Grammia figurata, (DRURY)

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 : 538-543

GRAMMIA FIGURATA (DRURY) ( FIGS 17, 52 , 83 , 110 )

Bombyx figurata Drury, 1773 : pl. 12, f. 4

Euplagia ceramica Hübner, [1820] : 180.

Arctia celia Saunders, 1863: 59 . syn. nov.

Arctia snowi Grote, 1875: 197 .

Arctia excelsa Neumögen, 1883: 70 .

Arctia phyllira ab. lugubris Hulst [1887]: 182. Apantesis figurata var. preciosa Nixon, 1911: 232 . Apantesis figurata (Drury) ; Franclemont, 1983: 117. Grammia figurata (Drury) ; Ferguson & Opler, 2006: 11.

Type material: Bombyx figurata : described from an unstated number of specimens, although a female is illustrated; the type material is lost; the type locality is Virginia [ USA] .

Euplagia ceramica : location of type material not known.

Arctia celia : described based on a male specimen from Toronto, ON, [ Canada] [CNC, examined]. The holotype is typical of the northern populations of figurata , which exhibit yellow hindwings and often have the subterminal forewing band well developed. The name celia has been variously applied to northern G. figurata and G. franconia sp. nov., but the celia holotype is clearly the same species as the northern figurata forms.

Arctia snowi : the female type specimen is in the USNM (type # 33673) [not examined]. Type locality: KS [ USA] .

Arctia excelsa : male holotype from NC in USNM ( Smith, 1938a) [not examined] .

Arctia phyllira ab. lugubris: this name was proposed to describe a variant with entirely black hindwings, and is therefore an unavailable infrasubspecific name.

Apantesis figurata var. preciosa : this name was proposed to describe a form of figurata with yellow hindwings, and is therefore an unavailable infrasubspecific name.

Diagnosis: Most specimens of G. figurata can easily be recognized by absence or reduction of the forewing subterminal band, and the fusion of the hindwing postmedial and subterminal spots into one relatively even-bordered marginal band. In the south-eastern Great Plains where the range of G. figurata overlaps with G. f-pallida , G. figurata can be separated with the characters given under G. f-pallida . Some variants of G. figurata from northern populations (Great Lakes/New England) can be similar to G. franconia and G. williamsii , see Diagnosis under those species.

Description: Head – Palps black, frons and vertex pale to pinkish buff; male antennae moderately bipectinate, rami averaging 5.10 ¥ 10 - 1 mm, (N = 9); female antennae slightly biserrate; dorsal scales dark brown to black; eyes well developed, mean diameter 1.04 mm (N = 9). Thorax – Vestiture black with pale to yellowish buff borders on vertex, patagia, and tegulae; black ventrally, base of coxa with yellowish buff scales; coxa and femur black, tibia and tarsus variable, pale buff dorsally, black ventrally, or entirely black. Abdomen – Dorsal ground colour yellow, orange or pink, slightly paler near apex, or entirely black, particularly females; medial and lateral markings black; pale buff ventrally with variable lateral black markings, or entirely black. Forewing – Male forewing length averaging 15.77 mm (N = 6 males); dark brown to black dorsally, vein lines absent; bands pale buff to ivory; medial band usually well developed, not extending beyond postcubital stripe; postcubital variable in length, extending only to postmedial or slightly past subterminal; medial band well developed, with slight angle at M 3 or straight; subterminal band absent to well developed; fringe and anal margin varying from pale buff to dark brown and combinations thereof; costa dark or variably pale-lined, up to 2/3 of the distance from base; ventral markings similar, but slightly paler and with distinct yellowish cast; wing shape in females slightly more elongate, but otherwise similar. Hindwing – Ground colour highly variable, ranging from yellow to scarlet and vivid pink; black markings variable in extent but postmedial and subterminal elements nearly always confluent to form a broad marginal band; medial spot usually prominent, antemedial spots varying from well developed and streaked basad, to absent; hindwing occasionally entirely black, or with very restricted coloured medial area; similar ventrally, but dark markings with a paler yellowish cast; sexes similar. Male genitalia – Distal portion of valve gradually tapering to rounded apex; clasper reduced, median ridge moderately developed; uncus broad-based, process evenly tapered to point, 2.5 ¥ as long as width of base; juxta two ¥ wider than height; aedeagus with dorsad curve at 2/3 of distance beyond base; medial chamber of vesica slightly longer than wide, minutely scobinate; distal chamber kidney-shaped, twice as long as wide, coarsely scobinate; diverticula moderately developed. Female genitalia – Ductus bursae unsclerotized; corpus bursae pear-shaped, 3.3 ¥ width of ostium bursae; signa round, relatively small, averaging 2.2 ¥ 10 - 1 mm, coarsely scobinate; posterior apophysis slightly longer than papillae anales.

