A new species of Eupsilia Hübner phenotypically allied to E. cirriplaea (Franclemont, 1952) and E. sidus (Guenée, 1852) is described from northeastern North America. Identification of E. schweitzeri, n. sp., is most reliably made on the basis of larval morphology or genetic data, although most adults can be determined using subtle forewing features, including their derived forewing scale type. The adult, genitalia, forewing scales, and larvae of the new species are illustrated. Keys are provided to adults and larvae of the seven northeastern Eupsilia. Eupsilia walkeri (Grote, 1864) is synonymized under E. vinulenta, n. syn. Scopelsoma colorado (Smith, 1903) is removed from synonymy with Eupsilia sidus and given valid species status as Eupsilia colorado, rev. stat., and a neotype is designated for E. sidus.
Eupsilia (Hübner, 1821) species are among the most abundant fall- and spring-active moths in the Northeast (Wagner et al. 2011). On warm winter and early spring nights, they account for many of the orange-brown moths seen in car headlights when driving through wooded habitats, and often outnumber all other moths at bait from January to early April, with densities sometimes reaching 50–200 or more moths at a single bait patch. There are seven species of Eupsilia in the northeastern United States (Forbes 1954, this work)—one of the more common of which has gone undescribed until now. The new taxon is named after Dale F. Schweitzer, who first recognized the new species based on larval characters. This new species is commonly confused with Eupsilia cirripalea Franclemont 1952, E. sidus (Guenée, 1852), and E. vinulenta (Grote, 1864). All seven Eupsilia fly sympatrically and synchronically at some locations (e.g., East Rock State Park in New Haven, Connecticut). Their abundance, species diversity, variability, and overlapping co-occurrence, collectively made it possible for this common eastern moth to go unnamed.
In this paper we describe the new species, illustrate its adult and larval stages, and provide an adult key and the first larval key for eastern Eupsilia. We identify and illustrate three types of forewing scales possessed by Eupsilia. We briefly describe the life history, as far as known (there is little knowledge of late instar feeding habits because larvae leave their natal hosts and are likely nocturnal). We also compare the new species to the western taxon, E. colorado (Smith, 1903), which appears to be closely allied to E. sidus and our new species.
Materials and Methods
The adult description of E. schweitzeri is based on reared specimens from Connecticut and New Jersey. The larval description of E. schweitzeri is based on living larvae, preserved larvae, and larval images. Larvae were compared to reared specimens and larval images of E. cirripalea, E. devia (Grote, 1875), E. morrisoni (Grote, 1874), E. sidus, E. tristigmata (Grote, 1877), and E. vinulenta. High-resolution photographs of forewing scales were taken with a MacroSolutions® stacking system. Genitalia of the male holotype and female paratype were prepared and mounted according to Lafontaine (2004). Eighty-two slide mounted genitalic preparations made by John G. Franclemont were examined from the CUIC; as were 3 additional slides made by Tim McCabe in the NYSM. COI sequences were generated by the Barcodes of Life Project (BOLD); sequences for two E. schweitzeri specimens have been submitted to BOLD. Additional sequences of the new species have also been submitted by J. Bolling Sullivan. Adult paratypes of E. schweitzeri (n=87) have been deposited in the following collections:
Canadian National Collection, Ottawa, Ontario, Canada.
Cornell University Insect Collection, Ithaca, NY, USA
Eric Quinter Collection, Windham, CT, USA.
New York State Museum, Albany, NY, USA.
Peabody Museum of Natural History, Yale University, New Haven, CT, USA.
University of Connecticut, Storrs, Connecticut, USA.
