Speyeria diana ( Cramer, 1777 )

Dunford, James C., 2009, Taxonomic overview of the greater fritillary genus Speyeria Scudder and the atlantis - hesperis species complexes, with species accounts, type images, and relevant literature (Lepidoptera: Nymphalidae), Insecta Mundi 2009 (90), pp. 1-74 : 11-13

Speyeria diana ( Cramer, 1777)

( Figure 5 A-B, 7)

Papilio Nymphalis Phaleratus Diana Cramer, 1777: 4 , 148.

Argynnis diana (Cramer) [ Holland 1931].

Speyeria diana (Cramer) [ dos Passos and Grey 1945a].

Common names. Diana fritillary, Great Smokies fritillary, Ozark Diana fritillary.

Type deposited. Holotype (male) (see Miller and Brown 1981) at Natural History Museum, London ( Figure 7 ).

Type locality. The original description (Cramer 1977) did not contain a collection date, sex of specimen, or series data; “en Virginie ”. Defined by dos Passos and Grey (1947) based on putative holotype (see Miller and Brown 1981) as Jamestown, James City County, Virginia.

Type label data. “ex collection Tring Museum, ex collection Felder, ex collection M. J. C. Sylvius van Lennep”.

Identification, taxonomy, and variation. Adult wingspan ranges from 88-112 mm. Both sexes are distinctive and superficially unlike other greater fritillaries. Adults are sexually dimorphic with the male’s general appearance orange and black and the female’s blue and black ( Figure 5 A-B). A similar sexual dimorphism occurs in western North America with S. nokomis and with Eurasian Argynnis that range through regions of higher rainfall and higher summer temperatures ( Hovanitz 1963b). Males bear black wing bases and are orange distally while females are black basally and bluish distally. The veins in the forewing of the male are thick and dark. Speyeria diana also lacks silver spots on the ventral hindwings, the discal bars are completely obliterated, and the postmedian and submarginal spots are greatly reduced, distinguishing them from most other Speyeria . A rare form of the female occurs that has green instead of blue on the hindwings ( Opler and Krizek 1984). No subspecies has been designated for S. diana ; however, there is some variability in individuals, but this is not abundantly apparent at the population level. Clark and Clark (1951) reported differences in wing facies due to elevational changes in Virginia populations. Female genitalia in S. diana differ from most other Speyeria by having a secondary bursal sac, closely allying S. diana with S. cybele and S. aphrodite . In the male the digitus is distinct, widening distally bearing an abrupt ventral angle with an outline unique to S. diana . Females are especially fecund with well over a thousand ova recorded ( Ross and Henk 2004). Eggs are light yellow when they are deposited, and turn gray by day four or five, reflecting the color of the developing larva ( Allen 1997; Ross and Henk 2004). Mature larvae are approximately 65 mm in length, velvety black to purple with rows of black spines that are red to orange basally. Dorsal spines are proportionately longer than those located laterally. There occasionally is a double row of white spots located dorsally. The larval head capsule is orange above and black below, but is more angulate than those of closely related S. cybele and S. aphrodite . Pupae are approximately 30 mm in length, mottled light brown and red, and bear tubercles on the abdomen. Duration of the pupal stage is approximately 20 days.

Range. It is currently restricted to the interior highlands of Arkansas, Oklahoma and Missouri ( Carlton and Nobles 1996; Rudolph et al. 2006). It is also known in the southern Appalachians from western Virginia, West Virginia to northeast Georgia and Alabama ( Scott 1986b; Moran and Baldridge 2002). Moran and Baldridge (2002) recorded it from 14 different Arkansas counties, 11 of these representing county records, indicating that it is more widespread than previously thought. It was extirpated in southeastern Virginia in about 1951 ( Opler and Krizek 1984; Scott 1986b), and is considered uncommon or extirpated in many other parts of its range. Historical populations in the Midwest and the Virginia Piedmont were extirpated in the 1800s ( Opler and Krizek 1984; Rudolph et. al. 2006). Dos Passos and Grey (1947) listed records from the following states: Pennsylvania, Virginia, West Virginia, North Carolina, South Carolina, Georgia, Alabama, Kentucky, Indiana, Tennessee, Illinois, Arkansas. The Illinois records are reported from Clay, Union, and Vermilion counties, where it may have had a regular occurrence in the southern third of the state ( Irwin and Downey 1973).

