferguson, MSU. 5. Adult habitus: Virbia fergusoni Zaspel, Male: GA: 7 mi. NE Fargo, Clinch Co., 13.vii.1981, D.C. Ferguson// Reared ex ovo on Taraxacum, USNM. 6. Adult habitus: Virbia laeta Guérin-Méneville, Female: Willard Missouri 19.vi., A.E. Brower//Eubaphe laeta Guérin- Méneville Treatii Grote, USNM. 7. Adult habitus: Virbia laeta Guérin-Méneville, Male: Marathon County Wisconsin, Township of Stettin 6.vii.1978, Russell A. Rahn, USNM. 8. Adult habitus: Virbia rindgei Cardé, Male: Custer Park South Dakota, Lawrence County, 1.vii.1962, F., P., & M. Rindge, AMNH. 9. Adult habitus: Virbia rindgei Cardé, Female: Spearfish Caynon, Black Hills South Dakota, 1.vii.1939, A.C. Frederick, USNM. ZASPEL, WELLER and CARDÉ : Virbia of America north of Mexico 117 118 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(3)