National Museum of Natural History, formerly United States National Museum, Washington DC, USA
Material Examined. HOLOTYPE male (Figs 1, 14, 15). CT: Windham Co., 0.7 mi. ESE Windham 240ft. junction Follett Rd. and Potters Brook // f#1 coll. in bait trap bred ex ova on Prunus serotina, emerged: ix.21-xi.8.2013, Eric L. Quinter, collector // Barcodes of Life Project DLW-000168, Specimen ID CNCLEP 00116373 // Barcode of Life leg removed DNA extracted // Genitalia CNC slide # ♂ 16800 (CNC). Paratypes (adults) (51 males, 36 females). Connecticut: New Haven County. Ex Ovis, pair A87, from CONN.: New Haven, West Rock, coll. March 1987 f white spot, ♂ yellow Ecl. August 18–September 15 1987 reared Dale Schweitzer // Sleeved late April to mid June, started on apple, 4th instar to Carya glabra, last instar on Juglans nigra at Dedham, Mass. (18 ♂, 6 f); same data as above except: Ecl. August 22-August 31 1987 Reared indoors on cuttings: oaks, hickory, apple, wild cherry etc. April-June 1987 (1♂, 2 f); same data as above except: Reared Dale Schweitzer Sleeved late April to mid June 1987 on apple at Dedham, Mass. (6♂, 1 f) (PMNH, UCMS); New Haven east base West Rock Jct. Baldwin Dr & Wintergreen Ave 11 March 1982 D. F. Schweitzer // McCabe gen. slide 1729 (1f) (NYSM); New Haven east base West Rock Jct. Baldwin Dr & Wintergreen Ave 17 August 1981 D. F. Schweitzer // [tlm 81-12 Reared ex ovo on Quercus ilicifolia] // McCabe gen. slide 1683 (1♂) (NYSM). Windham County. CT: Windham Co., 0.7 mi. ESE Windham 240ft. junction Follett Rd. and Potters Brook // f#1 coll. in bait trap bred ex ova on Prunus serotina (1 f), emerged: ix.21-xi.8.2013, Eric L. Quinter, collector // Specimen ID CNCLEP 00116374 // Barcode of Life leg removed DNA extracted (CNC). Ct: Windham Co., 0.7 mi. ESE Windham 240ft. junction Follett Rd. and Potters Brook // [ex ova] Eric Quinter, [adults emerged] ix.21-xi.2013, host: Prunus serotina (9 f, 12 ♂) (ISIC, UCMS, USNM); Ct: Windham Co., 0.7 mi. ESE Windham 240ft. junction Follett Rd. and Potters Brook // [ex ova] Eric Quinter, [adults emerged] Sept. 22–Oct. 4, 2007, host: Prunus serotina (10 ♂, 12 f) (ISIC). Massachusetts: USA: Miles Standish SF Mass.: Plymouth Co 26 Sept. 1987 D. F. Schweitzer // [tlm 87-4 Reared ex ovo on: Quercus ilicifolia] (2 ♂) (NYSM, UCIC); USA: Miles Standish SF Mass.: Plymouth Co 26 Sept. 1987 D. F. Schweitzer // [tlm 87-4 Reared ex ovo on: Quercus ilicifolia] // McCabe gen slide 5076 (1f) (NYSM). New York: Albany County. USA: Albany Pine Bush N.Y.: Albany County 42.33.43 — 73.51.54 24 November 1979 T. L. McCabe 100 m // McCabe gen. slide 506 (1♂) (NYSM. Orange County. USA: Bear Mtn. N.Y.: Orange Co. 41.18.14 — 74.00.24 26 Sept. 1989 T. McCabe 400 m // [tlm 89-13 L6 found feeding on Cirsium] // McCabe gen. slide 5306 (1♂) (NYSM). Tompkins County. Six Mile Creek Tompkins Co., N.Y. August 1958 JG Franclemont [Reared ex ovo on Prunus serotina] (2 ♂) (NYSM).
Etymology. We name this new species Eupsilia schweitzeri, after Dale F. Schweitzer, whom we credit for first recognizing this new species in the late 1980s.
