A revision of the genus Protorthodes McDunnough with descriptions of a new genus and four new species (Lepidoptera, Noctuidae, Noctuinae, Eriopygini)

Abstract The genus Protorthodes McDunnough is revised to include 15 species including P. ustulata Lafontaine, Walsh & Ferris, sp. n., from southwestern United States, P. texicana Lafontaine, sp. n., from Texas and Mexico, and P. mexicana Lafontaine, sp. n., from Mexico. A new genus, Nudorthodes Lafontaine, Walsh & Ferris, is proposed for two species formerly included in Protorthodes, P. texana (Smith, 1900), comb. n., and P. variabilis (Barnes & McDunnough, 1912), comb. n., and N. molino Lafontaine, Walsh & Ferris, sp. n., described from southern Arizona. A key to species, descriptions, illustrations of adults and genitalia, and distribution maps are included.

Diagnosis. Adults. Males and females of similar size (forewing length 11-17 mm). Vestiture of palpi, head, and thorax of long apically-forked or apically-serrated scales, sometimes with a slightly-raised central tuft near front of thorax. Head -Labial palpus porrect, apical segment about ½ as long as second segment. Frons rounded, covered with strap-like scales projecting over it from sides and top. Eye covered with surface hair. Male antenna biserrate (like toothed edge of a saw) to bipectinate (lateral processes mainly parallel sided like a feather) with lateral processes 0.5-4.0 × as long as central shaft. Female antenna filiform, setose ventrally. Thorax -Wings: Forewing ground color typically gray, brown, or orange; pattern variable, typically with reniform and orbicular spots with a pale or black outline, usually lower part or all of reniform spot filled with gray, which is darker than the ground color; postmedial line dentate or with inner element straight, outer element broken into series of dots on veins; subterminal line pale and sinuate in most species with dark wedges or shading along inner margin. Hindwing white to fuscous. Legs: Tibiae without spiniform setae. Tarsal segments 1-4 with three ventral rows of spiniform setae, and four ventral rows on tarsal segment 5. Abdomen -Base of abdomen without basal abdominal brushes. Eighth abdominal sternum of male with tuft of long setae on a short eversible coremata. Male genitalia -Uncus typically slender, slightly swollen mesially, tapered at apex to hook-like process. Valve broadest beyond middle, tapered to slight "neck" defining apical cucullus; sacculus more heavily sclerotized dorsally than ventrally with dorsal part crenulate and setose in some species; clasper a sclerotized plate in middle of valve distal to sacculus from which arises a long, heavily-sclerotized ampulla projecting posterodorsally, extending almost to, or beyond, dorsal margin of valve; ampulla centrally swollen in most species; digitus arising from large sclerotized plate in middle of valve, tapered posterolaterally into heavily-sclerotized pointed or blunt process projecting below ventral margin of valve at neck of cucullus; cucullus covered with long inward-projecting setae and with no defined apical corona. Vesica usually twisted or coiled above base with numerous pouches or diverticula basally and subbasally; a long, heavily-sclerotized basal or subbasal cornutus in most species; vesica 1-2 × as long as aedeagus. Female genitalia -Corpus bursae thin and membranous, rounded or oval, without obvious signa. Appendix bursae typically with one or two short coils. Ductus bursae variably sclerotized, usually about as long as corpus bursae. Abdominal segment eight about 2 × as long as wide; anterior apophyses 0.5-2.0 × as long as abdominal segment eight; posterior apophyses folding near middle, about 2.5 × as long as anterior apophyses. Ovipositor telescoping and projecting well beyond end of abdomen in most specimens. Anal papillae long and tapered, 0.5-1.0 × as long as abdominal segment eight; anal papillae lightly sclerotized, setae mainly confined to apical area.  Todd (1982). Taeniocampa bostura: holotype ♂, examined, AMNH. Type locality: Canada, British Columbia, Kaslo. Diagnosis. Protorthodes curtica is the dominant species of Protorthodes in the Pacific Northwest, especially in the intermontane region between the Cascades and the Rocky Mountains. Superficially, adults usually can be identified by the darkreddish tint on the forewing ground color, the pale, even-curved subterminal line that follows the wing margin, the even band of dark shading along the inner edge of the subterminal line, and the reniform is faintly outlined by a pale line. Forewing length varies from 12 to 16 mm. The male antenna of P. curtica is narrowly bipectinate, similar to those of P. incincta, the species with which P. curtica is most likely to be confused. The antenna of P. curtica is not as wide with the maximum width of the antenna being 2.6-2.9 × as wide as the central shaft, whereas it is 3.7-3.9 × as wide in P. incincta. The range of P. curtica is mainly to the west of the range of P. incincta. Populations of P. incincta where its range overlaps that of P. curtica usually have gray-brown or buffy-brown forewings with an irregular pale subterminal line that is margined on the inner side with dark shading that usually forms a series of larger wedges in the area distal to the reniform spot. The reniform spot of P. curtica is outlined with a dark line with a pale line inside this, and the forewing apex is not as acutely pointed as in P. curtica. The male and female genitalia of P. curtica and P. incincta are similar, but the digitus of P. curtica tends to be pointed where it projects beyond the ventral margin of the valve, whereas in P. incincta the apex of the digitus is tapered to a point.
Distribution and biology. Protorthodes curtica occurs from the interior of southern British Columbia southward in the West Coast states, mainly to the east of the Cascades and Coastal Ranges, to southern California. It occurs in the Rocky Mountains in Idaho and Montana and in the Ruby Mountains of Nevada. Adults occur from late June until mid-October, mainly in dry forested habitats. The larva was described by Crumb (1956) and Godfrey (1972). ( Diagnosis. Adults of Protorthodes eureka can be recognized by relatively small size (forewing length: 11-13 mm), narrow forewings, and longitudinally-streaked pattern resulting from the dark-colored veins and lighter-brown color between the veins. A series of black sagittate spots is on the inner side of the almost straight subterminal line. The male antenna and male and female genitalia of P. eureka are similar to those of P. curtica, but the right clasper of P. eureka extends beyond the dorsal margin of the valve by about 1/3 of the length of the clasper, whereas in P. curtica it exceeds the dorsal margin of the valve by less than 1/5 of its length.