Life history. Habitat is mostly Upper Austral to Transition Zone in deciduous and pine woodland near streams, rich forested valleys and mountainsides. Clark and Clark (1951) noted that thick undergrowth, usually with alders and rhododendrons, is usually present. Females will walk on the forest floor, laying single eggs on dead leaves and twigs near Viola spp. , mostly in late summer. Larvae emerge in the late fall and hibernate until the following spring when they commence feeding on violet leaves and flowers. Adult males begin flying one week earlier than females and patrol woodland habitats. Females likely mimic Battus philenor (L.), Limenitis astyanax (F.), and some papilionids in various parts of the species range ( Scudder 1889; Poulton 1909; Ehrlich and Ehrlich 1961; Adams and Finkelstein 2006). However, Hovanitz (1963b) hypothesized that there may an environmental relationship affecting wing coloration and patterns by noting that they may be correlated to the high humidity and temperatures where S. diana occurs. Flight period is mid-June through early August, rarely into September. In Arkansas’s Ouachita Mountains, male S. diana emerge as early as late May and females emerge approximately 7-10 days later ( Rudolph et al. 2006). Females have been observed as late as mid October in northern Georgia ( Adams and Finkelstein 2006). Speyeria diana is of conservation concern and the cause of extirpations and range contractions are likely due to habitat alteration ( Allen 1997), harvest of old growth forests (Hammond and McCorkle 1983), strip mining (Vaughan and Shepard 2005a), and loss of nectar plants ( Moran and Baldridge 2002; Rudolph et. al. 2006). The Xerces Society currently lists S. diana as vulnerable (Vaughan and Shepard 2005a).

Larval host plants. Viola papilionacea , V. cucullata , V. cornuta , V. sororia ; also partially reared on Vernonia noveboracensis (Compositae) (Scudder 1975; Tietz 1972; Scott 1986b; Robinson et al. 2002).

Adult food resources. Reported to visit milkweeds including swamp milkweed and butterfly weed, ironweed, red clover, dung, carrion, damp soil, wads of grass, vomitus of coyotes, and human sweat ( Opler and Krizek 1984; Krizek 1991; Opler and Malikul 1998; Rudolph et al. 2006). Rudolph et al. (2006) listed several plant species as primary nectar sources in Arkansas including Asclepias tuberosa , Monarda fistulosa , Cirisium carolinianum , and Echinacea purpurea ; Asclepias syriaca was recorded as a nectar source in western Virginia ( Krizek 1991).

Figures

Figure 5. Habitus images of adult Speyeria spp., each image with dorsal (left) and ventral (right) view. A) Speyeria diana, male. B) Speyeria diana, female. C) Speyeria zerene gunderi (Comstock), male; Nevada. D) Speyeria callippe harmonia dos Passos and Grey, male; Nevada. E) Speyeria mormonia artonis (Edwards), male; Nevada. Images by James C. Dunford.

Figure 6-10. Type images for Speyeria spp.; dorsal, ventral, and label images. 6) Speyeria idalia neotype, described as Papilio Nymphalis Phaleratus Idalia, male, AMNH. 7) Speyeria diana holotype, described as Papilio Nymphalis Phaleratus Diana, male, BMNH. 8) Speyeria cybele neotype, described as Papilio Nymphalis Phaleratus Cybele, female, BMNH. 9) Speyeria aphrodite neotype, described as Papilio Nymphalis Phaleratus Aphrodite, male, AMNH. 10) Speyeria nokomis neotype, described as Argynnis Nokomis, male, AMNH. Images 6 and 9-10 by James C. Dunford; 7-8 by Kim Goodger Buckmaster.