Eupsilia schweitzeri (Figs 1–3) is closely similar to E. sidus (Figs 5, 6) and E. cirripalea, especially when weather worn or having overwintered. Fresh adults usually have three bands of violaceous scales (inside the antemedial, outside the postmedial, and inside of the dark crenulated adterminal lines). E. sidus tends to have a stronger sienna aspect to the FW color; E. cirripalea tends to have a more uniform brown aspect to the FW color; and E. colorado has a light-brown to tan FW with a faint cast of umber (bearing in mind that specimens we had for study are 120 years in age). E. schweitzeri and E. sidus tend to have a more conspicuous silverywhite, pale to fiery-orange, or stramineous reniform spot on the FW than is typical for either E. cirripalea or E. colorado (Fig. 4). The outer FW margin is distinctly crenulated in E. cirripalea; modestly crenulated in E. schweitzeri, but decidedly less crenulated in E. colorado and E. sidus. The FW scales beyond the postmedial line of E. cirripalea, E. sidus, and E. schweitzeri have strongly recurved outer teeth (excelsior scales of Franclemont); E. cirripalea and E. sidus often have greater numbers of modified scales distal to the postmedial relative to E. schweitzeri. (see also Figs 17–22). Distal to the postmedial line the FW scales of E. colorado are less recurved than those of E. cirripalea, E. schweitzeri, and E. sidus, and are essentially unmodified (flattened and shingled). (Contrary to keys and claims by earlier workers, E. morrisoni also possess modified wing scales (Fig. 18), and these tend to be in the basal half on the wing, sparser in number, and less overlapping.) In the male genitalia, the new species lacks the diagnostic medial blister on the juxta of E. cirripalea (Franclemont 1952, Forbes 1954). It can be separated from E. sidus and E. colorado by its more robust (broader) valve (in E. colorado and E. sidus the valve is more elongate and narrowed through much of its length) (Figs 10, 12, 14); the cucullus is relatively broader than that of that of E. cirripalea, E. colorado, and E. sidus. The digitus is longer and more attenuated in E. cirripalea and E. schweitzeri than that expressed in E. colorado and E. sidus.
Description of Adult
Head. Setae orange to reddish brown often darkened or hoary apically. Some scales forked apically on head but not on antennae. Lamellar, tan to grey scales on dorsal surface of antennae. Thorax. Thorax densely covered with hairlike setae. Scales forming middorsal ridge. Thorax concolorous with wings, ranging from orange brown to purple or reddish brown. Apices of setae often deeply forked and hoary. Forewing (Figs 1–3). Orange to reddish brown with hoary, gray violet color, often with purple luster especially basad of antemedial line and over outer third of wing. Sinuous postmedial line bulging outward beyond cell with dark dentate projections sometimes present on R5, M2, M3—Cu2. Reniform spot small, orange, yellow, or white, often flanked by upper and lower satellite spots; orbicular almost always absent, but sometimes faintly discernible; claviform obsolete. Reared and freshly flown individuals have three bands of violaceous scales: inside antemedial, outside postmedial, and inside of dark crenulated adterminal line. Outer margin of wing convex, rounded, modestly crenulated. Fringe concolorous or slightly paler than FW ground color. Indistinct medial line of darker scales, jogging outward to reniform spot through cell. Hindwing (Figs 1–3). Uniformly fuscous with only slightly pinker fringe, often concave between M1 and M3. Crescent-shaped reniform spot weakly expressed. Ventral surface of hindwing with poorly differentiated discal spot and postmedial bands (relative to many Eupsilia). Abdomen. Fuscous, more or less concolorous with hindwings. Male genitalia (Figs 14, 15). Very similar to those of E. cirripalea and E. sidus. Uncas strongly hooked, drawn into spine; gnathos subquadrangular, juxta pentagonal with truncated dorsal margin and lacking medial conical projection (of E. cirripalea); valve elongate, roughly four times as long as wide. Prominent corona, with outer margin slightly curved, running nearly in same axis as lower valve margin, bearing 20–31 bristles. Digitus very long, projecting past lower margin of valve; cavity between cucullus and corona shallow. Inflated vesica with six basal pouches (in two groups of three); near midlength, rosette of ca. 25–35 thickened setae, with finely attenuated, twisted apices. Female (Fig. 16). Bursa copulatrix with four signa, two of which are rudimentary, i.e., only slightly differentiated, from the walls of the bursa; all four extend nearly full length; mesal surface of signa serrulate. Numerous concentric creases of bursa microserrulate; caudal end of bursa copulatrix with two pouches. Ductus bursae more strongly sclerotized about antrum opening to pouch at midlength.
Diagnosis of Living Final Instar (Figs 26, 28). Thoracic venter from T1–A1 bright cherry red, contrasting with following abdominal segments, which are pale and largely unpigmented. Spiracular stripe thin, broken or absent between T1 and A1; thickened on A7 and A8. Middorsal and subdorsal stripes, white, thin, sometimes interrupted; supraspiracular stripe mostly obliterated. Ground color of dorsum red brown over thorax but giving way to frosty grey or brown over abdomen (this same area becoming brown in the mature larvae and prepupa); laterally, brown below subdorsal and above supraspiracular stripe; contrasting with reddish-brown, maroon, or wine supraspiracular region. Middle Instars (Figs 23–25) with more gray over abdomen; red pigments in supraspiracular more evident, stripes white, and abdominal subventer and venter mostly unpigmented.