Protorthodes incincta (Morrison, 1874)
Diagnosis. Protorthodes incincta is so variable in appearance that it is almost easier to identify it be eliminating the other species. The ground color varies from pale whitish gray, through various shades of brown, orange, and gray to blackish gray. Forewing length varies from 11-14 mm. The moths are most likely to be confused with those of P. curtica that mainly occurs father to the west. Protorthodes incincta can be distinguished by the more irregular pale subterminal line, concentration of dark sagittate marks proximal to the subterminal line to the area distal to the reniform spot, and other characters of maculation, antenna, and male genitalia listed under P. curtica. The palest forms (e.g., Fig. 16) occur in xeric areas of Nevada, Arizona and New Mexico and previously were known as P. indra, and the most contrastingly marked forms (e.g., Figs 11, 12) are localized in southern Wyoming, Colorado, and northern New Mexico and previously were known as P. daviesi. However, both forms blend into more typical forms in these areas and do not differ in structural characters or barcodes from other populations.
Distribution and biology. Protorthodes incincta is mainly a species of the western Great Plains and dry open forests of the Rocky Mountain region, with range extensions into the Great Basin, the American Southwest, and eastward in relict prairie areas into the Great Lakes region. Adults occur from early June until early October. The larva was described by Crumb (1956) and Godfrey (1972). Diagnosis. The name of this species is derived from the silvery-gray ground color of the forewing. This, in combination with the prominent black basal dash on the forewing and the white somewhat translucent hindwing, make this the easiest species of Protorthodes to identify. In some males there is a narrow black terminal line and broken postmedial line on the hindwing and in the female the hindwing has a pale smoky tint to the white color. Forewing length varies from 13 to 16 mm. The male antenna is biserrate, unlike other species in this group, with the maximum with of the antenna being 1.9-2.1 × as wide as the central shaft. The male and female genitalia are similar to those of the other three species in the P. incincta group (P. curtica, P. eureka, P. incincta), surprising, because of the divergent external appearance of the moth. However, the cucullus is smaller, the two digiti apically truncated and more symmetrical, the subbasal diverticula in the vesica are smaller, and the basal cornutus larger than in the other three species in the group.

Protorthodes argentoppida
Distribution and biology. Protorthodes argentoppida has a very limited range, occurring in xeric forested areas of various mountain ranges in New Mexico and in the White Mountains in east-central Arizona. Adults occur from mid-May until early July. The immature stages are unknown. ( Diagnosis. Protorthodes mulina is easily recognized by the orange or yelloworange ground color of the forewing with the maculation defined by darker orangebrown lines, and especially by the enlarged lower lobe of the reniform spot that is filled with a dark blue gray. Forewing length varying from 13 to 17 mm. The male and female genitalia are divergent from other species in the genus, although several features suggest a relationship to the P. incincta group; the vesica has a basal cornutus, the subbasal diverticula in the vesica are similar to those of species in the P. incincta group, but in P. mulina these structures are much larger than those of other species in the P. incincta group and the vesica is about twice as long. Unique features of the male genitalia are the enlarged, rounded cucullus, wider than the valve, unlike other species of Protorthodes, the clasper is almost straight with an abrupt 90°bend near the apex, and the digitus is reduced, lying entirely along the inner surface of the valve with the apex slightly expanded and forked. The female genitalia, like other species in the P. incincta group, has an amorphous sclerotized mass on the posterior part of the ductus bursae; the position and shape of the lobes of the corpus bursae are unique.