Biology: Grammia figurata is bivoltine in northern OH ( Rings & Metzler, 2002), but most records from the northern part of the range are for May through to June. In the south-west (KS, OK, TX), records range from April to October, undoubtedly representing more than one yearly generation.

Distribution: Southern ON and NH south to GA, west to CO and TX ( Fig. 110 ). Reports of this species from QC ( Handfield, 1999) are probably referable to G. franconia .

Molecular variation: Eleven specimens from eight localities exhibited five haplotypes in the eastern lineage, with a mean and maximum intraspecific divergence of 0.49 and 1.52%, respectively ( Table 2). Four haplotypes clustered together (EA30: EA28, Fig. 135), and also included G. virguncula and G. f-pallida haplotypes.

Remarks: This taxon displays remarkable phenotypic variation across its range, but there are no reliable data to suggest that more than one species is involved. The northern forms of G. figurata ( Fig. 17b, c), with yellow or often melanic hindwings, and welldeveloped forewing subterminal bands, differ considerably from the pink hindwing/unbanded forewing forms (east-central and southern USA), yet many populations (e.g. southern ON) display individual variation to the point where no two specimens are exactly alike. This has resulted in the long-standing misapplication of the name celia , as discussed above under ‘ Type material’ and Grammia franconia .

The taxonomic status of some CO populations is not yet fully resolved, but I am retaining them as G. figurata until more data become available; this pertains to the slightly larger taxon with a fully developed subterminal forewing band and hindwings with more discrete rather than banded markings (Morgan, Douglas, and Arapahoe Counties, CO); females resemble males in this taxon (A. Warren, pers. comm.), suggesting that it is closely related to or conspecific with G. figurata , not G. f-pallida .

GRAMMIA F- PALLIDA (STRECKER) ( FIGS 18, 53 , 84 , 111 )

Arctia f-pallida Strecker, 1878 ; Ann. Rep. Chief of Engineers, App. SS: 1860

Arctia quadranotata Strecker, 1878 ; Proc. Davenport Acad. Nat. Sci, 2: 271, pl. 9, f. 6 [Note: published in October 1878, after the June–July issue in which the description of f-pallida Strecker 1878 appeared ( Smith, 1938a)]

Apantesis sociata Barnes & McDunnough, 1910: 149 .

Apantesis moierra Dyar, 1914: 161 .

Apantesis f-pallida (Strecker) ; Franclemont, 1983: 117.

Apantesis quadranotata (Strecker) ; Franclemont, 1983: 117.

Apantesis sociata (Barnes & McDunnough) ; Franclemont, 1983: 117.

Grammia f-pallida (Strecker) ; Ferguson & Opler, 2006: 11.

Type material: Arctia f-pallida : male holotype from Rio Navajo , [Archuleta Co.,] Colo [rado], [ USA], in FMNH according to Smith (1938a) [not examined], but this specimen is currently not in the FMNH numbered colour slides of type specimens, and there are no types missing judging by the complete numbering sequence of slides. It therefore appears that the f-pallida type has been missing for some time .

Arctia quadranotata : described from ‘several examples’ from TX according to the original description, although Strecker (1899) subsequently listed a single female as the type material. The female syntype labelled ‘Dallas/ Tex. Boll’, ‘orig. type.’, ‘ Arctia Quadranotata / Streck/ Dallas, Texas./ orig. Type, J. Boll’, ‘ Arctiidae / genitalia slide/ No. 1096′, Lepidoptera Type/ Photograph No. 197/ Field Museum’, ‘ Lectotype ♀ / det. A. Watson 1966’, ‘ CNC / colour slide/ 662212’, ‘ LECTOTYPE / Arctia / quadranotata Strecker / B.C. Schmidt, 2009’ is hereby designated as lectotype to ensure the stability of this name [ FMNH, photograph examined].

Apantesis sociata : female holotype from Fort Wingate , New Mex [ico, USA] [ USNM, examined] .

Apantesis moierra : female holotype from Dallas , TX, [ USA], USNM type # 19075 [ USNM, not examined] .

