Grammia phyllira, (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 : 521-523

GRAMMIA PHYLLIRA (DRURY) (FIGS 6, 43, 75, 99)

Bombyx phyllira Drury, 1773 : pl. 7 f. 2.

Geometra b-ata Göze, 1781: 428.

Arctia oithona Strecker, 1878: 131 .

Arctia rectilinea French, 1879: 45 .

Arctia dodgei Butler, 1881: 136 .

Arctia rectilinea var. conspicua Stretch, 1906: 120 , pl. 8 fig. 2.

Apantesis favorita ab. favoritella Strand, 1919: 284. syn. rev.

Apantesis phyllira (Drury) ; Franclemont, 1983: 117.

Apantesis oithona (Strecker) ; Franclemont, 1983: 117. Grammia phyllira (Drury) ; Ferguson & Opler, 2006: 11.

Grammia phyllira (= avoritella [Strand]); Ferguson & Opler, 2006: 11. [misspelling]

Type material: Bombyx phyllira : described from a female specimen [lost] from New York .

Geometra b-ata: described from an unknown number of specimens; type locality and type material not known.

Arctia oithona : A female syntype [FMNH, photo examined] labelled ‘ Arctia oithona / Dallas, Tex. Streck./ orig. Type; J. Boll’, ‘30.’, ‘11. V.’, ‘172.’, ‘orig./ type’, LECTOTYPE ♀ / det. A. Watson 1966’, ‘ Lepidoptera Type/ Photograph No. 196/ Field Museum’, C.N.C./ colour slide/ 662209′, ‘ LECTOTYPE / Arctia / oithona Strecker / B.C. Schmidt, 2009’ is hereby designated lectotype to stabilize the name. The type locality is Dallas, [TX, USA].

Arctia rectilinea : described from two male and two female syntypes, presumed lost. The type locality is Carbondale [Illinois?] according to Smith (1938a) .

Arctia dodgei : based on a female holotype [BMNH, photograph examined]; the holotype bears no locality data, although the original description states that dodgei may be a ‘Mexican representative’ of phyllira ; phyllira is not known to occur in Mexico.

Arctia rectilinea var. conspicua : types probably lost; type locality not stated; conspicua is an unavailable infrasubspecific name, referring to variants with partially developed forewing vein lines.

Apantesis favorita ab. favoritella: Strand (1919) based this name on Hampson’s (1901: 406) description of Arctia favorita ‘Ab. 1’, which he describes as having the ‘forewing bands absent, except the postmedial and subterminal’ ( Hampson, 1901). As this form is not known to occur in G. favorita , and because Hamspon’s concept of G. phyllira included not only G. phyllira (s.s.) but also G. figurata and G. f-pallida , it is most likely that his ‘Ab. 1’ describes the unlined form of G. phyllira ; favoritella is therefore transferred to the synonymy of phyllira .

Diagnosis: The lined form of G. phyllira prevalent throughout the Great Plains could only be mistaken for G. parthenice , but G. phyllira can easily be distinguished by the pinkish cast on the forewing underside (cream in G. parthenice ), and the straight, not angled postmedial line. The unlined form is similar to Grammia placentia and some specimens of G. figurata , but G. figurata has yellow-buff ventral forewing bands, not pink; G. placentia can be distinguished by its entirely dark, not spotted ventral abdomen, larger size, wider space between the medial and postmedial forewing band at the costa, and females which lack forewing bands.