Distribution. Because E. schweitzeri represents a cryptic species, its range is poorly circumscribed. We have examined specimens from southeastern Ontario, Massachusetts, and New York, southward to Pennsylvania and New Jersey, though the moth becomes scarce in the southern parts of New Jersey. Its range extends farther south in mountainous areas; confirmed records extend through mountainous areas of North Carolina (Bo Sullivan unpubl. data) into northern Georgia (Adams 2015). What appears to be the same taxon occurs as far west as Missouri according to Dale Schweitzer (personal communication).
Barcoding. Of 154 barcoded Eupsilia from North America, with reads >500 base pairs, E. schweitzeri clusters as a single entity (n=10) outside of E. cirripaleamorrisoni-sidus-tristigmata-vinulenta assemblage (Don Lafontaine unpubl. data). Thus, there is little suggestion from CO1 that E. schweitzeri is especially close to any of the three species with which it has been confused in collections, i.e., E. cirripalea, E. sidus, or even E. vinulenta. (Known specimens of E. colorado are too old for barcoding—see below.)
Key to Adults of Eastern North American Eupsilia . Our key extends the effort of Forbes (1954). All characters refer to forewing characters unless otherwise noted. Species identifications in Eupsilia can be challenging. In some species and forms, no single character may work to assure certain identification, especially in flown or overwintered individuals. Likewise both male and female genitalia are frustratingly uniform across the genus.
1. FW postmedial line not crenulate; basal, antemedial, and postmedial lines of pale scales; outer side of reniform often marked by crenulate tan line 2
FW postmedial line crenulate; basal, antemedial, and postmedial lines mostly of dark scales; outer side of reniform not marked by crenulate tan line 3
2. Ground color gray without reddish tints; basal third of FW gray; excelsior scales absent devia
Ground color with orange or reddish tints; basal third of FW concolorous with rest of wing; modified scales sometimes present basad to postmedial line morrisoni
3. Reniform with dark spot at its base; orbicular often distinct; basal line usually including a dark spot; often with dark subapical spot along costa tristigmata
Reniform without dark spot on its lower edge; orbicular obsolete; basal line without dark spot; without dark subapical spot along costa 4
4. FW scales distal to postmedial line lamellar (scales below the reniform are shown in Fig. 22) vinulenta
Many FW scales distal to postmedial line of excelsior type, i.e. deeply cleft with outer spines curled and overlapping, and scale edges often rolled upwards (as in Figs 17, 19, 20) 5
5. Male juxta with central conical projection; female bursa with two signa; outer margin of forewing crenulated; ground color of FW more uniformly brown (and less reddened) than those that follow) cirripalea
Male juxta without central conical projection; female bursa with four signa (two of which are weakly expressed); outer margin of forewing crenulated or smooth; ground color of FW more reddish brown and sometimes with violet luster 6
6. Three bands of violaceous scales (inside antemedial, outside postmedial, and inside of dark crenulated adterminal line) (in fresh individuals); FW color more brown than red; outer margin modestly crenulated; FW scales distal to postmedial often with numerous lamellar scales; valve robust, 4x longer then broad; digitus longer and more attenuated than that of sidus (cf. Figs 10, 14) schweitzeri
Lacking three bands of violaceous scales; FW color more reddish than brown; outer margin crenulate; FW scales distal to postmedial dominated by excelsior scales; valve slender, 5x longer than broad; digitus shorter and less attenuated than that of schweitzeri (cf. Figs 10, 14) sidus
Key to Last Instar Northeastern Eupsilia Larvae
Note: Our key works best for fifth and submature sixth instars. Fully fed last instars and prepupae lose diagnostic coloration and become muddied in color to the extent that coloration characters become unreliable. Likewise, early instars of northeastern Eupsilia can be so similar that identifications based on this key, or our images, should be considered provisional. For example, the diagnostic red thoracic venter of E. schweitzeri (in late instars) is seen in early instars of other Eupsilia. We also caution that some larval characters given in previous works, i.e., the absence of subdorsal white stripes on T1 in E. tristigmata, are not deemed reliable. All seven eastern Eupsilia are figured in Wagner et al. (2011).