Diagnosis: The reduced forewing banding pattern, with complete or nearly complete absence of the w-shaped forewing subterminal band is characteristic of only G. f-pallida and most G. figurata . Compared to G. figurata , G. f-pallida is generally smaller (forewing length <34 mm vs.> 34 mm), and the sexes have a dimorphic forewing pattern in G. f-pallida (female G. f-pallida always have nearly or entirely dark forewings), whereas female G. figurata exhibit a pattern similar to that of males. Although the differences in wingspan between G. f-pallida and G. figurata can be subtle, the difference in body size is more marked, reflected by the smaller eye diameter in G. f-pallida , which is less than 0.90 mm (mean 0.81 mm), greater than 0.90 mm (mean 1.04 mm) in G. figurata . Grammia f-pallida and G. figurata remain distinct where the ranges overlap in the south-eastern Great Plains; the two species are sympatric and synchronic at Fort Supply, OK, flying together in late August.

Description: Head – Palps black; frons and vertex pale to reddish buff, or black and buff laterally; male antennae moderately bipectinate, rami averaging 4.43 ¥ 10 - 1 mm, (N = 6); dorsal scales dark brown to black; eyes well developed, mean diameter 8.14 ¥ 10 - 1 mm (N = 6). Thorax – Vestiture black with pale to yellowish buff borders on vertex, patagia, and tegulae; females usually entirely dark brown – black; dark brown to black ventrally, base of coxa with pinkish buff scales; coxa and femur black, tibia and tarsus variable, pale buff dorsally, black ventrally, or entirely black. Abdomen – Dorsal ground colour pink, not noticeably paler near apex; medial and lateral markings dark brown to black; entirely black ventrally. Forewing – Male forewing length averaging 13.9 mm (N = 6 males); dark brown to black dorsally, vein lines absent; bands pale buff to ivory; medial and postmedial band and postcubital stripe well developed in males; subterminal band absent, or reduced to slight v-shaped bar at M 3; antemedial band sometimes expressed as costal bar; fringe, anal margin, and costa dark brown, anal margin rarely pale at base; forewing entirely dark brown in females, or with bands reduced to one or two bars or spots in discal cell; ventral markings similar, but slightly paler and with yellowish cast; wing shape in females slightly more elongate, but otherwise similar. Hindwing – Ground colour pinkish red, rarely yellow (CO); black markings restricted to broad, relatively smoothbordered margin consisting of merged postmedial and subterminal elements; medial discal spot prominent, antemedial spots usually absent, sometimes present and streaked basad; similar ventrally, but dark markings with a paler yellowish cast; sexes similar. Male genitalia – Distal portion of valve gradually tapering to rounded, slightly pointed apex; clasper reduced, median ridge moderately developed; uncus broadbased, process evenly tapered to point, two¥ as long as width of base; juxta two ¥ wider than height; aedeagus with dorsad curve at 2/3 distance beyond base; medial chamber of vesica slightly longer than wide, minutely scobinate; distal chamber kidneyshaped, twice as long as wide, coarsely scobinate; diverticula well developed. Female genitalia – Ductus bursae unsclerotized; corpus bursae pear-shaped, three ¥ width of ostium bursae; signa round, relatively large, averaging 3.0 ¥ 10 - 1 mm, coarsely scobinate; posterior apophysis equal in length to papillae anales.

Biology: Collection dates for adults indicate that G. f-pallida is bivoltine in at least parts of its range, with dates from the Davis Mountains in western TX ranging from late April to early May and July through to August. South-eastern CO/western OK records are for late August, and NM and UT for June/July and late August. In central CO it flies in June and July. The early stages are undescribed.

Distribution: Distributed from south-eastern UT and CO south through to eastern AZ, NM, and eastern TX ( Fig. 111 ). Ferguson et al. (2000) also give a record for west-central Nevada, and it almost certainly occurs in Mexico, although I have seen no specimens from there.

Molecular variation: Grammia f-pallida samples consisted of two specimens from two localities (CO, OK) with a single haplotype of the Eastern lineage ( Table 2, Fig. 135). This haplotype differed by only two base pairs from a haplotype shared by G. virguncula and G. figurata ( Fig. 135).

GRAMMIA INCORRUPTA (HY. EDWARDS)

( FIGS 19, 54 , 85 , 112 )

Arctia incorrupta Hy. Edwards, 1881 ; Papilio 1: 38 Arctia ochracea Neumögen, 1883 ; Papilio 3: 71 (preocc. by Stretch, 1872)

Arctia nevadensis var. sulphurica Neumögen, 1885 ; Entom. Amer. 1: 93

Apantesis geneura incorrupta (Hy. Edwards) ; Franclemont, 1983: 117.

Grammia nevadensis complex [in part] Ferguson et al., 2000: 48, 130.

