A revision of the genus Bryolymnia Hampson in North America with descriptions of three new species ( Lepidoptera , Noctuidae , Noctuinae , Elaphriini )

Th e three known North American species of Bryolymnia Hampson, 1908 are reviewed and three additional species are described as new. Two additional species, Elaphria ensina (Barnes, 1907) and Cryphia viridata (Harvey, 1876) are transferred to Bryolymnia as new combinations. Th e North American species are compared with related species in Central America. Adults of 11 species and male and female genitalia, where available, are illustrated.


Introduction
Th e genus Bryolymnia Hampson, 1908 includes 17 species occurring from western United States to Argentina (Poole 1989).An additional two species transferred to Bryolymnia, one from Elaphria Hübner, one from Cryphia Hübner, and the three species described below, bring the total to 22 species.
Th e genus is associated with the Elaphrini on the basis of the weakly sclerotized area on the costa of the male valve near the apex of the digitus, and by the membranous transverse suture in the sacculus about 2/3 from its base that isolates the apical triangular sclerite that usually is more heavily sclerotized than the basal part of the sacculus.
Bryolymnia can be recognized by the large, rounded or lobed plate-like process arising from the sclerotized rod that connects the dorsoanterior rod of the clasper to the costal margin of the valve.In most species this process extends dorsally beyond the dorsal margin of the valve.Superfi cially, in most species of Bryolymnia the medial fore wing area is pale and contrasts with black patches in the basal area, before or distal

JBW
Personal collection of J. Bruce Walsh, Tucson, Arizona, USA.NMSU New Mexico State University, Las Cruces, New Mexico, USA.USNM National Museum of Natural History (formerly, United States National Museum), Washington, District of Columbia, USA.

Dissecting methods and genital terminology
Dissection of genitalia and terms for genital structures and wing markings follow Lafontaine (2004).
Diagnosis.Adults.Males and females of similar size; fore wing length 9-18 mm.Head -Male antenna fi liform to beadlike, occasionally very slightly biserrate; setae tending to group into tuft on each side in species with slightly biserrate antenna.Female antenna fi liform, minutely setose ventrally.Frons rounded and smooth, or slightly bulging in middle, covered with short broad scales.Eye rounded, surface smooth, without surface hair.Labial palpus porrect, apical segment 0.3-0.4× as long as second segment.Th orax -Th orax: covered with broad apically serrated scales; scales tend to form slightly raised dorsal tuft on prothorax and spreading, slightly doubled dorsal tuft on metathorax.Wings: Fore wing ground color typically of brown and black with areas of white or pale green, especially medial area.Hindwing white to fuscous.Legs: Covered with short broad scales; tarsi with three ventral rows of spiniform setae.Abdomen -Base of abdomen with basal abdominal brushes, levers, levers and pockets in most species.Eighth abdominal sternite of male with slightly eversible coremata with a tuft of long setae.Male genitalia -Uncus cylindrical, thin, tapered from base to spine-like apex.Tegumen broad, truncated apically, ventrally expanded on posterior surface into round projecting lobe on each side.Valve long and mainly parallel sided, most often bent dorsally at point of weakness of costa; costa a sclerotized thickening on dorsal margin of valve, but dorsal margin with weakly sclerotized area at point where digitus bends away from costa; a slightly expanded cucullus with partial or complete corona on apical margin; digitus mainly a sclerotized band fused to mesial surface of valve adjacent and partially fused with costa, except at weakened area of costa where digitus bends obliquely posteroventrally across valve with apex tapered and free from surface of valve; apical part of clasper a simple curved (or slightly S-shaped) rod projecting posteriorly on inner surface of valve, with minute teeth or short subbasal process in some species; ventral base of clasper extending as a thin sclerotized rod to apex of sacculus; dorsal base of clasper diagnostic, expanded into large, rounded or lobed heavily sclerotized plate extending above dorsal margin of valve; sacculus 0.3-0.4× length of valve, mainly weakly sclerotized, with membranous transverse suture at 2/3 from base and with apical 1/4 more heavily sclerotized.Aedeagus weakly and patchily sclerotized, often with apical part of aedeagus extended as variably sclerotized straps onto base of vesica.Vesica short, about 1.5-2.0× length of aedeagus; vesica variably armed with "brushes" of long thin spines, stout cornuti, often projecting obliquely from a strongly sclerotized base, or with fi elds of minute spinules.Female genitalia -Corpus bursae thin and membranous, rounded, oval, or elongated, with one or two long signa in some species.Ductus