1. Venter green (including thorax), mostly unpigmented (sometimes pinkish anteriorly) 2
Venter flushed with red or pink 4
2. Thick white spiracular stripe on A1–8; mottled green to brown dorsum; supraspiracular areas black below faint white subdorsal line; subventer unpigmented E. devia
Thin white spiracular stripe often inconspicuous; dorsum concolorous or slightly differentiated in color from supraspiracular area; subventer pigmented or unpigmented 3
3. Dorsum and supraspiracular areas often somewhat contrasting; supraspiracular area usually with deep to smoky wine tints; middorsal and subdorsal lines thin and variously present, former more white in color; ground color usually some shade of brown E. morrisoni
Dorsum and supraspiracular areas usually concolorous; both dorsum and supraspiracular area usually with deep to smoky wine tints; middorsal and subdorsal lines thin, often obliterated; ground color typically velvety black to purplish brown E. vinulenta
4. Pink to red on thoracic and abdominal venter (Fig. 29), dorsum dark mottled gray to an olive green brown; supraspiracular area wine colored E. sidus
Pink to red confined to thoracic and A1 venter 5
5. Abdominal subventer usually pink to red above prolegs (Fig. 30) E. tristigmata
Abdominal subventer unpigmented above prolegs (Figs 27, 28) 6
Thoracic venter pink (Fig. 27); supraspiracular area reddish brown to pink brown E. cirripalea
Results and Discussion
It is easy to understand why this common species, occurring in one of the most densely populated and well-collected regions of North America, went undescribed for so long. Adults are phenotypically difficult to distinguish from those of E. cirripalea and E. sidus. The genitalia of all seven northeastern Eupsilia species are similar, even between species that would easily be separated by wing pattern, coloration, and scale shape, e.g., E. vinulenta and E. schweitzeri. Where E. cirripalea, E. sidus, and E. schweitzeri co-occur, for some adult phenotypes, the only certain means of identification are genital and larval characters. Female genitalia are indistinguishable from those of E. sidus.
Biology and Distribution.
Female Eupsilia lay eggs singly on twigs and buds in the early spring. Early instars of Eupsilia are believed to feed on a wide taxonomic range of woody plants in the spring, fashioning loose leaf shelters (Fig. 24) or entering those made by other Lepidoptera (Wagner et al. 2011). By the fourth or fifth instar Eupsilia abandon their shelters. Where they go is a mystery. In two decades of caterpillar collecting in Connecticut, we have never found a late instar on foliage during the day. Their dark coloration is suggestive that the larvae feed at night. It is our suspicion that the late instars are broadly polyphagous on woody and non-woody plants, and perhaps other organic matter on or near the ground (Wagner et al. 2011). We once found a last instar Eupsilia apparently feeding on dog food in a basement bulkhead. It is unclear if larvae ascend trees at night to feed, as is commonly the case with many climbing cutworms, e. g., Metaxaglaea Franclemont and related genera. The European Eupsilia transversa is recorded from forbs (e.g., dandelion), and a wide range of woody plants; additionally, it is reported to feed on the caterpillars of other Lepidoptera (Manley 2008). Eupsilia larvae mature in late spring, and then burrow underground and form a cell where they enter diapause for two to three months, before pupating (Wagner et al. 2011). Adults begin issuing in September in Connecticut.
Head capsule measurements for E. schweitzeri indicate that there are six larval instars (Fig. 31): 1st: = 0.3 mm (n=6, 0.3 mm); 2nd: = 0.5 mm (n=10, 0.4–0.5 mm); 3rd: = 0.8 mm (n=7, 0.8–0.9 mm); 4th: = 1.4 mm (n=8, 1.2–1.5 mm); 5th: = 2.2 mm (n=8, 2.2 mm); 6th: = 3.2 mm (n=16, 3.1–3.3 mm). The head capsule width ratios between successive instars were generally consistent with those observed by Dyar, which ranged from 1.3–1.7 (Dyar 1890, Berg and Merritt 2009). Observed ratios in E. schweitzeri ranged from 1.4 between the last two instars and 1.8 between the 3rd and 4th instars, with a mean, median, and mode of 1.6 across the six instars.