Protorthodes mulina
Distribution and biology. Protorthodes mulina undoubtedly has a wide range in Mexico, occurring as far south as the state of Chiapas, but is known from very few localities. In the United States it occurs from western Texas to southeastern Arizona. Adults are found in May and June and again from mid-August to early November, probably representing two generations. The larva was described by Crumb (1956) and Godfrey (1972).   Krogerus, 1954: 20. Type material. Taeniocampa oviduca: type material lost. Type locality: North America. Note: no species is likely to be confused with Taeniocampa oviduca in the areas of eastern North America from where Guenée's material originated, mainly the Southeast (Georgia and Florida), and the Northeast (mainly New York and eastern Canada), so no neotype is proposed. Taeniocampa capsella: syntype ♂, BMNH, examined. Type locality: New York, Albany. Protorthodes lindrothi: holotype ♂, CNC, examined. Type locality: Canada, Newfoundland, Badger.
Diagnosis. Protorthodes oviduca is mainly a boreal-zone species occurring in Canada from coast to coast. Because of its mainly northern and eastern distribution, its range overlaps that of only two other species, P. curtica in British Columbia and P. incincta in southern Canada and the Rocky Mountain region. Adults can be recognized by the reddish-brown coloration of the forewing and the contrastingly pale outline of the reniform and orbicular spots with the reniform spot usually entirely filled with dark shading. The male antenna is strongly bipectinate, 3.8 to 4.1 × as wide as the central shaft. The forewing length varies from 11 to 14 mm. This and the next two species (P. orobia and P. melanopis) form a structurally similar species group in which the ampulla of the clasper is relatively short and teardrop shaped, and does not extend to the dorsal margin of the valve; the digitus is a broad triangular sclerotized plate extending from the costal margin of the valve and tapers ventrally into heavily-sclerotized process with the apex covered with short spines and minute setae and ends near the middle of the valve, not near the "neck" of the cucullus as in other species. The female genitalia of the group are characterized by the sclerotized area toward the posterior end of the ductus bursae, which is heavily sclerotized on each side of the ductus with the central area being only lightly sclerotized. Protorthodes oviduca can be distinguished from the other two species in the group by the characters given in the key.
Distribution and biology. Protorthodes oviduca occurs across boreal and temperate areas of Canada and northern United States with extensions in eastern US to central Florida and southern Alabama, and in the mountains in the West as far south as Colorado and Utah. In some areas (e.g., Ohio, Michigan) it is found only in sandy habitats (Eric Metzler, pers. comm.). Adults occur mainly from mid-May to early July with occasional records as late as mid-August. The larva was described by Crumb (1956) and Godfrey (1972). ( Diagnosis. Protorthodes orobia is closely related to P. oviduca, but can be recognized by the gray-brown color of the forewing with a dusting of white scales that gives the wing a hoary appearance; the maculation is defined by thin white lines with the transverse lines represented on the costa by seven wider white spots that immediately separate this species from P. oviduca. Unlike P. oviduca, the reniform and orbicular spots of P. orobia are concolorous with the rest of the forewing, being defined only by the white outline. The white subterminal line of P. orobia is white and contrasting, not yellow buff with red shading on its inner margin as in P. oviduca. Structurally, the two species differ by the characters of the vesica given in the key.

Protorthodes melanopis
Diagnosis. Protorthodes melanopis is most easily recognized by the pale gray-brown forewing ground color, which emphasizes the contrast between the reniform spot, orbicular spot, and the ground color. The pale hindwings of the males contrast with the forewings. Forewing length varies from 11 to 14 mm. In males the hindwing is white and translucent, with some fuscous shading on the veins and wing margin; in females the hindwing is covered with a fuscous sheen, darker on the veins and wing margin. The antennal pectinations and wing size are the same as in P. oviduca. Structurally, the species differs from P. oviduca by the vesica character given in the key. The coil in the vesica in P. melanopis is mesial rather than basal, and the dorsal lobe on the sacculus tapers to a pointed process, whereas the dorsal lobe of the sacculus is rounded in P. oviduca and P. orobia.
Distribution and biology. Protorthodes melanopis occurs across the southern United States from western Texas to southern California. Its range extends as far north as southern Utah and as far south as northern Mexico. Adults were collected from late February to early May and again from mid-August to late September. The larva was described by Godfrey (1972). Etymology. This species is both Texan and Mexican, thus the name texicana. Diagnosis. Protorthodes texicana can be recognized by the brown forewings with the transverse lines defined by pale buff and dark-brown lines, and the reniform and orbicular spots darker brown than the ground color with a pale buff outline. Protorthodes texicana is most likely to be confused with P. oviduca or P. orobia, however, P. texicana but does not have the reddish tints to the forewing of P. oviduca, and has by the less contrasting white lines on the forewing than P. orobia. The male genitalia of P. texicana are characterized by large, heavily-sclerotized, spinulose hood-like structure that projects posteriorly from the dorsal margin of the sacculus, and the clasper-digitus combination that creates a structure with three processes; the homologies of this assemblage are best interpreted by comparison with the more rudimentary form found in P. oviduca. The female genitalia are characterized by the large, heavily-sclerotized, double-lobed plate covering the ostium bursae ventrally.

