A revision of Admetovis Grote, with the description of a new species from western North America (Noctuidae, Noctuinae, Hadenini)

Abstract The genus Admetovis Grote is revised. Admetovisicarussp. n. is described from the mountains of western North America. A lectotype of Admetovisoxymorus Grote is designated. Illustrations of the adults, male and female genitalia, and distribution maps are presented, together with an identification key. The classification of the genus is reviewed resulting in its reassignment to the tribe Hadenini from Orthosiini.


Introduction
Admetovis Grote is a small genus of three western North American noctuid moths. The species are similar superficially, with a distinctive flame-like mark on the distal third of the otherwise gray forewing. Grote named the genus and first species in 1873. Two species, Admetovis oxymorus Grote and Admetovis similaris Dyar, have been known filling that contributes to palest portion of "flame" mark. Hindwing: Margin between M1 and M3 straight or concave; ground color pure white or slightly mottled light tan to brownish gray; veins, discal spot, and terminal line light to dark gray; fringe white to light gray with darker stripes, paler than ground. Abdomen. Male with or without basal coremata on segment I and pockets on ventrolateral segment III; when present weak, comprised of few filaments attached directly to base with pockets shallow, or strong, paint brush-like, filaments arising from rod with bulblike base, sometimes with very small medial accessory brushes, with pockets deep. Dorsal tufts on segments I-III. Female sternite VII sclerotized on distal half, with posterior median cleft 0.33-0.58 × segment length, blunt posterolateral lobes on side of cleft cover ostium bursae and part of ventral segment VIII. Male genitalia: Uncus relatively short, curved ventrad, base cylindrical, distal half widened and flattened dorso-ventrally, slightly concave dorsally with flange-like raised edges, tapering to truncate apex with small ventral spine; dorsal edges and ventral apex with numerous short setae. Juxta broadly shield shaped, wider than tall, smooth (A. similaris) or with median thorn-like spine directed posterodorsad (2 species). Valve length ~ 5 × width, S-shaped with slight curve dorsad near mid-point and 90° lateral bend distal to clasper at ⅔ distance from base to apex, tapered evenly from broad base to end of clasper, narrowing abruptly at bend to thin neck of cucullus, ventral margin of mid valve thickened with short blunt lateral projection ventral to distal clasper; sacculus length 0.4-0.5 × valve, sclerotized strongly, extending to or slightly above dorsal valve, distal costal margin near dorsal attachment of valve humped dorsad or anvil shaped, covered densely with minute setae; clasper strongly sclerotized, a bowl-like concavity dorsal and distal to sacculus, distal margin a narrow flange-like rim extended to blunt triangular dorsal and broad convex ventral projections, dorsal projection longer than (two species) or similar (one species) to ventral projection; digitus absent; cucullus triangular, 0.6-1.0 × mid-valve width, with slightly rounded margin bearing simple corona of 15-25 claw-like setae. Phallus tubular, length 6-7 × width. Vesica length (including right and left limbs) 1.0-1.5 × phallus, bent slightly ventrad at base then divided into nearly equal limbs directed 90° right and left to form with phallus a "T;" right limb comprised of distal vesica, tapered to ductus ejaculatorius at apex, with subapical broad short ventral diverticulum; left limb variable between species, tapering to conical or thin apical cornutus, bearing an additional thin acute cornutus at dorsal base (absent or diminutive in some specimens of all species) and anteriorly-directed broad conical subapical diverticulum (lacking in A. icarus). Female genitalia: Papilla analis asymmetrically conical, apex near dorsum, length 1-1.5 × width, covered sparsely on base and mid-portion by thin hair-like setae, more densely on apex by short setae. Abdominal segment VIII slightly longer than wide, venter length 2 × dorsum, covered sparsely by short hair-like setae; anterior apophyses length 0.57-0.77 × abdominal segment VIII; posterior apophyses length 2.75-3.35 × anterior apophyses. Ductus bursae length 1.14-1.32 × abdominal segment VIII, strongly sclerotized, flattened dorsoventrally, length ~ 3 × width, ventral surface smooth, dorsum rugose; ostium bursae simple. Corpus bursae length 3.9-4.6 × abdominal segment VIII, elongate, gourd shaped, anterior portion curved dorsad and leftward, membranous ovate anterior end widest, width 0.35-0.43 × bursa length, 3-4 longitudinal irregular string-of-beads signa evenly spaced on surface (left lateral signum absent in two species); appendix bursae projected ventrad and leftward from broad origin at junction with posterior corpus bursae at ductus bursae, length 0.12-0.25 × total corpus bursae length, sclerotized lightly, apex blunt, ductus seminalis at dorsum anterior to apex.