seminalis at posterior end of corpus bursae in most species.Appendix bursae absent in most species; one species in North America (B.semifascia) with rounded, sclerotized appendix bursae at junction of corpus bursae and ductus bursae.Ductus bursae with length, shape, and amount and position of sclerotization highly variable and species-specifi c.Ventral wall of ostium bursae sclerotized and forming a plate that projects posteriorly.Anterior and posterior apohyses rod-like, 1.0-1.5 × length of abdominal segment eight.Anal papillae lightly sclerotized rounded posteriorly, covered with mixture short and long setae.Diagnosis.Superfi cially Bryolymnia viridimedia can be recognized by large size (fore wing length: 14-18 mm), extensive pale-green shading in the medial area, and the course of the antemedial line on the fore wing.On the fore wing the antemedial line has a rectangular indentation between the cubital and anal veins, and an outward loop below the anal vein; the basal area is black, except for the area below the anal vein, which is pale green; the subterminal and terminal areas are mainly brown with gray scales (mainly adjacent to the lower part of the postmedial line, and with a dark-fuscous truncated wedge-shaped patch on the costa.Th e hindwing is pale fuscous with darker shading on the veins, discal spot, postmedial line, and wing margin.Bryolymnia viridimedia is similar to B. poasia Schaus, 1911 from Costa Rica (Figs. 4, 5, 26, 35) and B. marti (described below).Bryolymnia poasia is smaller (fore wing length: 12-14 mm) than B. viridimedia, the fore wing medial area is white or whitish buff , not green, and the genitalia diff er.Male genitalia.In B. viridimedia the cucullus is short, similar in width to the rest of the valve, and has a partial corona on the dorsal-apical margin; the clasper is a simple fi nger-like process with a large rounded or slightly lobed sclerotized plate on the dorsal-anterior rod at the base of the clasper.Th e vesica has two dense patches of long spines ventrolaterally on the right, one postmedial and one preapical, the latter on a short diverticulum.In B. poasia (Fig. 26) the valve is straighter and narrower toward the base, the rounded sclerotized plate on the dorsal-anterior rod at the base of the clasper is larger and more rounded, and the vesica is short with a single large basal cornutus on a large base.Female genitalia.Th e corpus bursae is oval, 4 × as long as abdominal segment eight and 0.8 × as long as the ductus bursae.Th e ductus bursae is long and narrow with irregular sclerotized ridges longitudinally, especially mesially and anteriorly.Th e ostium bursae has an elongated heavily sclerotized plate in the ventral wall with rounded sides and a deep central notch posteriorly.

Key to species of
Distribution and biology.Bryolymnia viridimedia occurs from southeastern Arizona (Huachuca and Santa Rita Mountains) southward in the Sierra Madre Occidental to the Mexico City area.Collecting dates range from early July to mid-September.Bryolymnia bicon occurs south of the range of B. viridimedia, from the State of Veracruz in central-eastern Mexico southward to Costa Rica.Etymology.Th e species is named in honor of Marti Romero, who fi rst collected the species and was extremely helpful in collecting most of the type series.instead of gray-green shading, a less prominent medial line, and a smaller black dot in the orbicular spot.Description.Adults.Male and female similar in size, color, and maculation.Head -Male antenna with fl agellomeres very slightly swollen laterally; setae tending to group into two or three clusters on each side of each segment.Palpi and head a mixture of broad, apically serrated pale-brown and blackish-brown scales predominating.Th orax -Covered with scales similar to those on head, except blackishbrown scales predominating; a slightly divided dorsal tuft on metathorax.Legs: Covered with blackish-brown scales with white band in middle of tibiae and at apices of tarsal segments.Tibia without spiniform setae.Wings: Fore wing length: 11-12 mm.Dorsal fore wing ground color black in basal and antemedial area, pale whitish green in medial area, and a mixture of black, brown and greenish gray in subterminal and terminal areas; subterminal line with concave notch between cubital and anal veins, and bending abruptly toward wing base below anal vein; medial line obscure or diff use in most specimens; postmedial line black, slightly serrated with a series of short wedge-shaped extensions projecting into medial area; subterminal line evident only near costa where blackish gray in upper part of subterminal area contrasts with paler greenish gray in upper part of terminal area; terminal line black; fringe dark gray with white scales at end of veins; orbicular spot usually obscure but outlined partially or completely in black in some specimens; reniform spot shaped like fi gure 8 but upper part usually obscure, lower part white with black outline and small black central dot.
