Grammia behrii, (STRETCH)

Schmidt, B. Christian, 2009, Taxonomic revision of the genus Grammia Rambur (Lepidoptera: Noctuidae: Arctiinae), Zoological Journal of the Linnean Society 156 (3), pp. 507-597 : 549-552

GRAMMIA BEHRII (STRETCH) ( FIGS 24, 58 , 88 , 117 )

Arctia behrii Stretch, 1872: 75 , pl. 3, f. 11–12.

Arctia shastaensis French, 1889a: 35 .

Arctia shastanensis Hampson, 1901 [misspelling] Callarctia nevadensis Smith, 1938a: 129 .

Apantesis nevadensis (Grote & Robinson) [in part]; Franclemont, 1983: 117.

Grammia behrii (Stretch) ; Ferguson et al., 2000: 48; Ferguson & Opler, 2006: 11; Ferguson & Schmidt, 2007: 46.

Type material: Arctia behrii : the original description was based on two male and two female syntypes from Downieville , [Sierra Co.], CA [ USA]. The types are lost, and a neotype was designated by Ferguson & Schmidt (2007). The type locality is Jackson Cr [eek], 39 50′N 120 39′W, Plumas Co [unty], CA , USA [ CNC] .

Arctia shastaensis : based on one male specimen that was missing the hindwings and abdomen. The French types may be destroyed ( Lafontaine et al., 1991). The illustration in French (1889b: 162) however leaves little doubt as to the identity of this name. Type locality: Upper Soda Springs, Siskiyou County, near Mt. Shasta [CA, USA] [type lost?].

Diagnosis: Most specimens of G. behrii are easily recognized by the very wide, complete antemedial band on the forewing, all-dark thorax, bright pinkishred hindwing with markings reduced or absent, and the yellowish forewing bands. Additionally, it is larger overall compared to G. nevadensis , with which G. behrii is most likely to be confused. Some forms of G. ornata can be similar to G. behrii , but G. behrii lacks the hindwing antemedial markings of G. ornata , has longer antennal rami, and flies in late summer not spring.

Description: Head and Thorax – Entirely dark brown, almost black; Male antennae with dark brown and scattered yellow scales along rami; longest rami averaging 7.01 ¥ 10 - 1 mm, 4.06¥ longer than intersegmental distance (N = 6). Female antennae slightly biserrate, less so than in G. nevadensis . Thorax dark brown ventrally, vestiture of coxae and femora usually at least partly yellow, legs dark brown to black, may be marked with yellow. Abdomen – Dorsally concolorous with hindwing ground colour, with mid dorsal and lateral segmental black spots, sometimes merging into a solid stripe; underside of abdomen black, segments bordered distally with transverse yellowish bands. Anal tuft black or mixed black and yellowish. Forewing – black, with the usual pattern of light bands typical of the nevadensis complex, except that their colour is bright ochreous yellow, and the antemedial band is usually conspicuously wider than other bands, up to 5 mm wide; The medial band may be missing or reduced to one or more small spots at or near the position of the discal spot in other moths. The antemedial, medial (if present) and postmedial bands commonly cross the postcubital stripe to the anal margin, but the basal band never does. Fringes blackish, yellow tipped. Hindwing – Ground colour saturated orange-pink, rarely yellow. Dark markings usually reduced to two to three postmedial spots, and subterminal band reduced to narrow marginal black marks; Fringe yellowish. Sexes similar in wing pattern, females exhibiting brighter, more saturated colours as in other species of the nevadensis group. Male genitalia – Distal portion of valve gradually tapering to rounded apex; clasper poorly developed; median ridge moderately developed; uncus broad-based, process evenly tapered to point, 2.5¥ as long as width of base; juxta 1.75¥ wider than height, dorsal concavity broad, relatively deep; aedeagus with dorsad curve at 2/3 distance beyond base; medial chamber of vesica short, about as long as wide, minutely scobinate; distal chamber kidney-shaped, twice as long as wide, coarsely scobinate; diverticula moderately developed. Female genitalia – Ductus bursae unsclerotized; corpus bursae slightly pear-shaped, three¥ width of ostium bursae; signa round or ellipsoid, averaging about 2.5 ¥ 10 - 1 mm, coarsely scobinate; posterior apophysis slightly longer than papillae anales.