Protorthodes texicana
Description. Adult. Male and female similar in size, color, and maculation. Forewing length: 12-14 mm. Head -Male antenna with individual segments bipectinate, 3.8-4.0 × as wide as the central shaft (anterior rami 1.9-2.1 × as wide, posterior rami 1.4-1.5 × as wide). Female antenna filiform, minutely setose ventrally. Palpi and head clothed with spatulate apically-forked setae, pale buff or gray basally, darkbrown patch subapically, with pale-gray and pale buff tips creating a brown color with a hoary overlay. Thorax -Covered with similar scales to those of head; with slightly raised prothoracic and metathoracic tufts. Legs: Appearing speckled with mixture of buff and darker gray-brown scales. Tibiae without spiniform setae. Tarsi with three ventral rows of spiniform setae. Wings: Dorsal forewing pale brown with dusting of darker-brown scales; subbasal, antemedial, postmedial, and subterminal lines buff, partially bordered by darker-brown scales; reniform spot gray-brown, darker than forewing and with pale-buff outline, with slight constriction on anterior and posterior margin, giving it a figure 8 shape; orbicular spot similar in color, rounded or obliquely oval; fringe with two irregular rows of spatulate scales, pale-brown at base, darker toward tips. Dorsal hindwing pale fuscous basally with darker fuscous on discal spot, wing veins, and marginal area of wing; fringe pale fuscous white with irregular darker fuscous medial line. Male genitalia -Uncus mainly evenly tapered from base to apex except slightly swollen laterally subbasally, sparely covered with long hair-like setae, apex compressed and flattened. Valve gradually tapered from base, abruptly constricted subapically to define a rounded cucullus; sacculus sclerotized, about 0.4 × length of valve, dorsal margin extended into elongated, hollow, hood-like process with outer surface covered by spines, its apex extending to base of ampulla of clasper; ventral base of clasper extending from sacculus ending in rounded lobe from which arises a heavily-sclerotized, partially articulated, tear-drop-shaped process projecting posteriorly; dorsal base of clasper articulates with long heavily-sclerotized ampulla of clasper with apex spatulate, extending posterodorsally beyond costal margin of valve; digitus appearing to be a heavily-sclerotized extension from editum; apex of digitus a stout apically blunt process with short conical setae-tipped spines at apex; cucullus densely covered with long, stiff setae projecting anterodorsally, and no apical corona; central part of transtilla densely spinulose; juxta a long narrow plate, broad basally, tapered posteriorly with subapical constriction. Aedeagus extended ventrally into short spine-like tip free of vesica. Vesica about 2 × as long as aedeagus; vesica with submedial coil and ½ postmedial coil, the latter with short diverticulum on posterior side; vesica without spines or cornuti. Female genitalia -Corpus bursae membranous, rounded, without signa. Appendix bursae with two short coils, arising from posterior dorsal surface of corpus bursae. Ductus bursae 1.5 × as long as corpus bursae, anterior 2/3 membranous, posterior 1/3 covered with thin, longitudinal sclerotized striations; a heavily sclerotized, double-lobed plate covering ostial part of ductus bursae and anterior-ventral part of abdominal segment eight; abdominal segment eight 1.5 × as long as wide, lightly sclerotized; anterior apophyses 0.5 × as long as abdominal segment eight; posterior apohyses folding near middle, about 2.5 × longer than anterior apophyses. Anal papillae long and tapered, 0.5 × as long as abdominal segment eight; anal papillae lightly sclerotized, long setae sparsely scattered over surface, short setae abundant near apex of papillae.
Distribution and biology. The immature stages are unknown. Protorthodes texicana is known from two areas, west-central Texas and southern Mexico. Adults were collected between late March and late May and in early October. The immature stages are unknown. Etymology. This species is known only from Mexico, thus the species name mexicana. Diagnosis. Protorthodes mexicana is closely related to P. texicana. Protorthodes mexicana can be recognized by the pale whitish-buff color of the forewing, the white translucent hindwing. In the male genitalia the ventral process of the clasper assemblage is apically spatulate in P. mexicana (apically tapered in P. texicana), and the vesica has two full coils (1½ in P. texicana).

Protorthodes mexicana
Description. Adult male. (Female unknown). Forewing length: 12 mm. Head -Male antenna with individual segments bipectinate, 3.8 × as wide as the central shaft (anterior rami 1.9 × as wide, posterior rami 1.4 × as wide). Palpi and head clothed with spatulate apically-forked setae, pale whitish, some with pale-brown patch subapically, with whitish-buff tips. Thorax -Covered with similar scales to those of head; with slightly raised prothoracic and metathoracic tufts. Legs: Appearing speckled with mixture of whitish-buff and gray-buff. Tibiae without spiniform setae. Tarsi with three ventral rows of spiniform setae. Wings: Dorsal forewing pale whitish buff brown with dusting of pale-brown scales; subbasal, antemedial and postmedial lines very faint, indicated by paler lines bordered on each side by scattered pale-brown scales; subterminal line more distinct because of darker shading in terminal area and outer part of subterminal area adjacent to it; reniform and orbicular spots slightly darker than ground color, outlined in white; terminal area darker than remainder of forewing because of more numerous gray-brown scales; fringe similar in color to subterminal area with base of fringe whitish-buff. Dorsal hindwing white, translucent, with trace of darker scaling on veins and wing margin; fringe white with scattered pale fuscous scales. Male genitalia -Similar to those of P. texicana with following exceptions: lower process of clasper apically flattened and enlarged, spatulate (apically tapered and tear-drop shaped in P. texicana); spine-covered dorsal process of sacculus acutely tapered to point (posterior apex blunt and rounded in P. texicana); vesica with two coils (1½ coils in P. texicana).
Distribution and biology. The female and immature stages are unknown. The species is known only from the type locality where the holotype was collected in late April. Diagnosis. Protorthodes perforata can be recognized by the pale whitish gray to buffy gray color of the forewings with the reniform and orbicular spots darker gray than the ground color and each is outlined by a contrastingly pale line. Forewing length varies from 12 to 14 mm. The hindwing is dirty white with fuscous shading on the wing margins and veins in both sexes. This species is most likely to be confused with P. rufula in southern California where their ranges overlap. The forewing in P. rufula is darker and more mottled than that of P. perforata, and P. rufula usually has shades of red in the ground color. Males of P. perforata can readily be distinguished from those of P. rufula by brushing the scales away from the apices of the valves; in P. perforata the apices of the digitus on each valve are similar, barely extend beyond the ventral margins of the valve, and the apices curl to project anteroventrally. In P. rufula, by contrast, the left digitus projects posteroventrally, whereas the right digitus is much longer and projects posteriorly and extends beyond the apex of the right valve (Fig. 66). On dissection, the valves of P. perforata are very wide postmedially with both the dorsal and ventral margins rounded and convex; the ampulla of the clasper is almost straight, bending posteriorly only near the apex; the heavily-sclerotized dorsal lobe of the sacculus is much larger and more convex than in P. rufula; and the large cornutus in the vesica is near the middle, whereas in P. rufula it is much closer to the base of the vesica; also the cornutus projects at a right angle to the axis of the vesica and parallel to the aedeagus. In the female genitalia of P. perforata the sclerotized mass of the ostium bursae covers almost the entire ductus bursae and is rounded anteriorly.