Description: Head – Palps black; frons and vertex pale buff; male antennae strongly bipectinate, rami averaging 5.84 ¥ 10 - 1 mm, (N = 6); dorsal scales dark brown to black, sometimes with scattered buff scales; eyes well developed, mean diameter 1.07 mm (N = 6). Thorax – Vestiture black with pinkish buff to pale buff borders on vertex, patagia, and tegulae; sides of thorax dark brown ventrally, with orange-pink vestiture at base of coxae; legs buff-coloured and dark brown to black dorsally, predominantly black ventrally. Abdomen – Dorsal ground colour pinkish red, apex often pale buff; medial and lateral markings black; pale to yellowish buff ventrally, lateral black spots not confluent. Forewing – Male forewing length averaging 16.9 mm (N = 6 males); dark brown to black dorsally, vein lines complete, reduced or absent; bands rich creamy buff to pinkish buff; medial, postmedial, subterminal bands and postcubital stripe generally well developed, antemedial reduced to thickened bar at costa or absent entirely; postmedial not angled at M 3; transverse bands not extending beyond postcubital stripe; fringe entirely cream, costa variably cream and dark; anal margin with relatively broad cream line; ventral markings similar, but dark colours slightly paler, and light bands with distinct yellow-pink hue. Hindwing – Ground colour pink to orange-pink; antemedial spots lacking, medial spot absent to well developed; postmedial spots relatively large, but not confluent with reduced subterminal band; fringe pink and pale buff; ventral dark markings slightly paler but otherwise similar to dorsum. Male genitalia – Distal portion of valve tapering to pointed, slightly rounded apex; clasper moderately developed, median ridge moderately developed; uncus broad-based, process evenly tapered to point, two¥ as long as width of base; juxta 1.5¥ wider than long, dorsal margin with broad u-shaped concavity; aedeagus with dorsad curve at 2/3 distance beyond base; vesica with basal and medial chamber approximately equal in length and width, not scobinate; distal chamber medium-sized, twice as long as width of medial chamber, kidney-shaped, finely scobinate; vesica as long as or slightly shorter than aedeagus. Female genitalia – Ductus bursae unsclerotized, slightly narrowed medially; corpus bursae globose, three¥ width of ostium bursae; signa round to slightly elliptical, relatively large, coarsely scobinate; coil of appendix bursae elbowed; posterior apophysis 1.5–2¥ longer than papillae anales.

Biology: Adults have been collected from March into October in the southern USA, indicating two or possibly three generations. Bivoltine as far north as SD, IL, NY, and southern ON with flights in May–June and August–September. Prairie province and central ON records indicate one peak flight in mid July. Grammia phyllira occurs in dry, open woodland and grassland. Although common in much of its southern range, this species becomes very localized further north, occurring only in sandy prairie habitats in western Canada.

Distribution: Found from QC ( Handfield, 1999) and New England south to FL, west to TX, CO, and AB ( Fig. 99 ).

Molecular variation: Eight specimens of G. phyllira from five localities (ON, KY [2], AB, CO) exhibited seven haplotypes distinct from other Grammia haplotypes by at least 9%. Divergence among G. phyllira haplotypes was high, with a maximum of 2.8% ( Fig. 133 , Table 2). Although intraspecific variation was high compared to that of other monophyletic Grammia species clusters, there was no indication that molecular variation was geographically structured, with highly divergent haplotypes co-occurring in the Ottawa, ON region (PY5 and PY6: Fig. 133 , Appendix S1). Haplotypes did not cluster according to wing phenotype (lined vs. unlined), with PY1 to PY4 haplotypes corresponding to lined forms, and PY5 to PY7 being unlined forms, although all samples of lined forms were most similar to each other ( Fig. 133 ).

Remarks: Grammia oithona was for a time considered specifically distinct from G. phyllira ( Franclemont, 1983) , based on the presence of forewing vein lines in G. oithona and absence thereof in G. phyllira . It is now clear that both forms are present in most populations, with varying frequency. A reared series of specimens from a female specimen of the lined form show specimens varying from unlined to lined and partly lined [Chicago, IL; CNC]. Over 95% of specimens from Great Plains populations (AB, MB, KS, OK, northern TX) are of the lined form, whereas the unlined form predominates in eastern North America; ON and IL populations are predominantly unlined or partially lined, and north-western GA populations are slightly more than half unlined (J. Adams, pers. comm.), and lined forms are extremely rare in FL.

Although it is clear that lined and unlined forms per se do not represent two different species because reared F 1 siblings exhibit both forms (B. C. Schmidt, unpubl. data), the possibility that two species (both with lined and unlined forms) are currently included under G. phyllira should not be ruled out; the Great Plains populations ( oithona ) are on average smaller with a more elongate wing shape, and the divergence in molecular variation between eastern and western populations is larger than the average intraspecific variation within other monophyletic mtDNA Grammia lineages. Molecular and phenotypic variation of geographically intermediate populations needs to be evaluated.