Distribution and ecology. Admetovis species occur in western North America, from the Rocky Mountains and Arizona-Mexico border west to the Pacific Coast and north to southern British Columbia. They undoubtedly occur in Mexico but the distribution there is unknown.
The flight period of adults is from early spring (February or March) to as late as August depending on the species and locality. Based on limited information for Admetovis oxymorus the larvae feed on woody shrubs and are most likely climbing cutworms (Mc-Farland 1975). All species in the genus are nocturnal and are attracted readily to light.
Discussion. Admetovis was historically classified in the subfamily Hadeninae, where it was placed since the early twentieth century (Hampson 1905, McDunnough 1938, Hodges et al. 1982. It was reassigned recently to the tribe Noctuinae: Orthosiini (Lafontaine and Schmidt 2010) stemming from changes in the composition of the Noctuidae (Fibiger and Lafontaine 2005). Although the higher classification, including the recognition of Orthosiini as a tribe, is fairly recent (Fibiger and Lafontaine 2005) the relationship of Admetovis to other genera in this tribe is not obvious and had not been recognized widely. More recent molecular approaches to noctuid classification based on mtDNA barcode sequence data (Zahiri et al. 2017) support the current classification of North American Orthosiini in that nearly all included genera cluster together, but with the exception of Admetovis -this genus instead groups with genera in the Hadenini, which Godfrey also indicated as being the most closely related group to the orthosiine genera. This warrants a re-examination of Godfrey's (1972) interpretation of larval hypopharynx structure, which forms the basis of the Orthosiini as currently defined. Godfrey (1972) included nine genera in the Orthosia group ("Group 8"), noting that two other genus-groups were likely closely related based on the morphology of the hypopharynx, namely the spining pattern and especially the transverse cleft, the latter shared across the three groups. Godfrey's three "transverse-cleft" groups comprise the Anarta-group, Polia-group and Orthosia-group, the first two now combined in the tribe Hadenini and the third comprising the Orthosiini. Although Godfrey separated the Orthosiini and Hadenini morphologically by two characters (lack of setae above the spinneret and larger spines on the proximolateral hypopharynx in Orthosiini), exceptions occur in both tribes (Godfrey 1972). Admetovis differs in the shape of the spinneret and the finely granular (vs. smooth) larval integument from all other Orthosiini, but these characters appear to be autapomorphic because they also do not occur in the Hadenini.
Importantly, Admetovis is the sole constituent of Godfrey's Group 8 where adults do not emerge in early spring. Most species in the Orthosiini emerge very early in the season, and some, such as Orthosia praeses (Grote) and Egira hiemalis (Grote), are the first non-overwintering moths to fly in late winter. While A. similaris can be found as early as February in the deserts of the Southwest, most Admetovis differ from other orthosiines in that they fly later in the year from late spring through summer.
Finally, the male valves of Admetovis, S-shaped with triangular cuculli, are more similar to those of most hadenine moths (e.g., as shown for European species in Hacker et al. 2002) than they are to orthosiines (shown similarly in Ronkay et. al. 2001), as are the everted vesicae. Females of both tribes are rather simple in genital structure and as a result are not highly diagnostic.