Protorthodes perforata (Grote, 1883)
Distribution and biology. Protorthodes perforata occurs across the southern United States from western Texas to southern California and southward to central Mexico. According to material examined, the range of P. perforata lies to the east of that of P. rufula with the westernmost records of P. perforata being from the desert areas in interior southern California. Adults were collected from early April to early July and again from mid-August to late October. The immature stages are unknown. Diagnosis. Protorthodes rufula is much more varied in color than P. perforata with the forewing varying from pale whitish buff, through various shades of red and orange, to brown. Unlike in P. perforata, almost all specimens show some areas or patches of rufous shading and the ground color is much more mottled in P. rufula. Forewing length ranges from 13 to 16 mm, averaging larger than in P. perforata. In the male genitalia, the right digitus projects posteriorly beyond the end of the valve, whereas on the left valve it is shorter and projects posteroventrally below the valve; the ampulla of the clasper is evenly curved in a rounded arc on both valves; the dorsal lobe of the sacculus is elongated and narrow with the dorsal margin slightly concave and extending into a rounded densely setose lobe anteriorly. The vesica has a long cornutus about 1/3 from the base that projects along the axis of the vesica toward the end of the aedeagus. In the female genitalia the ostium bursae is covered by a heavily-sclerotized ventral plate that is lobed and convex posteriorly and concave where it meets the ductus bursae anteriorly; the ductus bursae is elongated, about 2 × as long as the ostial plate, and is sclerotized with elongated ridges and folds.

Protorthodes rufula (Grote, 1874)
Distribution and biology. Protorthodes rufula is a species of the far west with most records being from the Pacific Coast, and the coastal mountain ranges from northern Washington to southern California -to the west of the known range of P. perforata in southern California. Adults were collected from mid-April to mid-June in the North (from mid-February in southern California) and again from early August to late October. The larva was described by Crumb (1956) and Godfrey (1972).  Etymology. The name ustulata is Latin and refers to the burnt-orange color of with body and forewings of the species.