Biology: Adults have been captured between early August and late September, with most records from mid September. Females do not come to light, and are partially or entirely diurnal. Two female specimens from Los Angeles County were collected in flight in mid-afternoon. Larvae have been collected on Lotus humistratus E. Greene (Fabaceae) and Amsinckia (Boraginaceae) in Los Angeles County. Grammia behrii inhabits dry, low- to mid-elevation lithosol flood plains and balds in the Siskiyou – northern Sierra Nevada ranges, but is absent from serpentine barrens (J. Troubridge, pers. comm.).

Distribution: Occurs from Portland, OR south to Los Angeles and Kern Counties, CA, with most records from the Siskiyou and Sierra Nevada ranges ( Fig. 117 ). Ferguson et al. (2000) also gave a record for west-central ID.

Molecular variation: Three specimens exhibited three haplotypes, all of which were paraphyletic with respect to G. nevadensis ( Table 2, Fig. 134 ). Grammia behrii haplotypes showed a maximum of 0.9% intraspecific divergence, including one shared haplotype with G. nevadensis from central Washington (BE1; Fig. 134 ).

Remarks: Grammia behrii is a rarely collected species that was for many years relegated to the synonymy of G. nevadensis , but as discussed by Ferguson & Schmidt (2007), is a distinct species. The two are sympatric in Siskiyou County (Hornbrook) in northern CA, and possibly also in the Greenhorn Mountains, Kern County.

GRAMMIA BRILLIANS SCHMIDT SP. NOV.

( FIGS 27, 60 , 119 )

Type material: Holotype ( Fig. 27) – ♂. UT, [Garfield Co.], Bryce Canyon , Water Canyon, 6800′, 10.vii.2005, S. Mueller, DNA voucher # BCS-1087, ‘barcode’ voucher BCSC-244 [ USNM] . Paratypes – 2 ♂♂. Same data as holotype [ USNM, CNC]

Etymology: The name brillians reflects the bright, saturated colours of the hindwing.

Diagnosis: The wing pattern, colour, and male antennal structure are most similar to G. bowmani , but brillians lacks antemedial and medial markings on the hindwings (usually present in bowmani ), and the postmedial forewing band curves basad near the costa, not straight or nearly so as in bowmani . Internally, the medial and distal chamber of the male vesica are longer and more elongate in brillians , covered in smaller, more numerous spinules; the median ridge is reduced in brillians , well developed in bowmani . In comparison, Grammia williamsii tooele , like bowmani , also has a straight postmedial forewing band, shorter male antennal rami, and fewer forewing bands. Grammia nevadensis is sympatric with G. brillians , but flies later in the season and has longer male antennal rami, and sympatric G. nevadensis populations have much paler and washed-out colours, and are unlikely to be confused with G. brillians .

Description: Male:Head – Frons, vertex, and palps entirely dark brownish black; eyes fully developed; male antennae moderately bipectinate, rami averaging 5.42 ¥ 10 - 1 mm, 3.4¥ longer than intersegmental distance (N = 2); antenna with dark brown dorsal scales, or dark brown with buff scales near apex of rami. Thorax – Vestiture entirely dark brownish black dorsally and ventrally; a few light brown scales at base of forecoxa; legs entirely dark brown with buff apex of tibia. Abdomen – Ground colour pink dorsally with buff scales at apical tuft, medial and lateral markings brownish black; predominantly black ventrally, segments with buff lateral and distal margins. Forewing – Mean forewing length 16.1 mm (N = 2 males); ground colour dark brownish black, fringe variable, entirely pale buff or black and buff; bands ivory in colour, antemedial band incomplete, extending toward inner margin beyond postcubital stripe; medial and postmedial bands curved basad near costa, extending beyond postcubital stripe when well developed; similar ventrally, with colours less bright and a slight yellowish flush near wing base. Hindwing – Ground colour bright orange-pink, patterned with black; antemedial and medial markings absent, or medial markings reduced to a single, indistinct discal spot; postmedial spots variable, but relatively reduced, not confluent with subterminal markings; fringes variably black and buff; at anal angle; ventral and dorsal pattern similar, colours less bright ventrally and with a slight yellow flush near wing base. Male genitalia – Distal portion of valve tapering evenly to more or less rounded apex; median ridge poorly developed, median ridge reduced; uncus long evenly tapered, typical of genus; juxta broad nearly twice as wide as high; aedeagus relatively broad, distal end curving dorsally, and opening to the right as in other Grammia ; vesica with kidney-shaped distal chamber, relatively elongate medial chamber; medial and distal chamber scobinate, spinules relatively small.