The weight of evidence (barcodes, biology, and adult genitalia structure versus somewhat equivocal larva hypopharyngeal structure) suggests that Admetovis is better classified in the Hadenini rather than the Orthosiini. A definitive phylogeny of the Noctuinae tribes, using multiple molecular and morphological markers, is still needed. As such, we recommend Admetovis as a fertile subject for further investigation. LGC, OSAC, TM. Three specimens from Sandon, British Columbia (J. W. Cockle) at the CNC were also examined, but are excluded from the type series because they are worn.

Key to species of Admetovis (adults)
Diagnosis. Admetovis icarus is the only species in the genus with a dusky hindwing with a straight outer margin distal to the cell between veins M1 and M3. Admetovis icarus is distinguished easily from A. similaris by the pure white hindwing of the latter species. It is only subtly different from A. oxymorus in habitus and can easily be confused with it, especially in the Pacific Northwest where both species occur. While the hindwing margin shape is the most reliable character for separating these species short of dissection, there are subtle differences in color. Admetovis icarus tends be more mottled on the forewing with a darker "flame-mark," and its hindwing is consistently darker brownish gray than that of A. oxymorus. The forewing orbicular stigma of A. icarus is often more conspicuous than in either other species, with pale filling outlining a dark central ocellus.
Structurally, males of A. icarus differ from both other species by the presence of weak basal abdominal coremata lacking a strong core; A. oxymorus has fully developed coremata with stout rods and A. similaris lacks them altogether. Admetovis icarus and A. oxymorus both differ from A. similaris in having a median spine on the juxta. In the valves, the setose dorsal protuberance of the sacculus is triangular to anvil shaped in A. icarus, convex in A. oxymorus, the dorsal process of the clasper is short and triangular in A. icarus, longer and curved in A. oxymorus, and the cucullus is relatively small and rounded in A. icarus, larger and triangular in A. oxymorus. The relative sizes of the cuculli are apparent readily if the two species are compared after the scales are removed with a brush. The vesica of the phallus of A. icarus is the simplest in the genus, with a relatively short left limb lacking a subapical diverticulum.
Females of A. icarus, like those of A. oxymorus, differ from those of A. similaris in having three rather than four signa on the corpus bursae. They can be differentiated from those of A. oxymorus by the shape of the corpus bursae, narrow with a small bulbous anterior portion in A. icarus; curved and broader with a bulbous anterior end in A. oxymorus. This difference is quantified in the Key as a ratio between width of the corpus bursae and the length of the appendix bursae.
The barcode of A. icarus (BOLD:AAD7456) differs from both other species by about 3.5 %. There is no intraspecific variation in three samples from Washington and Utah.
Description. Head -Structure of male and female antennae, eye, palpus, and haustellum as for genus. Dorsal antenna tan with scattered gray scales. Scape off white. Labial palpus with nearly equal mixture of pale tan and dark gray scales. Frons tan, gray periph-erally. Dorsal head scales white-tipped gray. Thorax -Dorsum as for genus; venter scales long, hair-like, white-tipped dark gray. Legs as for genus; tarsal segments dark gray banded distally with off-white. Wings: Forewing: length 16-17 mm (males), 17 mm (females), length 2.3-2.4 × width, outer margin scalloped weakly; ground lead gray, most mottled in genus due to more whitish scales, especially near base; medial area distal to reniform stigma and postmedial area whitish tan to tan, darkest distally; basal, antemedial, and medial lines as for genus; postmedial line dark gray, conspicuous for genus, double with strong inner and incomplete weak outer components, scalloped strongly with dark gray distal extensions on veins; postmedial white with preceding red-brown and black shade; terminal line black, interrupted at veins; fringe gray with light tan base, pale checkering at veins; claviform stigma dark gray, filling light gray; orbicular stigma nearly round, double, outer component darkest, filling light gray, whitish gray, or pale tan, central ocellus gray or mixed gray and brown; reniform stigma moderately large, kidney shaped with slightly larger posterior end, dark gray, incomplete at posterior end, filling of medial ⅓ light tan and distal ⅔ whitish tan or gray. Hindwing: margin straight between M1 and M3; powdery dusky gray