Ex ova lots have been reared on Carya glabra, Juglans cinerea, Malus sylvestris, Prunus serotina, and Quercus ilicifolia. Our wild larvae of E. schweitzeri, have been collected as middle instars in southern New England from Carpinus caroliniana, Cirsium sp., Hamamelis virginiana, Nyssa sylvatica, Prunus serotina, Quercus spp., and Vaccinium spp. We suspect that E. schweitzeri is a generalist on woody shrubs and trees in early and middle instars, but becomes even more generalized in diet breadth in late instars, consuming a wide variety of plants (e.g., Cirsium) and perhaps even other types of organic matter.
Because all Eupsilia (across both the Nearctic and Palearctic) are similar in size, we think it likely that head width measurements of any wild-collected Eupsilia larva would generally conform to those given in Figure 31. We would be interested to learn of wild larval collections of 4th, 5th, and 6th instars, and upon what substrates they were observed feeding.
Although all seven northeastern Eupsilia may cooccur (e.g., in southern Connecticut), E. sidus and E. schweitzeri are largely allopatric over much of their range. E. sidus is usually a denizen of “dry oak forests, savannas, woodlands, ridgetops, and barrens from Wisconsin, southern Ontario, and central New Hampshire south to northern Georgia and Texas” (Wagner et al. 2011) (this range may conflate records of E. schweitzeri, e.g., records of E. sidus from Ontario may in fact refer to the heretofore unrecognized E. schweitzeri). E. schweitzeri is more ecologically generalized. It is abundant in both mesic and dry oak woodlands, but also occurs in bottomlands, along the edges of wetlands, in upland woodlands and forests, ridgetops, and other forested communities. In most habitats it is only outnumbered by E. vinulenta and E. morrisoni.
Wing Scales. Eupsilia have derived forewing scales. We confirm the presence of three scale types: lamellar, excelsior, and hooked (Figs 17–22). Lamellar scales, common to Lepidoptera (Scoble 1992, Kristensen & Simonsen 2003), predominate in E. devia, E. morrisoni, and E. vinulenta (Fig. 22) and in the former species, virtually all scales are of the lamellar type. In addition, the lamellar scales of E. devia are wider than those found in the six other northeastern species. Excelsior scales are those where the outer teeth of forewing scales tend to be elongated and recurved; in many cases, scale edges are rolled upward to the extent that they almost enclose the dorsal scale surface. Forbes (1954) used the term “shredded wheat” effect to describe the wing surface appearance resulting from the overlap and interlocking of the modified scales.
Excelsior scales occur in E. cirripalea, E. morrisoni, E. schweitzeri, E. sidus, and E. tristigmata, (Figs. 17–21), and in lower frequency and with an intermediate degree of recurvature of the outer teeth in E. colorado and E. vinulenta. E. tristigmata best represents the scales we describe as hooked; i.e., just the outer (two) teeth are recurved, sometimes as much as 180º (Fig. 21). The outer teeth also tend to be much darker in coloration in E. tristigmata.
Eupsilia walkeri (Grote, 1864). With the possibility that Eupsilia walkeri might refer to our new species, we compared our type series to Grote's description, which he based on a single specimen from Philadelphia. The type of E. walkeri is believed to be lost (Franclemont 1952). Grote's description of E. walkeri makes no mention of purple or related violaceous colors (yet he used purplish twice as a descriptor in his description of E. vinulenta, which appeared in the same publication, immediately following the account for E. walkeri). Violaceous scaling on the forewings is among the most reliable features for the recognition of E. schweitzeri adults, and is usually present basad to the antemedial line. Grote's characterization of the terminal line in E. walkeri as “very clearly defined, semi-lunulate,” further eliminates E. schweitzeri as a possibility (as well as E. cirripalea or E. sidus and others, a point we return to below). Grote uses ochrecous to describe the forewing color of E. walkeri and “rich dull red” to describe the forewing of E. vinulenta. Ironically, the only Eupsilia in the Northeast that is accurately characterized as ochraceous are the pale (overwintered) forms of E. vinulenta (Fig. 9), the most phenotypically variable Nearctic Eupsilia. Both E. schweitzeri and E. sidus are a richer red than Grote's rendering of E. walkeri.