Female: unknown.

Biology: Adults come to light, and the type series was collected on 10 July, the only known flight date. Nothing else is known of its biology.

Distribution: Known only from the type locality in Bryce Canyon National Park, UT ( Fig. 119 ).

Molecular variation: The single sequenced specimen exhibited a unique haplotype, at least 0.8% divergent from other haplotypes in the western lineage ( Table 2; Fig. 134 ).

Remarks: Although I hesitate to describe a species from only three specimens taken at a single locality, particularly in such a phenotypically variable group, it is clear from the morphological and molecular data that this taxon is distinct from other species in the nevadensis group. Grammia brillians may be endemic to southern UT; a similar pattern of endemism is seen in the noctuids Protogygia pectinata Lafontaine & Copablepharon opleri Lafontaine ( Lafontaine, 2004) . The complex geological history of the Colorado Plateau and the edges of the Great Basin are likely to have played an important role in the evolutionary radiation of the nevadensis group.

GRAMMIA FERGUSONI SCHMIDT SP. NOV. ( FIGS 35, 68 , 92 , 127 )

Type material: Holotype ( Fig. 35A) – ♂: CA, Inyo County, Mono Pass , 12 000′, 1.viii.1978, C.D. Mac- Neill [ CNC] . Paratypes – 6 ♂♂ 6 ♀♀. CA: Tuolumne County, Sonora Pass, 9623′, 20.vii.1953, T. W. Davies [ LACM]; Tuolumne County , Leavitt Peak , 11 050– 12 000′, 29.viii.1967, P. A. Opler , 1 ♂ [ UCB]; Tuolumne County, near Sonora Pass, Leavitt Peak , north slope, 10 500′, 22.viii.1975, J. Wiseman , 1 ♂, [ LACM]; Mono County, Lake Marie, 12.viii.1944, R. Mattoni , 2 ♀♀ [ LACM]; Mono County, White Mtns., Blanco’s Corral , 10 000′, 7.vii.1953, D.D. Linsdale , 1 ♂ [ UCB]; Inyo County, Mono Pass , 12 000′, 1.viii.1978, C.D. MacNeill , 1 ♀ [ CNC]; same locality, 10.viii.1958, J.M. Burns , 1 ♂ [ UCB] same locality and date, C.D. Mac- Neill , 1 ♀ [ UCB]; Mono County, White Mtns, Barcroft , 13 000′, 28.vii.1958, C.D. MacNeill & W.W. MacGuire , 1 ♂ 1 ♀ [ USNM]; Fresno County, Mono Pass , 37°25.44′N 118°46.36′W, 12 040′, larva collected 18.viii.1998, reared on Taraxacum, L. Crabo , 1 ♀ [Lars Crabo personal collection].

Etymology: The name honours the late Douglas C. Ferguson, whose research contributed significantly to the state of knowledge of North America’s arctiid moths.

Diagnosis: Grammia fergusoni is relatively distinctive among Grammia , and somewhat resembles a small, ‘pointy-winged’, brightly coloured G. nevadensis or G. behrii . The antennae are buff-coloured, not brownblack as in most nevadensis group species, and the male antennal rami are shorter than in G. nevadensis . The forewing is more pointed and elongate apically, and the hindwing more triangular. Internally, the male distal portion of valve is slightly hooked, not blunt or rounded. This species is diurnal, and is endemic to the alpine zone of the central Sierra Nevada.