brown with slight brassy sheen, discal spot and patchy marginal shade slightly darker with lighter patches near outer margin. Abdomen. Male coremata very weak, a few filaments arising from button-like base, lacking central stalk; pockets on ventrolateral segment III small, shallow. Female sternite VII posterior median notch 0.36 × length of the segment. Male genitalia: Uncus as for genus, wide distal portion only slightly tapered to blunt tip. Juxta height 1.5 × width, median spine present. Valve as for genus, length 5 × width, ventral mid-valve projection short, blunt; sacculus length ⅔ × valve, width 1 × valve, setose costal lobe asymmetric, anvil shaped with triangular pointed tip; clasper with short triangular dorsal and broad-based convex ventral processes of similar length, small for genus; cucullus relatively small, ~ 0.6 × valve width, rounded, corona 15-20 setae. Phallus tubular, length 6.8 × width. Vesica length 1.0 × phallus; diverticulum of right arm broad based, short; left arm tapered to straight spike-like apical cornutus, basal cornutus present or absent, diverticula absent. Female genitalia: papilla analis length 1.5 × width, longer and more pointed than in other Admetovis. Segment VIII as for genus; posterior apophysis length 0.75 × segment VIII; anterior apophysis length 2.75 × posterior apophyses. Ductus bursae relatively smooth, rugae limited to near junction with corpus bursae. Corpus bursae length 4.6 × abdominal segment VIII, elongate, narrow, curved weakly, bulbous anterior portion relatively small, diameter ~ 0.4 × total corpus bursae length; signa on dorsal, right lateral, and ventral sides; appendix bursae simple sac-like posterior extension of corpus bursae, length 0.16 × corpus bursae.
Geographic variation. This species is fairly uniform throughout most of its range. The population on the Wasatch Plateau of central Utah is paler and more mottled than those from elsewhere.
Etymology. The species epithet refers to Icarus, son of Daedalus in Greek mythology. Icarus used wings that his father had made to escape from the island of Crete but flew too close to the sun, thereby falling to his death in the sea. The flame marking on the distal forewing and high elevation habitat of this moth bring to mind his story. It is a noun in the genitive in apposition to the generic name.

Distribution and ecology. Admetovis icarus occurs in the mountains of western
North America, mostly in the Rocky Mountain region. Records extend from central Utah and central Colorado to the Selkirk Mountains of southeastern British Columbia, including a record from northeastern Oregon. Farther west there are scattered records from the Okanagan region of south-central British Columbia and Chelan and Kittitas counties in the northern Cascade Range, Washington. It replaces A. oxymorus in Utah and Colorado but is partially sympatric with it in the Pacific Northwest.
Admetovis icarus has been collected almost exclusively in high-elevation forests near tree line, although the habitat in Kaslo and Nelson, British Columbia is transition zone forest. Collection dates are from late June to early August.

Admetovis oxymorus
Type material. Admetovis oxymorus was described from two syntypes, one each from Sierra Nevada and Rocky Mountains (Grote 1873), with an illustration of the California type on plate 4, figure 5. The depicted California female is clearly identifiable as the species that is widespread along the West Coast, an important distinction since based on current known distributions the other type specimen from the Rocky Mountains is most likely A. icarus.
We found neither type specimen in major collections known to contain Grote type specimens, including AMNH and NHML, and both specimens are most likely lost. This conclusion is supported by the fact that Poole (1989), in the Lepidopterorum Catalogus series, appears to not have examined the syntypes or known their whereabouts. He lists the type material noncommittally as "Type(s)" and cites "AMNH London [sic]" as the repository collection. In order to fix the identity of the name, we hereby designate a female specimen in the CNC labeled "CA, Sierra Co., 2 mi. E. Bassetts, 5300' [1615 m], Hwy. 49, SNFC-SF State U., 4-7.Jul.2007, P. A. and E. Opler" as Neotype with a red label designating it as such.
Diagnosis. This species and A. icarus have dorsal hindwings with gray or tan ground color and are distinguished easily from A. similaris that has a pure white hindwing with white or thin gray veins. Admetovis oxymorus is best distinguished from A. icarus by the shape of the hindwing. The margin between veins M1 and M3 is slightly concave in A. oxymorus but straight in A. icarus.