Dorsal hindwing white with varying amount of fuscous, especially on veins, discal spot, postmedial line, and wing margin; females averaging slightly darker than males; fringe fuscous with white basal line and sometimes with white on outer margin.Male genitalia -Uncus cylindrical, tapered to spine-like apex.Valve somewhat oval, broad mesially, tapered apically with dorsal margin angled upward at membranous part of costa; cucullus with weak corona on apical half of outer margin; digitus long and tubular, fused to costal margin of valve to costal angle, then free of valve and slightly spatulate; apical part of clasper slender and very slightly upcurved, dorsal part of basal rod extending to dorsal margin of valve enlarged into apically rounded process extending beyond dorsal margin of valve; apical part of sacculus beyond postmedial suture more heavily sclerotized than basal part.Aedeagus irregularly sclerotized with narrow sclerite extending onto vesica ventrally to base subbasal bend in vesica.Vesica about 1/2 as long as aedeagus with postmedial cornutus tapered to sharp point.Female genitalia -Corpus bursae membranous, oval, slightly shorter than ductus bursae, with single long signum ventrally and ductus seminalis arising on right near junction with ductus bursae.Ductus bursae long and slender, membranous except for lightly sclerotized band about 3/4 from posterior end.Ostium bursae with broad sclerotized plate in ventral wall with deeply concave posterior margin.Etymology.Th e name mixta is from the Latin mixtus, meaning mixed or mingled and refers to the blotchy confused appearance of the fore wing maculation.
Diagnosis.Bryolymnia mixta can be recognized the mottled black and brown fore wing pattern that obscures the maculation.It is most similar to B. ensina, but in B. mixta there is much more black shading in the medial area than the basal area, the opposite of B. ensina.In the male genitalia the sclertoized plate at the base of the clasper is somewhat mushroom shaped, narrow basally and abruptly expanded apically, and the apex of the digitus also is abruptly expanded.In the vesica there is a narrow sclerotized rod that extends from the apex of the aedeagus to a long spine-like medial cornutus.In B. ensina the sclerotized plate on the basal arm of the clasper is tapered to a broadly rounded apex, and the digitus is tapered apically.In the vesica of B. ensina there is a broad, heavily-sclerotized lobe extending from the apex of the aedeagus onto the vesica and the apical 1/2 of the vesica is covered with minute spines.
Description.Adults.Female unknown.Head -Male antenna with fl agellomeres very slightly swollen laterally; setae tending to group into two or three clusters on each side of each fl agellomere.Palpi and head a mixture of broad apically serrated pale-brown and blackish-brown scales, the latter predominating.Th orax -Covered with scales similar to those on head, except blackish-brown scales predominating; a slightly divided dorsal tuft on metathorax.Legs: Covered with blackish-brown scales with white band in middle of tibiae and at apices of tarsal segments.Tibia without spiniform setae.Wings: Fore wing length: 12 mm (2 specimens).Dorsal fore wing ground color brown heavily dusted with blackish-brown scales, especially in lower part of medial area, to a lesser degree in lower part of basal and subterminal areas, and forming a dark streak distal to reniform spot and in costal part of subterminal area; antemedial and postmedial lines black but partially obscured by scattered black scales on fore wing; reniform and orbicular spots with some gray scales but mainly obscured by mottled ground color; subterminal line and apical part of terminal area pale brown; fringe dark gray.Dorsal hindwing fuscous, darker on discal spot, veins, and wing margin.Fringe pale buff with fuscous medial line.Male genitalia -Uncus cylindrical, tapered to spine-like apex.Valve strap-like with ventral margin slightly convex near middle and dorsal margin slightly angled at point of weak sclerotization; apex of valve with well-developed corona on outer margin; digitus long and tubular, mainly fused to costal margin of valve except for short spatulate apex; apical part of clasper slender and upcurved, dorsal part of basal rod extending to dorsal margin of valve enlarged into apically rounded process with narrow base, somewhat mushroom shaped; apical part of sacculus beyond postmedial suture slightly more heavily sclerotized than basal part.Aedeagus irregularly sclerotized with narrow sclerite extending onto vesica on right to base of submedial cornutus.Vesica about as long as aedeagus with tapered cornutus about 2/3 as long as width of vesica.Female genitalia -Unknown.