Grammia incorrupta (Hy. Edwards) ; Ferguson & Opler, 2006: 11, Ferguson & Schmidt, 2007: 42.

Type material: Arctia incorrupta : A lectotype for A. incorrupta was designated by Ferguson & Opler (2006), but no details pertaining to the specimen are provided, other than the fact that it was collected at Prescott, AZ (as were three of the original syntypes, according to the description). All specimens from Prescott were examined among Ferguson’s material at the USNM, and no specimen bears any label indicating that Ferguson had intended it as the lectotype. According to the original description, A. incorrupta was based on three female specimens from Prescott, AZ and one male from [The] Dalles, OR ( Edwards, 1881). According to Smith (1938a), there were three incorrupta specimens from the Edwards collection in the AMNH at the time, two males from ‘Arizona’ and one male from ‘[The] Dalles, Ore[gon].’ As noted by Smith (1938a), Edwards probably either mistook the AZ males as females, or the mix-up was the result of a typographical error. At any rate, all evidence suggests that the two Edwards specimens remaining at the AMNH are syntypes, one from ‘Oregon’ with a ‘No. 8130 Coll. Hy. Edwards’ label, the second from Prescott, AZ with a ‘No. 8128, Coll. Hy. Edwards’ label (S. Rab-Green, pers. comm., 2005). As the OR specimen is in all likelihood referable to Grammia nevadensis superba ( incorrupta is not known to occur in OR), I designate the AZ specimen as lectotype, bearing the labels: ‘Prescott, Arizona’, ‘No. 8128 / Collection Hy. Edwards’, ‘ Arctiidae / genitalia slide No. 1262’, ‘ Lectotype by A. Watson 1967’, ‘ LECTOTYPE / Arctia / incorrupta H. Edwards / B.C. Schmidt, 2009’.

Arctia ochracea: Unavailable primary homonym of ochracea Stretch (1872) . The holotype is the same specimen subsequently designated as the holotype of sulphurica (see below) according to Neumögen (1883), although the specimen no longer bears any labels indicating that it is the type of ochracea Neumögen.

Arctia nevadensis var. sulphurica : proposed as a replacement name for ochracea Neumögen 1883 , a homonym of ochracea Stretch (1872) (a junior subjective synonym of Grammia ornata ). The holotype of sulphurica in the USNM bears the labels ‘ Arctia / nevadensis / v. sulphurica . / Type. Neumgn.’, ‘TYPICUM/ SPECIMEN’, ‘Prescott/ Arizona.’, ‘Col./ B. Neumögen.’, ‘Type No./ 33679/ U.S. N.M.’, ‘ ♂ genit. on/ slide/ Sept. 6, 1931 / MES 1536’. Sulphurica is an unavailable infrasubspecific name, based on a colour variant.

Diagnosis: As a result of the significant phenotypic variation in G. nevadensis , G. incorrupta was long confused with this species. Most of this confusion stems from the similarity of G. n. geneura with incorrupta , as both taxa are relatively large, broad-winged and brightly coloured. Most specimens of G. incorrupta can be distinguished from G. nevadensis by the longer antennal rami, broader forewing shape, reduced hindwing markings, and the larger signa of the female bursa. Males very rarely express the alldark thorax phenotype (I have seen only three specimens like this) common in G. n. nevadensis and G. n. superba , and the entirely pale thoracic collar (patagia) seen in about 1/4 of G. incorrupta specimens does not occur in any nevadensis populations. Grammia incorrupta is multivoltine, whereas G. nevadensis has a single yearly flight. I have not been able to find consistent differences in the male genitalia. A comparison of characters is given in Table 3.