Admetovis oxymorus is the only species in the genus with fully developed male coremata; the coremata of the others are either vestigial or absent completely. The cucullus of the male valve of A. oxymorus is broadest in the genus, more than 2 × width The barcode of A. oxymorus (BOLD:AAD7455) differs from those of the other Admetovis species by slightly more than 3.5 %. Intraspecies variation within A. oxymorus is approximately 0.2 % (n = 7; British Columbia, California, Oregon).
Distribution and ecology. Admetovis oxymorus occurs in the American West between the Rocky Mountains and Pacific Coast, as far north as extreme southern British Columbia. Most records are from the western part of this area, where it occurs throughout much of California, Oregon and Washington. The range is limited to Idaho and immediate vicinity in the Rocky Mountains.
Admetovis oxymorus is most commonly collected in hilly or mountainous areas with at least some trees, and occurs in a variety of habitats from riparian areas in steppe to near timberline.
Adults of A. oxymorus have been collected from late May until early August, with most records from mid-June through July. High-elevation populations fly latest, often in late July or August. It is nocturnal and comes to light.
Admetovis oxymorus is the only species in the genus for which the early stages are known. Reared larvae from southern California accepted elderberry (Sambucus mexicanus Presl.) in captivity (McFarland 1975). Godfrey (1972) illustrated the head and hypopharyngeal complex of the mature larva.
Discussion. This species has until now been confused with A. icarus. Prior records of A. oxymorus from Utah and Colorado are referable to A. icarus. Barnes, 1904 Figs 5, 6, 11, 13, 15, 17 Admetovis similis Barnes, in Dyar 1903: 157. Nomen nudum. Admetovis similaris Barnes, 1904: 200. Type material. Three male and three female syntypes from Southern California and Arizona are at NMNH. All are typical of the species indicating that lectotype designation is unnecessary.

Admetovis similaris
Diagnosis. Admetovis similaris is the easiest species in the genus to identify without examining structural characters. It is the only one in which the hindwing ground color is pure white.
Structurally, both sexes differ in several respects from those of the other two species. Males lack completely basal coremata, present to some degree in the other two. The juxta of A. similaris is smooth, lacking the median spine that is found in both other species. The left arm of the vesica bears the largest subapical diverticulum in the genus. Females have four signa on the corpus bursae, three in the other two species. The corpus bursae are similar otherwise to that of A. oxymorus in that the anterior end is bulbous, but the appendix bursae is more strongly curved in A. similaris. The barcode of A. similaris (BOLD:AAB7673) differs from both other Admetovis species by at least 3.5 %. Intraspecies variation is less than 0.9 % (n = 19; Arizona, California, Washington).
Distribution and ecology. Admetovis similaris is a species of open habitats in the Southwest, California, and Pacific Northwest. It is found near the border with Mexico from western New Mexico to the coast of southern California, thence north to southcentral British Columbia. Although its distribution is mostly in the region near the Pacific Coast it does not occur near the ocean north of the San Francisco Bay area. In the Pacific Northwest A. similaris is common on the Columbia Plateau, in the adjoining Cascade Foothills, and at low elevations in the Blue Mountains. Interestingly, it is absent from similar steppe habitats in southeastern Oregon and southern Idaho and it does not occur elsewhere in the Great Basin. Admetovis similaris almost certainly occurs in Mexico as it is found very close to the Mexican border both in Arizona and in California. This species favors the most xeric environments of any Admetovis, as dry as the Sonora and Mojave deserts. Northern populations fly most commonly in sage steppe. The flight time is during spring and early summer, typically earlier in the year than either of the other two species.
The early stages are unknown. Geographic variation. The color and pattern of this moth are uniform across its range. Specimens from deserts of the Southwest tend to smaller than those from elsewhere.
Discussion. Dyar (1903) included this species in his list of North American Lepidoptera as Admetovis similis prior to its proper description by Barnes in 1904. The Dyar mention lacks a description or illustration and is therefore a nomen nudum.