Protorthodes ustulata
Diagnosis. Protorthodes ustulata looks like a dark burnt-orange form of P. perforata and was usually identified as either P. perforata or as P. oviduca in collections because of the orange in the ground color and the contrasting pale outlines around the reniform and orbicular spots. The species appears to be most closely related to P. perforata and P. rufula, not just because of the superficial similarity, but because of the heavily-sclerotized spiculate lobe on the dorsal part of the sacculus, a structure restricted to these three species. Beyond that, however, Protorthodes ustulata is unique within the genus in several features: in the male genitalia the valves are very long and slender and the apices easily broken in brushing or dissection; the digitus is vestigial, ending in small lobe free of surface of valve at ventral margin of valve; the spine-covered dorsal lobe on the sacculus is oblique to the longitudinal axis of the valve. In the female genitalia, the ostium bursae and posterior part of the ductus bursae form an elongated, narrow, heavily-sclerotized tube, and the appendix bursae has two coils, both features otherwise found only in Protorthodes alfkenii.
Description. Adult. Male and female similar in size, color, and maculation. Forewing length: 12-15 mm. Head -Male antenna biserrate with projections (rami) tapered, anterior rami about as long as width of central shaft, posterior rami half as long. Female antenna filiform, minutely setose ventrally. Palpi and head clothed with spatulate apically-forked setae, dark orange or brown, some with white tips giving head a hoary look, white tips more frequent on margin of prothoracic collar (patagia). Thorax -Covered with similar scales to those of head; with slightly raised prothoracic and metathoracic tufts. Legs: Appearing speckled with mixture of dark-orange, brown scales, some white tipped. Tibiae without spiniform setae. Tarsi with three ventral rows of spiniform setae. Wings: Dorsal forewing a burnt-orange color, tending to be darker toward costal and outer edge of forewing; subbasal, antemedial, postmedial, and subterminal lines whitish gray, partially bordered by dark-brown scales; reniform spot kidney shaped, to markedly constricted mesially, so almost figure 8 shaped, upper part gray brown, lower part blackish gray, overall darker than forewing and with contrasting whitish-gray outline; orbicular spot obliquely oval, slightly darker than ground color, outlined in whitish gray; terminal line dark brown; fringe similar in color to forewing except base yellow with fine yellow streak at end of each wing vein. Dorsal hindwing pale fuscous basally with darker fuscous on discal spot, wing veins, and marginal area of wing; fringe buff white with fuscous medial line. Male genitalia -Uncus slender, evenly tapered from base to apex, sparely covered with long hair-like setae, apex tapered to point. Valve elongated and narrow, about 5 × as long as width at base of clasper, gradually narrowing to long "neck" before rounded cucullus densely covered with long, stiff setae projecting anterodorsally, with no apical corona; sacculus sclerotized, about 0.4 × length of valve, ending posteriorly in spine-covered oblique lobe, somewhat mushroom shaped, with sides extending posteroventrally almost to ventral margin of valve and anterodorsally over dorsal margin of valve; clasper about 1/3 length of valve, curved in slight arc to project posterodorsally over dorsal margin of valve; digitus vestigial, a slightly raised, lightly sclerotized rod extending to base of "neck" of cucullus and ending in slightly raised lobe projecting posteroventrally to ventral margin of valve; juxta a long posteriorly-tapered plate with middle cleft and lightly sclerotized almost to anterior end, posterior apices on each side of cleft pointed and heavily sclerotized. Aedeagus long and cylindrical, 12 × as long as medial width, mainly lightly sclerotized except for posterior half of ventral margin, which ends in slightly-sclerotized lobe. Vesica about 1.5 × as long as aedeagus; vesica with slight basal coil from which arises a curved, tapered diverticulum, and a medial coil with a minute diverticulum; apex of vesica swollen; vesica without spines or cornuti. Female genitalia -Corpus bursae membranous, rounded, without signa. Appendix bursae with two short coils, arising from posterior dorsal surface of corpus bursae. Ductus bursae 0.6 × as long as corpus bursae, anterior 1/3 membranous, posterior 2/3 covered with smooth heavily-sclerotized plate, slightly wider anteriorly than posteriorly; ostium not differentiated from ductus bursae; abdominal segment eight slightly longer than wide, more lightly sclerotized posteriorly; anterior apophyses slightly longer than abdominal segment eight; posterior apophyses folding near middle, about 2 × as long as anterior apophyses. Anal papillae long and tapered, 0.75 × as long as abdominal segment eight; anal papillae lightly sclerotized, long setae sparsely scattered over surface, short setae abundant near apex of papillae.
Distribution and biology. Protorthodes ustulata occurs from southeastern Wyoming southward to the Guadalupe Mountains in western Texas and westward to central and southeastern Arizona and northern Mexico. Adults were collected from early April to mid-May and again from early August to early October. The immature stages are unknown. Note. The name Perigea perplexa was listed without description by Smith (1893) and credited to Grote by virtue of Grote distributing the name through his check lists, which lacked any description. Grote described Perigea alfkenii in 1895 and lists Perigea perplexa as his check list manuscript name for it. Hampson (1909) lists Perigea perplexa as an unavailable senior synonym of Perigea alfkenii and credits the name to Smith. Franclemont and Todd (1983) simply list Protorthodes [Perigea] perplexa (Grote, 1895) as a synonym of Protorthodes alfkenii. The name, neither validated, nor made available, should be deleted from the synonymy.

Protorthodes alfkenii (Grote, 1895)
Diagnosis. Protorthodes alfkenii is an extremely variable species in terms of size, ground color, and pattern, yet with practice it usually is easily identified by a combination of features. The orbicular spot usually is rounded, surrounded by a thin black line, and the spot itself usually is paler than the ground color, often contrasting so; the reniform spot is oblique, unlike other species in the genus, with the lower part of the spot projecting toward the anal angle of the wing; the light and dark marks on the forewing, and the tendency for longitudinal streaks on the wing, give the forewing a busy appearance instead of the softer, more even ground pattern of most other species. The hindwing in males, and many females, is white, often with a slight pearly sheen, with fuscous shading confined to the veins and outer part of the wing. Some females have more extensive fuscous shading on the hindwings, but usually a pearly sheen is still evident. Forewing length varies from 11 to 14 mm. The male genitalia differ from those of other species of Protorthodes in that the digitus is long and pointed, projecting below the ventral margin of the valve at a right angle, the ampulla of the clasper is almost straight, projecting dorsoposteriorly toward the upper part of the cucullus, then bending abruptly through 90°near its apex to project ventrally. The vesica has 2½ medial coils and projects anterolaterally to the right from the end of the aedeagus. In the female genitalia the appendix bursae has two full coils and is at the posterior end of the corpus bursae on the left side. The ductus bursae is long and cylindrical, about 7 × as long as wide with the posterior 4/5 th lightly sclerotized.
Distribution and biology. Protorthodes alfkenii is the most abundantly collected species in the American Southwest, including southern California. It occurs from central Oregon, southern Idaho, central Wyoming, and northwestern Texas southward to southern Mexico. It occurs mainly in open arid woodlands and although its range surrounds the Great Basin, it is largely absent from the Basin. Adults were collected in the South from April until late June and again from early September to early November. In the Pacific Northwest they fly mid-July to late September. The larva was described by Crumb (1956) and Godfrey (1972). (Barnes & McDunnough, 1912) Figs 44, 45, 69, 86. Map 13 Eriopyga antennata Barnes & McDunnough, 1912a: 21. Type material. Syntypes 4 ♂, 3 ♀, USNM, examined. Type locality: Arizona, Redington. Diagnosis. Males of Protorthodes antennata can easily be identified by the very long pectinations of the antennae, which gives them a feathery appearance. The other diagnostic feature, which applies to females as well, is the form of the reniform spot. In P. antennata the reniform spot is not outlined like in other species of Protorthodes; there is a series of tiny white dots that partially define the reniform spot, and a series of tiny yellow dots that form a partial outer border of the spot. Forewing length varies from 10 to 14 mm with females tending to be larger than males. The male genitalia are characterized by the short, broad, apically-truncated digitus, and the very narrow, posteriorly tapered juxta. The apex of the aedeagus has a double field of flattened scale-like sclerites that create a lizard-skin appearance. In the female genitalia, the ductus bursae is inflated mesially and covered with minute sclerotized spicules; the ductus is mainly membranous except for a narrow sclerotized "collar" at the posterior end.