Distribution and biology.Bryolymnia mixta is known only from the Patagonia Mountains in southeastern Arizona.Adults were collected in late June and mid-July.Diagnosis.Bryolymnia ensina is a medium-sized species (fore wing length: 12-14 mm), with a mainly brown or orange-brown fore wing with blackish-brown shading in the upper half of the antemedial area, in the subterminal area, especially toward the costa, and as one or two dark streaks between the reniform spot and the subterminal line.It looks like a large form of Bryolymnia biformata, but the dark shading on each side of the paler medial area is much more patchy than in B. biformata.Th e hindwing is pale fuscous basally with darker fuscous on the veins, discal spot, and on the outer 1/3 of the wing.Bryolymnia ensina appears to be most closely related to B. mixta on the basis of the male genitalia, but the details of male genitalia and the mainly pale medial area of the fore wing allow the two species to be readily separated.Male genitalia.Th e male genitalia are similar to those of B. mixta, but in B. ensina the uncus is swollen dorsally toward the base, not cylindrical, the dorsal process from the basal arm of the clasper is rounded and tapered, not basally constricted, the digitus is fused to the inner surface of the valve, not free of the valve and apically spatulate as in B. mixta, and the clasper has a short dorsal branch about 1/3 from the base.Th e sclerotized extension of the aedeagus is broad, heavily sclerotized, and apically rounded, unlike the slender extension in B. mixta.Th e vesica has no cornutus, unlike many other species of Bryolymnia, and the apical half of the vesica is covered with minute spines.Female genitalia.Th ese are most likely to be similar to the female genitalia of B. mixta, which are unknown.Th e ductus bursae is short, 1/3 as long as the corpus bursae, and the posterior 2/3 is heavily sclerotized and fused into a broad posteriorly convex ostium bursae with a small pouch on each side of the ostium.Th e corpus bursae is oblong, 3 × as long as wide, with two long thin signa.
Distribution and biology.Bryolymnia ensina occurs in coniferous forests from southeastern Arizona (Huachuca Mts.) and southwestern New Mexico (Pinos Altos Mountains) southward in the Sierra Madre Occidental to the State of Durango.Collecting dates range from mid-June to mid-July.Etymology.Th e name biformata is from Latin and refers to the two color forms of this species.

Bryolymnia biformata
Diagnosis.Bryolymnia biformata can be recognized by the contrast between the gray-brown or pale yellow-brown shading of most of the medial area compared to the blackish-brown shading in the basal and subterminal areas and in the medial area between the reniform spot and subterminal line.Th e contrast in the shading is much more sharply defi ned than in either B. ensina or B. mixta, although B. biformata went unrecognized for many years because the two forms were treated as additional forms of B. ensina.Th e genitalia are signifi cantly diff erent from those of B. ensina.Th e male genitalia are similar to those of B. ensina in that the uncus is swollen basally, the clasper has a dorsal process near the base, and the sclerotized lobe on the basal extension of the clasper is tapered and rounded.However, the vesica in B. biformata does not resemble that of B. ensina but is more like that of B. mixta in having a postmedial cornutus, although much stouter and angled, and no apical spines.Th e valve is tapered apically, unlike that of either B. ensina or B. mixta.In the female genitalia the ductus bursae is almost as long as the corpus bursae, and the ostium bursae, unlike that in B. ensina, lacks a pouch on each side of the ostium bursae.