In his 1952 treatment of Eupsilia, Franclemont regarded E. walkeri (Grote) to be a synonym of E. sidus:
“ As identified here, walkeri Grote is a synonym, and judging from the figure and description it was based on a worn hibernated specimen. Grote's type is also lost, but was probably also from the vicinity of Philadelphia since it was in the collection of the Entomological Society of Philadelphia.”
Forbes (1954) and subsequent workers have followed Franclemont. However, based on Grote's original description, remarks made above, and other arguments, we disagree with Franclemont's decision, and treat E. walkeri as synonym of E. vinulenta (Grote). As noted above, Grote states that E. walkeri possessed a clearly defined, semi-lunulate terminal line. Such is absent not only from E. schweitzeri but also E. cirripalea and E. sidus, and is commonly present in E. vinulenta. Grote's description of E. walkeri states: “transverse anterior [antemedial] line distinct, darker shaded, straight, composed of two narrow lines with a central light shade.” Yet in E. sidus the antemedial line is poorly differentiated, especially relative to that of E. vinulenta. In addition to the character evidence, we note that E. vinulenta is among the most common Eupsilia in eastern Pennsylvania. E. sidus is an oak barrens species, at least at the latitude of Philadelphia, that we associate with scrub oak and low-bush blueberry communities, a habitat absent from the vicinity of present-day Philadelphia. In sum, we are convinced that Grote based his description of E. walkeri on a flown (perhaps overwintered) individual of E. vinulenta, and could not have been either E. schweitzeri or E. sidus.
Scopelsoma colorado (Smith, 1903), rev. stat. In his 1952 treatment of Eupsilia, Franclemont synonymized Scopelsoma colorado with E. sidus. The species is known only from the type series evidently collected by William H. Barnes in March and April, 1893–1895, in Glenwood Springs, Colorado. We were able to locate and examine 1 (of the 2 formerly present) specimens in the AMNH, including the lectotype male and its associated genitalic slide; 13 specimens in the USNM; and 2 specimens in the CNC. After examining the above, as well as three additional male genitalic dissections (Franclemont #7574, USNM 38903, USNM 118685) and three female dissections (CNC 16801, USNM 38901 [bearing the notation: (From spem. labeled “♂ cotype”) [sic]], USNM 38902), we conclude that Scopelsoma colorado is a valid species. The ground color ranges from tan or ochraceous (most) to reddish brown (few); violet scales are absent and any hint of red or orange typical of E. sidus is subdued; the antemedial line in E. sidus and to a lesser extent E. schweitzeri is nearly absent in E. colorado (Fig. 4). Most notably the reniform and associated satellite spots common to E. schweitzeri, E. sidus, and E. vinulenta are vague and illdefined in E. colorado save for the dark scales along the lower edge of the reniform spot (Figs 1–9). The forewing margin is less crenulate than that of E. schweitzeri or E. cirripalea; and especially that of E. vinulenta. The forewing scales are intermediate between lamellar and excelsior, and most similar in character and distribution to E. vinulenta. They lack the “shredded wheat” aspect that we (and Forbes) would attribute to E. schweitzeri, E. sidus, and E. cirripalea. Forewing scales in E. colorado distad to the postmedial line are lamellar, whereas great numbers of excelsior scales are present in this area in E. cirripalea, E. schweitzeri, and E. sidus. The male genitalia of E. colorado, based on dissections from Franclemont (#7574), the USNM material (#38,903), and J. Donald Lafontaine (#118685), are close to those of both E. sidus and E. schweitzeri, but with important differences. The coronal spines are stouter and fewer in number (14–17) than in other North American Eupsilia (Figs 10, 12, 14). The setal rosette at midlength on the vesica (Fig. 13) is like that of E. schweitzeri (Fig. 15) with 25–30 fine spines, and but differs from that of E. sidus (Fig. 11), whose vesica bears a run of fewer (ca. 20) stouter setae, and these are drawn out along the vesica, rather than pulled into a single rosette. Nor does E. colorado have the six small basal pouches, arranged in two groups, as seen in E. sidus and E. schweitzeri. In our preparations, the most basal set of pouches are much enlarged and give the basal portion of the vesica considerable girth. We note that the lectotype preparation from the AMNH, made by Smith, is badly damaged and distorted.