Description: Head – Frons, vertex, and palps with dark brown to black vestiture, frons and vertex varying from dark with buff borders to (rarely) entirely pale buff; eye fully developed, mean maximum diameter = 1.07 mm (N = 6); male antenna strongly bipectinate, longest branches averaging 7.55 ¥ 10 - 1 mm, or 4.31¥ intersegmental distance (N = 12; AZ, NM, CO); female antenna slightly to moderately biserrate; antenna dark-scaled in both sexes, occasionally with pale buff scales along vertex of shaft and rami. Thorax – Patagia, tegulae, and vertex of thorax usually dark-brown to black bordered with pale buff, patagia entirely pale buff in about 1/4 of specimens; yellow ventrally, legs dark brown with variable amounts of pale buff. Abdomen – Ground colour orange-pink, rarely yellow-orange, anal tuft pale buff; medial spots wider in females, particularly on last two segments; yellowish-buff to pale buff ventrally, dark brown to black markings consisting of two rows of spots, bordered distally at each segment margin by pale buff; females usually entirely or predominantly dark brown–black ventrally. Forewing – Mean forewing length = 18.6 mm (N = 9); usually with complete set of transverse bands, as in G. nevadensis , basal and antemedial bands rarely reduced or absent; bands pale buff to yellowish buff, rarely pinkish buff; fringes concolourous with bands, lacking dark scales; anal dash absent. Female forewing bands more yellowish than males. Hindwing – Ground colour pink to yellowish pink, very rarely yellow; markings most often restricted to three postmedial spots and traces of subterminal spots at margin, but never confluent; antemedial and medial spots generally absent or reduced, more prevalent in females; females with more vivid, brighter hindwing pink; ventral pattern similar, but colours less vivid. Male genitalia – Distal portion of valve varying from broad to slender, but not constricted near middle, and apex rounded; clasper reduced to moderately developed; uncus broad-based, process evenly tapered to point; juxta wider than long; aedeagus with dorsad curve at 2/3 distance beyond base; distal chamber of vesica kidney-shaped, scobinate. Female genitalia – Ductus bursae unsclerotized; corpus bursae with three round and one oval signa, roughly equal in size; appendix bursae with angled elbow, not evenly coiled, finely but conspicuously scobinate; posterior apophysis about 1.5 ¥ longer than length of papillae anales.

Biology: This species has two flights annually, with adult collection dates ranging from late April to early October. The peak of each flight varies according to local climate and seasonal rains ( Singer, 2000), peaking in June and October in south-eastern AZ ( Singer, 2000). Grammia incorrupta inhabits arid grasslands and open woodlands associated with the mountain ranges of the south-western USA and Mexico. Unlike species of the nevadensis group, females of incorrupta come to light.

Larvae feed on a large variety of primarily herbaceous, flowering plants, belonging to at least 80 species in 50 families ( Singer, 2000). Larvae parasitized by tachinid flies are able to ‘self-medicate’, increasing their uptake of toxic plants in response to parasitism ( Bernays & Singer, 2005).

Distribution: Grammia incorrupta occurs from southern CO and south-east KS south through AZ, NM, and western TX into Mexico (states of Aguascalientes and Nuevo Leon), and west to south-eastern CA ( Fig. 112 ).

Molecular variation: Four specimens of G. incorrupta from four localities (TX, AZ, NM, CO) all exhibited unique haplotypes, with a maximum intraspecific divergence of 2.3% ( Table 2). Grammia incorrupta haplotypes formed a discrete cluster (Incorrupta lineage, Fig. 133 ). Haplotypes of G. incorrupta were at least 3% divergent from G. ursina ( Table 2, Fig. 133 ).

Remarks: I have not seen specimens of incorrupta and nevadensis from the same site, although the ranges of the two should overlap in southern CO, southern UT, and possibly southern Nevada. Grammia incorrupta and G. bowmani fly together at Colorado National Monument, Mesa County, CO. The black thorax form is very rare in incorrupta ( Fig. 19B), and I have seen it only from Yavapai County, AZ and McGaffey, NM.

Figures

Figure 133. Neighbour-joining tree of cox1 ‘barcode’ haplotypes of Grammia species and Holarctia obliterata. Two-letter codes correspond to the species name in a given lineage. The Kodara Lineage includes Grammia quenseli (QE1), Grammia williamsii (WI1), and Grammia virgo (VI14) haplotypes. Haplotypes of the Western and Eastern lineages are shown separately in Figs 134 and 135, respectively. Numbers in parentheses indicate number of sampled specimens.

Figures 52–57. Genitalia of male Grammia, ventral aspect of valve and right lateral aspect of aedeagus (scale bar = 1 mm). 52, Grammia figurata (Drury). 53, Grammia f-pallida (Stkr.). 54, Grammia incorrupta (Hy. Edwards). 55, Grammia allectans Ferguson. 56, Grammia ursina Schmidt. 57, Grammia nevadensis (G. & R.).

Figures 79–86. Genitalia of female Grammia, ventral aspect (scale bar = 1 mm). 79, Grammia virguncula (W. Kby.). 80, Grammia quenseli (Paykull). 81, Grammia margo Schmidt. 82, Grammia franconia (H. Edw.). 83, Grammia figurata (Drury). 84, Grammia f-pallida (Stkr.). 85, Grammia incorrupta (Hy. Edwards). 86, Grammia ursina Schmidt.

Figures 108–113. Distribution of examined specimens of Grammia.