Description: Head – Vestiture of frons pale buff with black lateral border, vertex and palps black with pale buff borders; eyes reduced, averaging 5.8 ¥ 10 mm at greatest diameter; male antennae moderately bipectinate, rami averaging 3.3 ¥ 10 - 1 mm, two¥ longer than intersegmental distance (N = 3); antenna with pale buff dorsal scales, apical third of antenna black; female antennae slightly bipectinate. Thorax – Vestiture black with narrow, pale buff borders on vertex, patagia, and tegulae; dark brown to black ventrally with yellowish buff central tuft at base of forecoxae; legs predominantly pale buff, femur dark brown basally with yellowish buff apical half. Abdomen – Ground colour pinkish-orange, pale buff near apex; medial and lateral markings dark brown to black; pale buff ventrally, dark brown to black markings consisting of two rows of spots; females with ventral spots expanded and confluent, segments pale bordered distally. Forewing – Male forewing shape elongated toward apex, mean forewing length 14.4 mm (N = 3), females considerably larger with somewhat broader wing shape, forewing length 17.7 mm (N = 2); ground colour dark chocolate brown, fringe pale buff; banding pattern complete or with antemedial band reduced to costal spot and line caudad of postcubital stripe; medial band often wider than postmedial, the latter curved basad near costa as in G. nevadensis group. Hindwing – Ground colour bright pinkishorange, patterned with black; antemedial and medial markings often extending basally as a dash in males, reduced or absent in females; postmedial and subterminal markings usually confluent in males, discrete in females; fringe pale buff to orange, particularly near anal angle; ventral pattern similar to dorsum, colours slightly paler. Male genitalia – Distal portion of valve narrowing gradually, apex slightly hooked, point of hook directed ventrad; median ridge and ventral process well developed; uncus broad-based, process evenly tapered to point; juxta 1.5–2¥ wider than long; aedeagus with dorsad curve at 2/3 distance beyond base; vesica overall most similar to G. nevadensis , curving dorso-cephalad; distal chamber of vesica relatively elongate and kidney-shaped, scobinate, spinules relatively large. Female genitalia – Not examined.

Biology: Adult dates of capture are primarily in August, with a few in mid July. This is a diurnal species that flies in subalpine and alpine habitat.

Distribution: Known only from the central Sierra Nevada and White Mountains in CA ( Fig. 127 ).

Molecular variation: Not examined.

Remarks: The small size, narrow wing shape and antennal structure and colour suggest a close affinity to G. blakei , although the vesica structure is unlike this species, and is more typical of G. nevadensis .

Figures

Figure 134. Neighbour-joining tree of Western lineage haplotypes of Grammia species. Multiple symbols per row indicate multiple species and/or samples for a given haplotype. Squares indicate species with haplotype complements primarily or exclusively in the Western lineage, circles indicate predominantly Eastern lineage species. Grammia virgo haplotype VI15 (triangle) is the sole G. virgo haplotype outside of the Virgo lineage.

Figures 58–63. Genitalia of male Grammia, ventral aspect of valve and right lateral aspect of aedeagus (scale bar = 1 mm). 58, Grammia behrii (Stretch). 59, Grammia eureka Ferguson & Schmidt. 60, Grammia brillians Schmidt. 61, Grammia bowmani Ferguson & Schmidt. 62, Grammia williamsii (Dodge). 63, Grammia elongata (Stretch).

Figures 87–95. Genitalia of female Grammia, ventral aspect (scale bar = 1 mm). 87, Grammia nevadensis (G. & R.). 88, Grammia behrii (Stretch). 89, Grammia williamsii (Dodge). 90, Grammia elongata (Stretch). 91, Grammia blakei (Grt.). 92, Grammia fergusoni Schmidt. 93, Grammia ornata (Pack.). 94, Grammia hewletti (B. & McD.). 95, Grammia complicata (Walker).

Figures 114–119. Distribution of examined specimens of Grammia.

Figures 64–68. Genitalia of male Grammia, ventral aspect of valve and right lateral aspect of aedeagus (scale bar = 1 mm). 64, Grammia yavapai Schmidt. 65, Grammia favorita (Neum.). 66, Grammia blakei (Grt.). 67, Grammia cervinoides (Stkr.). 68, Grammia fergusoni Schmidt.

Figures 126–131. Distribution of examined specimens of Grammia.