Description.Adults.Male and female similar in size, color, and maculation.Head -Male antenna with fl agellomeres very slightly swollen laterally; setae tending to group into two or three clusters on each side of each segment.Palpi and head mainly of broad apically serrated dark-brown scales with a speckling of gray scales on sides of palpi.Th orax -Scales similar in shape to those on head, dark brown to blackish brown, similar in color to basal area of fore wing; a slightly divided dorsal tuft on metathorax.Legs: Covered with gray-brown scales with white band in middle of tibiae and at apices of tarsal segments.Tibia without spiniform setae.Wings: Fore wing length: 11-12 mm.Dorsal fore wing ground color variable by area; basal and antemedial areas and medial area between reniform spot and postmedial line dark brown to blackish brown; remainder of medial area usually either pale orange brown, reddish brown, with a few intermediates; subterminal and terminal areas a mixture of brown, hoary gray, and blackish brown with hoary gray predominating in subterminal area, brown in terminal area, and blackish brown mainly forming a patch in subterminal area adjacent to costa; basal, antemedial and postmedial lines black, narrowly bordered by a pale line; subterminal line diff use; reniform, orbicular, and claviform spots, and medial line, a shade darker than medial area ground color, not contrasting, but reniform spot sometimes with partial pale-gray outline; fringe dark gray brown with tiny yellow dots at base between wing veins.Dorsal hindwing pale fuscous with darker on discal spot, veins, postmedial line, and wing margin.Fringe pale buff with fuscous medial line.Male genitalia -Uncus mainly cylindrical, slightly swollen dorsally toward base, tapered to spine-like apex.Valve strap-like, mainly parallel sided but slightly tapered apically; apex of valve with corona on outer margin; digitus mainly fused to costal margin of valve except for short tapered apex; apical part of clasper slender and upcurved, with dorsal process near base; dorsal part of basal rod extending from clasper to dorsal margin of valve a large rounded process extending beyond dorsal margin of valve; apical part of sacculus beyond post-medial suture slightly less heavily sclerotized than basal part.Aedeagus irregularly sclerotized, with narrow sclerite extending onto vesica on right as far as base of cornutus.Vesica about as long as aedeagus, with large basally-stout, apically-angled postmedial cornutus.Female genitalia -Corpus bursae oval, slightly longer than ductus bursae with ductus seminalis at posterior end of corpus bursae.Ductus bursae with posterior 3/4 heavily sclerotized, slightly wider posteriorly, with irregularly sclerotized suture separating ductus bursae from heavily sclerotized rounded plate of ostium bursae.
Distribution and biology.Bryolymnia biformata is known only from the Huachuca, Patagonia, and Santa Rita Mountains in southeastern Arizona.Adults have been collected between mid-June and late July.Ferris & McFarland, 2007 Figs 17-19, 31, 39 Bryolymnia anthracitaria Ferris & McFarland 2007: 196 Type material.Holotype ♂.Arizona, Cochise Co., Huachuca Mts., Ash Canyon, 31°23.27'N,110°14.Diagnosis.Bryolymnia anthracitaria can be recognized by its grayish-brown forewing with almost all of the basal and antemedial areas blackish brown and with a thick blackish-brown streak extending from the orbicular spot (in some specimens from the antemedial line) to the terminal line.Th e reniform and orbicular spots are outlined in black and the postmedial line is black but the lines are thin, so they tend to be partially covered and obscured by the prominent dark streak.Th e subterminal line is pale and usually obscure, being most evident near the costa because of black shading in the upper part of the subterminal area adjacent to the line.Fore wing length ranges from 12-16 mm with males averaging 1.7 mm larger than females (Ferris and McFarland 2007).In the male the hindwing is pale fuscous with darker fuscous on the veins, discal spot, and wing margin.In the female the hindwing is dark fuscous, slightly paler toward the wing base.Male genitalia.Th ese are similar to those of Bryolymnia biformata, but in B. anthracitaria the dorsal surface of the clasper is serrated, the clasper is without a basal dorsal process, the digitus is much larger and widens apically into a plate that is serrated on both sides, and the vesica is more globular with two stout cornuti.Female genitalia.Th ese also are similar to those of Bryolymnia biformata, but in B. anthracitaria the corpus bursae is more elongated and the ductus bursae is completely sclerotized and markedly expanded anteriorly and posteriorly.Diagnosis.Bryolymnia semifascia can be recognized by the brown fore wing with blackish-brown shading and streaks, especially toward the hind margin of the wing, and usually with a contrasting white patch in the lower part of the medial area.Th e white patch, the best character to recognize this species, may consist of only a few scattered white scales, but in most specimens it covers most, sometimes all, of the lower half of the medial area.In many specimens light-gray scales form partial highlights to the otherwise obscure reniform and orbicular spots, and the postmedial and subterminal lines.Forewing length ranges from 11 to 14 mm.Th e hindwing is pale fuscous with darker fuscous on the veins, discal spot, postmedial line and wing margin with that of the female only slightly darker than in the male.Male genitalia.Th ese are most similar to those of Bryolymnia anthracitaria.As in B. anthracitaria the dorsal surface of the clasper is serrated, the clasper is without a subbasal process, and the digitus is large and heavily sclerotized.Th e main diff erences from the genitalia of B. anthracitaria are that the dorsal surface of the digitus is densely hairy, the clasper in much smaller, and the vesica has only a single slender tapered cornutus.Female genitalia.Th e corpus bursae is rounded, about 0.6 × as long as the heavily sclerotized, parallel-sided ductus bursae, and gives rise to a rounded lightly sclerotized appendix bursae ventrally at the junction with the ductus bursae.Th e sclerotized plate in the ostium bursae has a wide V-shaped notch on the posterior margin.

