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Taxonomic Revision of the Genus Grammia Rambur (Lepidoptera

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Taxonomic Revision of the Genus Grammia Rambur (Lepidoptera Zoological Journal of the Linnean Society, 2009, 156, 507–597. With 135 figures Taxonomic revision of the genus Grammia Rambur (Lepidoptera: Noctuidae: Arctiinae)zoj_496 507..597 B. CHRISTIAN SCHMIDT* Canadian Food Inspection Agency, Canadian National Collection of Insects, Arachnids and Nematodes, K.W. Neatby Building, 960 Carling Avenue, Ottawa, ON, Canada K1A 0C6 Received 25 September 2007; accepted for publication 25 June 2008 The genus Grammia Rambur (Lepidoptera: Noctuidae: Arctiinae) is revised to include 37 species, using an integrative taxonomy approach based on morphological, ecological, behavioural and molecular traits. Six species, Grammia margo sp. nov., Grammia yukona sp. nov., Grammia brillians sp. nov., Grammia fergusoni sp. nov., Grammia yavapai sp. nov., and Grammia ursina sp. nov. and three subspecies, Grammia speciosa celineata ssp. nov., Grammia virgo gigas ssp. nov., and Grammia nevadensis vivida ssp. nov. are described. Neotypes are designated for Callimorpha parthenice Kirby, Callimorpha virguncula Kirby, Arctia williamsii Dodge, and Arctia edwardsii Stretch. Lectotypes are designated for Euprepia gelida Möschler, Arctia speciosa Möschler, Arctia quadranotata Strecker, Arctia oithona Strecker, Arctia incorrupta Hy. Edwards, Apantesis tooele Barnes & McDunnough, and Arctia favorita Neumögen. Grammia celia (Saunders) is synonymized under Grammia figurata (Drury), and Grammia franconia (H. Edwards) is raised to species status. Apantesis favorita ab. favoritella Strand is revised to synonymy under Grammia phyllira (Drury). Holarctia M.E. Smith, previously treated as a monotypic subgenus of Grammia, is treated as a distinct genus, as is Palearctia Ferguson, stat. rev. Sequence analysis of the ‘barcode’ fragment of the mitochondrial DNA cytochrome oxidase I (cox1) gene proved to be of limited taxonomic value at the species level because of extensive gene tree nonmonophyly. Adults of all species are described in detail and illustrated, and male and female genitalia are illustrated (where known). Females of three species (Grammia favorita, Grammia brillians, and Grammia bowmani) remain unknown, and Grammia bolanderi is known only from the original description. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 507–597. ADDITIONAL KEYWORDS: Arctiidae – Arctiini – cytochrome oxidase I – DNA barcoding – Holarctia – mtDNA – Palearctia. INTRODUCTION three species occurring also in the Palaearctic. Within the Arctiinae (sensu Lafontaine & Fibiger, 2006), The genus Grammia Rambur, 1866 (Lepidoptera: Grammia are currently placed in the tribe Arctiini, Noctuidae: Arctiinae) is the most speciose of northern subtribe Arctiina (Franclemont, 1983; Ferguson, temperate tiger moth genera, here revised to include 1985; Ferguson & Opler, 2006), or subtribe Micrarc- 37 species. Family-group names of the Noctuoidea tiina (Dubatolov, 1996, 2003). Jacobson & Weller have recently been revised to reflect changes in the (2002) did not resolve the Arctiina as monophyletic phylogeny of the group (Lafontaine & Fibiger, 2006), based on morphological characters of immatures. with the Arctiidae now recognized as a subfamily of Grammia appears to be most closely related to the the Noctuidae, and previously delineated subfamilies Nearctic genera Apantesis Harris, Notarctia Smith, equal to tribes, and tribes as subtribes. This revised and Holarctia M.E. Smith, all of which have vein R nomenclature is followed herein. Nearly all Grammia 1 of the forewing arising before the distal end of the species are restricted to the Nearctic region, with discal cell (Fig. 1D, Ferguson, 1985). Members of the genera Chelis Rambur, Hyperborea Grum-Grshimailo, *E-mail: [email protected] Neoarctia Ferguson, and Holoarctia Ferguson can be © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 507–597 507 508 B. C. SCHMIDT Table 1. Taxonomic history of the genus Grammia This revision Ferguson (1985) Franclemont (1983) Forbes (1960)* Grammia Grammia Apantesis Apantesis subgenus Grammia ––– subgenus Mimarctia ––– Holarctia turbans (Grammia turbans) (Apantesis turbans) n/a* Notarctia Notarctia (Apantesis) n/a* Apantesis Apantesis (Apantesis) (Apantesis) Munroe (1941) Smith (1938a) McDunnough (1938) Neumögen & Dyar (1894) Callarctia Callarctia Apantesis Euprepia subgenus Callarctia subgenus Callarctia –‘Callarctia group’, in part subgenus Mimarctia subgenus Mimarctia –‘Mimarctia group’, in part subgenus Holarctia subgenus Holarctia –‘Orodemnias group’, in part subgenus Notarctia subgenus Notarctia –‘Cymbalophora group’, in part Apantesis Apantesis (Apantesis) ‘Apantesis group’ *Treated only north-eastern North