GRAMMIA PLACENTIA (J.E. SMITH)

(FIGS 7, 44, 76, 100)

Phalaena placentia Smith, 1797: 129 , pl. 65.

Arctia flammea Neumögen, 1881a: 9 .

Apantesis placentia (Smith) ; Franclemont, 1983: 117. Grammia placentia (Smith) ; Ferguson & Opler, 2006: 11.

Type material: Phalaena placentia : described from an unstated number of specimens, although only a female is illustrated; the Smith types are lost; the type locality is GA, [ USA] .

Arctia flammea : described from a female holotype from Indian River , FL, [ USA] [ USNM, not examined] .

Diagnosis: The wing pattern and colour is similar to G. phyllira , but G. placentia has broader, larger wings, the last abdominal segment is entirely dark not black and pale, and the roughly triangular area formed by the forewing medial and postmedial band is much wider at the costa, three to four¥ the distance at the costa than at the postcubital versus approximately two¥ in G. phyllira . Female G. placentia resemble those of G. f-pallida , but have discrete hindwing spots rather than a broad, even marginal band.

Description: Head – Palps black; frons and vertex creamy buff; male antennae strongly bipectinate, rami averaging 7.17 ¥ 10 - 1 mm, (N = 5); female antennae slightly bipectinate; dorsal scales dark brown to black; eyes well developed, mean diameter 1.16 mm (N = 5). Thorax – Vestiture black with creamy buff to pinkish borders on vertex, patagia, and tegulae; females usually entirely dark brown–black; entirely dark brown to black ventrally; legs dark brown to black, base of forecoxa bright pink-red in males, entirely dark in females. Abdomen – Dorsal ground colour deep pinkish red, last segment entirely black; medial and lateral markings black; entirely dark brown to black ventrally in both sexes. Forewing – Male forewing length averaging 19.6 mm (N = 5 males), relatively broad, more so than other Grammia ; dark brown to black dorsally, vein lines absent; bands creamy buff; medial, postmedial, subterminal bands and postcubital stripe well developed in males; females entirely dark brown or with medial and postmedial reduced to one to two pale bars or dots, rarely with vestigial, broken bands; fringe, anal margin, and costa cream; ventral markings similar, but dark colours slightly paler and with an overall pink-red cast. Hindwing – Ground colour deep pinkish red; black markings highly reduced, but medial discal spot prominent; postmedial spots discrete, small; subterminal band thin, or absent entirely; fringe pink and dark brown; ventral dark markings slightly paler and with pinkish cast. Male genitalia – Distal portion of valve elongate, tapering to rounded apex; clasper poorly developed, median ridge reduced; uncus broad-based, process evenly tapered to point, 1.5 ¥ as long as width of base; juxta 1.5 ¥ wider than long, dorsal margin with shallow, broad u-shaped concavity; aedeagus with dorsad curve at 2/3 distance beyond base; vesica with basal and medial chamber approximately equal in length and width, not scobinate; distal chamber large, three ¥ longer than width of medial chamber, kidneyshaped, scobinate; vesica slightly longer than aedeagus. Female genitalia – Ductus bursae unsclerotized, slightly narrowed medially; corpus bursae globose, three ¥ width of ostium bursae; signa round to slightly elliptical, relatively large, coarsely scobinate; coil of appendix bursae elbowed; posterior apophysis 1.5–2¥ longer than papillae anales.

Biology: Adult collection dates range from March to October, with abundance peaks in May and July, indicating at least two annual generations. Grammia placentia occurs in dry, sandy open wooded areas, primarily pine barrens of the Atlantic coastal plain. Both sexes come to light.

Molecular variation: Six specimens from four localities (GA [3], FL) exhibited four haplotypes, with a maximum intraspecific difference of 0.7% ( Table 2). Grammia placentia formed a monophyletic, highly divergent lineage with respect to all other Grammia , with a minimum interspecific divergence of 6% ( Table 2).

Distribution: Grammia placentia has a restricted distribution, occurring only in the south-eastern USA from NJ to FL. In the NJ pine barrens this species is rare and local in occurrence (D. Schweitzer, unpubl. data).

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 96–101. Distribution of examined specimens of Holarctia and Grammia.