Protorthodes antennata
Distribution and biology. Protorthodes antennata is rarely collected but can occasionally be locally common. It has a small distribution extending from central Arizona to northernmost Mexico. Adults seem to have an abbreviated flight period between mid-May and mid-June, and in October. The immature stages are unknown.   Smith, 1900: 476. Etymology. The generic name Nudorthodes is a reference to this species being removed from the other members of the Orthodes group of genera by the lack of hairs on the surface of the eye. From the Latin Nudus [bare] + Orthodes.
Diagnosis. The genus Nudorthodes differs from other genera in the Hadenini: Eriopygina in lacking visible hair-like setae on the surface of the eye, and by the very long vesica in males and appendix bursae in females. Males can be distinguished from those of most other eriopygine genera by the filiform antennae. The genus includes three species: N. texana (Smith, 1900), N. variabilis (Barnes & McDunnough, 1912), and N. molino Lafontaine, Walsh, & Ferris, sp. n.
Description. Adult: Head -Frons rounded; labial with apical segment about 1/4 as long as second segment; male antenna filiform, or very slightly constricted between segments (N. variabilis), setose ventrally; female antenna filiform, setose ventrally; eye rounded, without surface hairs; ocellus present. Thorax -Thorax clothed with narrow, spatulate, apically serrated scales that form a slightly raised tuft on the prothorax, and a partially divided tuft on the metathorax. Legs: middle and hind tibiae without spiniform setae and with three ventral rows of spiniform setae on tarsi. Wings: forewing venation typically quadrifine, cubital vein appearing four branched; hindwing with typical trifine venation (i.e., M2 reduced, about 2/3 down cell and parallel to M3. Abdomen -basal abdominal brushes and pockets absent; eighth abdominal sternite of male with a slightly eversible coremata with a transverse tuft of long setae. Male genitalia -Valva: symmetrical, basal ¾ of valve mainly parallel-sided except for dorsal lobe of sacculus (basal half of valve, mainly sacculus, massive compared to narrow apical half in N. molino); valve slightly constricted at ¾ from base to define a tapered densely setose cucullus with much stouter setae on apical and ventro-apical part of cucullus forming an irregular corona; digitus projecting posteriorly along middle of valve, bending ventro-posteriorly near base of cucullus and flattened into elongated plate extending beyond ventral margin of valve (apical part of digitus slightly s-curved and flattened into a foot-shaped structure in N. molino); clasper arising as a sclerotized rod at apex of sacculus projecting posteriorly with ampulla bending abruptly through 45°angle to project posterodorsally beyond costal margin of valve; sacculus heavily sclerotized, with a large dorsal process (lobe somewhat quadrangular in two species, and produced posteriorly in N. molino), without membranous flap; Uncus: decurved, cylindrical, tapered apically to a down-curved sharply pointed apex. Aedeagus: long and slender, about 7 × as long as wide, dorsal surface mostly membranous, lateral margins extended on to base of vesica; everted vesica tubular, about 4-5 × as long as aedeagus, basal area angled and with several diverticula, main part gently curved through 5 or 6 coils, vesica expanded and covered with short sclerotized spine-tipped granules toward apex. Female genitalia -Corpus bursae rounded, membranous, without signa and with inner surface covered with minute spicules. Appendix bursae arising from left posterior part of corpus bursae and extending anteriorly through several open coils; appendix bursae about 2 × as long as corpus bursae. Ductus bursae about at long as corpus bursae, mainly membranous, with a sclerotized collar-like ring at posterior end forming a slightly wider ostium bursae. Abdominal segment eight sclerotized, covered posteriorly with numerous long, heavily-sclerotized setae, many half as long as segment. Anterior apophyses rod-like, 1.2-1.4 × as long as abdominal segment eight. Posterior apophyses 1.6-1.9 × longer than anterior apophyses. Anal papillae lightly sclerotized, bullet shaped, gradually tapered to rounded apex; surface covered with long hair-like setae, especially toward apex.
Larva and habits. The only species of Nudorthodes known as a larva is Nudorthodes texana. Like Protorthodes the species are generally associated with xeric habitats. The larvae were described by Crumb (1956) and Godfrey (1972), and were separated from Protorthodes by both authors and described as being more similar to larvae of Homothodes McDunnough. The larva of Nudorthodes texana differs from those of Protorthodes species in that the larval skin is smooth, not granulose, the setae arise from sclerotized rings, not pinacula, the posterior part of the prothoracic shield is not contrastingly pale, sclerotized plates between the bases of the abdominal prolegs are lacking, and the apical seta of the labial palpus (Lp-2) is similar in length to the basal segment of the palpus (Lps-1).
Distribution. USA (Arizona, California, Nevada, Oregon, Texas, Utah, Washington).  Todd (1982), examined. Type locality: USA, Phoenix, Arizona. Diagnosis. Nudorthodes texana is the most widely distributed and common species in the genus. In most of its range adults can be recognized by the pale buffy-brown or gray-brown color of the forewings with darker shading around the reniform and orbicular spots and in the outer part of the subterminal area. The male and female genitalia of N. texana form the basis for the description of the genus Nudorthodes; it and N. variabilis are structurally very similar, differing mainly in the shape of the costal process of the sacculus and the relative lengths of the ampulla of the clasper in the male genitalia. The barcode of N. texana is most similar to that of N. variabilis, the two differing by 3.67-3.98 %.
Distribution and biology. Nudorthodes texana occurs from the intermontane region of Washington, Oregon, Nevada, and Utah southward to southern California and Arizona and southeastward to the Gulf Coast of Texas. Adults were collected rarely in the spring, mostly in March; the second generation occurs from mid-August to mid-November with the flight earlier in the North than in the extreme South. The larva was described by Crumb (1956) and Godfrey (1972).
Notes. A peculiar form of Nudorthodes texana (Fig. 49) occurs in southwestern Arizona in the vicinity of Yuma. The forewing is a mottled pale orange brown or reddish brown, without the dark shading between the reniform and orbicular spots characteristic of N. texana. The barcodes of this population differ from those of N. variabilis by 3.36-3.67 % and from those of N. texana by 4.43-5.05 %. However, many specimens from the Yuma area are intermediate in appearance between the mottled-orange form and the normal form of N. texana and these barcode with either haplotype clade, suggesting adults of the two haplotype groups are interbreeding, or did so in the past. For this reason we do not describe the Yuma form as a distinct species, but suggest more research on the two haplotype groups in this area is needed. Only five specimens of the deviant haplotype are known. (Barnes & McDunnough, 1912), comb. n. Figs 50,51,71,88;Map 15 Namangana variabilis Barnes & McDunnough, 1912b: 21. Type material. Namangana variabilis: syntypes, USNM, examined. Type locality: USA, California, San Diego.