Eupsilia sidus (Guenée, 1852). The type of E. sidus is missing (Franclemont 1952). Given the difficulty of adult identifications, even with dissection, and the taxonomic confusion within the genus, to promote nomenclatural stability we anchor the concept of the name as understood by Franclemont (1952), Forbes (1954), current on-line resources, and institutional collections, by hereby designating a neotype of E. sidus from New Jersey, the presumed type locality. (According to Franclemont (1952) “From what we know about the source of Boisduval's material described by Guenée, it seems safe to assume that the type of sidus was collected by Say in New Jersey, given to LeConte and sent by LeConte to Boisduval.”) The identity of the neotype (Fig. 5) was confirmed by larval characters, the male was dissected, and a leg was submitted for DNA barcoding. Data on the neotype labels are as follows: New Jersey, Cumberland Co., Millville, Poplar Drive f coll. at bait III.11.2010, Dale F. Schweitzer // bred ex ova on Prunus serotina at Windham, CT, by Eric L. Quinter, emerge[d]: IX.26-X.15.2010 // Eupsila sidus (Guenée), Det. Eric L. Quinter // CNC # ♂ 17072 // Specimen ID CNCLEP 00116372 // Barcode of Life Project // Leg removed, DNA extracted // Photographed J. D. Gill. The neotype has been deposited at the CNC, Ottawa, Ontario, Canada.
Eupsilia is a difficult taxon given the uniformity of the genitalia across species. Likely the new species would have gone undiscovered until the advent of barcoding had not the larval characters revealed that the name E. sidus was being applied to two different species. Given the conservative nature of phenotypic evolution in adult Eupsilia, it would not be surprising to learn that still other cryptic species of Eupsilia remain to be recognized and described from North America. Barcodes are suggestive that E. tristigmata represents more than a single entity and perhaps as many as three (Don Lafontaine pers comm.).
Eupsilia colorado has not been collected since 1895—and awaits rediscovery. Efforts to barcode specimens were unsuccessful. The type locality, Glenwood Springs, is noteworthy for its exceptional complex of moths that occur sparingly or nowhere else on the west slope of the Rockies (Don Lafontaine in litt, Terhune Dickel pers. comm.). Other notable lepidopterans known from the area include Acronicta exempta Dyar, 1922, Heterocampa rufinans (Dyar, 1921), Hyparpax venus Neumoegen, 1892, Orthosia flaviannula (Smith, 1899), and Phoberia ingenua (Walker, 1858). What is special about the conditions in the vicinity of Glenwood Springs remains a mystery; perhaps they relate to a unique paleoclimatological history, specialized edaphic factors, or unique aspects of its flora. Many of the above are oak feeders, and Quercus is a known host for many eastern Eupsilia (Wagner et al. 2011). Eupsilia colorado flies in March and April, when few moth collectors are afoot in the Rockies and weather conditions are especially challenging for nocturnal collecting. Larval sampling, especially on oak in the spring, might prove to be the easiest means to survey for the moth.
John G. Franclemont prepared 82 genitalic dissections of Eupsilia that were especially helpful over the course of our effort; his preparations were made available by Jason Dombroskie and Cornell University. Tim McCabe also made his genitalic preparations available for study. Eric Quinter supplied reared material of E. schweitzeri and E. sidus from which the type material was selected. Eric also guided us to relevant literature, specimens in the AMNH, and supplied considerable unpublished information regarding Eupsilia. Dale Schweitzer first alerted DLW to the existence of the new species and supplied some of the life history data reported here. Paul Goldstein (USNM) arranged for the loan of specimens of Eupsilia (Scopelsoma colorado). David Grimaldi provided access to the AMNH collection and the lectotype of Scopelsoma colorado. Don Lafontaine and Jocelyn Gill prepared key genitalic preparations and associated images for our manuscript. Bo Sullivan and Don Lafontaine shared relevant barcode data. Annette Evans and Mark Smith took the high resolution images of the forewing scales used in Figs 17–22. Don Lafontaine and Lawrence Gall made helpful suggestions on the submitted version of the manuscript that significantly improved the text. This study was funded in part by USFS Co-op Agreement 14-CA-11420004-138 and an award from the Richard P. Garmany Fund (Hartford Foundation) to DLW.