American fauna. very similar in wing markings and coloration, but are subsequent groupings, with the exception of the more distantly related to Grammia than Notarctia Apantesis group, which was subsequently recognized and Apantesis (Schmidt, 2007). as Apantesis (Table 1). Hampson (1901) included all As a result of their bright coloration, Grammia three currently recognized genera under Apantesis, species have received attention from professional and also including the species Atlantarctia tigrina (de amateur entomologists alike, including pioneers of Villers) [as Apantesis fasciata (Esper), a junior taxonomy and entomology such as Carolus Linnaeus, synonym]. McDunnough (1938) maintained the genus Dru Drury, Johan Fabricius, and Jean Baptiste Bois- name usage of Hampson (1901) (Table 1). duval. Despite their popularity, Grammia species can The species of Grammia were first revised by be notoriously difficult to identify, as a result of Marion Smith in an unpublished doctoral thesis often considerable intraspecific variation, conserva- (Smith, 1938a), which also included species now tive variation in genitalic morphology, and similar placed in the genera Apantesis and Notarctia. This wing pattern and colour among species-groups and work was unfortunately published only as a 12-page even among Grammia and other genera such as Holo- abstract (Smith, 1938b). Smith (1938a) included the arctia Ferguson, Chelis Rambur, and Palearctia Fer- modern concept of Grammia, Notarctia, and Holarctia guson (Ferguson, 1985). The majority of the 84 names (revised status, see below) as subgenera of Callarctia proposed for the 25 valid species (as defined here) Packard, and treated Apantesis separately (Table 1). prior to 1985 apply to variants or forms. Smith (1938a) recognized 22 species of Grammia, compared to the 24 species listed by McDunnough shortly prior to her revision in his ‘Check List of the HISTORY OF GRAMMIA TAXONOMY Lepidoptera of Canada and the United States’ The first Grammia species to be named was Bombyx (McDunnough, 1938). Five taxa treated as species by virgo, by Carolus Linnaeus (1758) in his seminal McDunnough (1938) were informally synonymized Systema Naturae, and the majority of Grammia by Smith (Arctia quadranotata Strecker, Apantesis names were proposed by A. R. Grote, B. Neumögen, gibsoni McDunnough, Arctia elongata Stretch, Arctia H. Strecker, and R. H. Stretch, all of whom figure intermedia Stretch, and Arctia oithona Strecker), and prominently in the nomenclature of North American three taxa were raised to species status (Arctia quen- moths. Neumögen & Dyar (1894) included the selii ab. complicata Walker, Arctia edwardsii Stretch, Grammia-group of genera (Grammia, Apantesis and Apantesis ornata hewletti Barnes & McDun- Harris, Notarctia M.E. Smith) in their treatment nough). Smith (1938a) did not describe any new of North American moths, uniting all these in species, but, more importantly, provided the genus- one genus, Euprepia Hübner, with five constituent group names Holarctia M.E. Smith and Notarctia.To species-groups. These groups were based on wing this end, Smith’s most significant contribution to pattern similarities, and largely did not correspond to Grammia taxonomy was in defining groups based on © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 507–597 REVISION OF THE GENUS GRAMMIA RAMBUR 509 structural characters of the larvae, pupae, and adult tion, occurring from the Arctic coast to central Mexico. genitalia, her work reflected in the modern concepts Dry, sparsely treed biomes are preferred Grammia of the two Grammia subgenera (Mimarctia and habitats, including tundra, open forest, rocky slopes, Grammia), Apantesis and Notarctia, although these grasslands, barrens, sand dunes, and gravelly or genera/subgenera were not formally recognized for rocky substrates. Although most western North nearly another 50 years (Table 1). American species occur at low to moderate elevations, Shortly after Smith’s work, Eugene Munroe sum- several species occur only in alpine and arctic habi- marized Grammia, Apantesis, and Notarctia in an tats. Relatively few species have distributions exclu- unpublished MSc thesis, wherein he maintained sive to eastern North America, but these species are the generic and subgeneric usage proposed by again usually associated with dry, open forest habi- Smith (1938a, b), and further grouped the subgenus tats such as oak/pine barrens and poor or shallow Grammia (as Callarctia, subgenus Callarctia) into soils such as sand hills, beaches, granite balds, and three species groups based on general wing pattern alvars. A few species are common in mesic and similarities (Munroe, 1941). No new species or syn- wetland habitats. Unlike many other Arctiinae, onymies were proposed by Munroe, although he rec- Grammia
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