Bryolymnia anthracitaria
Distribution and biology.Bryolymnia semifascia occurs from northern Colorado and southern Utah southward to southeastern Arizona and south-central New Mexico.Adults have been collected from mid-June to mid-September in conifer forests.Other material examined and distribution.USA: California: Alameda Co., Los Angeles Co., San Diego Co., Sonoma Co.
Remarks.Th is species previously has been associated with the genus Cryphia Hübner in the Bryophilinae.Chris Schmidt, working with DNA barcodes of Cryphia species noted that Cryphia viridata associated with Bryolymnia.Examination of the male genitalia confi rmed this generic association.Th e male genitalia are more weakly sclerotized than most other Elaphriini, but the membranous transverse postmedial suture of the sacculus, and the membranous weak area on the costal margin of the valve associate this species with the Elaphriini, and the sclerotized plate on the dorsal rod extending from the base of the clasper to the dorsal margin of the valve associates it with Bryolymnia within the Elaphriini.
Diagnosis.Bryolymnia viridata diff ers from all other congeners in the genus in having the fore wing ground color almost entirely green.Th ere usually is some fuscous shading over the ground, especially in the subterminal area, but not in the terminal area.In most specimens there is dark fuscous or black shading between the reniform and orbicular spots, on the costa above the orbicular spot, representing the upper part of the medial line, and on the costal part of the subterminal area.A rounded spot in the fold, possibly derived from the claviform spot, looks like a second orbicular spot and often has a similar dark central dot.Th e terminal line is a series of black dashes between the wing veins, and this in combination with a white fringe with black wedge-shaped spots at the end of the veins, give the fringe a checkered pattern.Fore wing length ranges from 12 to 15 mm.Th e hindwing is white with a variable amount of fuscous on the veins, discal spot, postmedial line and terminal line.Females tend to have more fuscous shading in these areas than do males.Male genitalia.Th e valve is more oval than in other species of Bryolymnia, broader mesially and tapered apically with the cucullus not diff erentiated from the rest of the valve and the apical corona weak and confi ned to the dorsal half of the outer margin.Th e clasper has a slight subbasal tooth dorsally and ventrally, and the sclerotized dorsal lobe on the dorsal rod extending from the base of the clasper is somewhat infl ated and lobed, not a fl at rounded plate as in most other species.Th e uncus is broad mesially and tapered toward each end, unlike other species.Both the aedeagus and vesica are unarmed, the latter being about as long as the aedeagus, slightly swollen at the base, then tapered to the apex.Female genitalia.Th e corpus bursae is oval, about 2 × as long as the ductus bursae, without signa, and with the ductus seminalis on a small appendix bursae posteriorly on the left.Th e posterior 1/3 of the ductus bursae is heavily sclerotized and this expands into a broad U-shaped ostium bursae with a very slightly convex posterior margin.
Distribution and biology.Bryolymnia viridata occurs in western California from Sonoma County north of San Francisco southward to San Diego County.Adults have been collected late May to mid-October.
Type material.Holotype ♂.Location of type unknown.Type locality: USA, California, San Diego.