Diagnosis.
Nudorthodes variabilis is a rarely collected species confined to coastal southern California. Adults are most likely to be confused with those of N. texana, but average larger (forewing length: 13 to 14 mm). In Nudorthodes variabilis, the medial line usually is prominent, extending obliquely from the costa to the reniform spot and as a straight line from there to the hind margin of the forewing, and the lower third of the reniform spot is filled with a well-defined dark blue-gray patch. In N. texana the medial line is absent or barely traceable, and the lower third of the reniform spot has diffuse dark shading. In southern California where the ranges of the two species overlap, the forewing ground color in N. variabilis is an even gray brown, sometimes with a slight reddish tint; whereas in N. texana the ground color is pale buff and powdery in appearance. In the male genitalia the dorsal lobe on the sacculus of N. variabilis is produced posteriorly towards the top, giving it a slight mushroom shape, whereas the posterior margin of the lobe in N. texana is straight and perpendicular to the longitudinal axis of the sacculus. The female genitalia of the two species appear to be indistinguishable.
Distribution and biology. Nudorthodes variabilis occurs along the coast of southern California from Santa Barbara County to San Diego County. Adults were been collected from late August until mid-September. The immature stages are unknown. genitalia -Similar to generic description except as noted. Valves mainly symmetrical except dorsal process of sacculus, and ampulla of clasper larger on right valve than on left valve. Valve widest at 1/3 from base, abruptly tapered near middle, then gradually tapered to ¾ from base to "neck" of cucullus; cucullus densely covered with posterodorsally-directed hairlike setae on anterior half of cucullus with stout setae curving to project anterodorsally on posterior half of cucullus; digitus abruptly-angled ventrally near neck of cucullus with apical part curved, ventrally flattened and foot shaped; ampulla of clasper expanded and lobed at base, then gradually tapered to apex, abruptly curved through 90°angle at ¾ to project dorsally, right clasper about 15% larger than left one; dorsal process of sacculus elongated and lobed posteriorly, process larger on right valve than on left valve; juxta an elongated flattened plate extending into series of oblique sclerotized ridges in membrane directed toward midline. Aedeagus and vesica similar to those of N. texana and other species but apical part of vesica more inflated. Female genitalia -Corpus bursae membranous, rounded, without signa; appendix bursae arising from posterior dorsal surface of corpus bursae, 1.5 × as long as corpus bursae and slightly coiled toward anterior end; ductus bursae about as long as corpus bursae, with expanded sclerotized ostium bursae on posterior ¼; abdominal segment eight as long as wide, lightly sclerotized; anterior apophyses 1.5 × as long as abdominal segment eight; posterior apophyses folding near middle, about 2 × as long as anterior apophyses. Anal papillae long and tapered, about as long as abdominal segment eight; anal papillae lightly sclerotized, long setae sparsely scattered over surface, short setae abundant near apex of papillae.

Nudorthodes molino
Distribution and biology. Nudorthodes molino is known from southeastern Arizona and southwestern New Mexico. Adults were collected from late May until late June and from late August until early November. The immature stages are unknown.