Speyeria cybele ( Fabricius, 1775 )

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 : 13-14

Speyeria cybele ( Fabricius, 1775)

( Figure 4B , 8 )

Papilio Nymphalis Phaleratus Cybele Fabricius, 1775: 516 .

Papilio Nymphalis Phaleratus Daphnis Cramer, 1775: 89 ; 1777: 152.

Argynnis Cybele aberration Baal Strecker, 1878: 111.

Speyeria cybele (Fabricius) [ dos Passos and Grey 1945a].

Common names. Great spangled fritillary, cybele fritillary, yellow-banded silver wing.

Type deposited. Holotype (female) (= neotype of dos Passos and Grey 1947; see Miller and Brown 1981) at Natural History Museum, London ( Figure 8 ).

Type locality. The original description ( Fabricius 1775) did not contain a collection date, sex of specimen, or series data; “Habitat in America”. Defined by dos Passos and Grey (1947) based on alleged holotype as New York City, New York County , New York .

Type label data. “ cybele, Fab., Syst. Ent. P. 516 n. 311 (1775), United States; Papilio Cybele Fabr. Sp. Ins. No. 477; NEOTYPE, Papilio Nymph. Phalerat. Daphnis ? Martyn, designated by dos Passos and Grey 1947, p. 6”.

Identification, taxonomy, and variation. There are approximately 10 described subspecies in the cybele complex. Adult wingspan ranges from 65-105 mm. The western races show a sexual dimorphism in which the ground color of the male is bright orange and the female is yellow with darker scales located near the base. Some, such as Speyeria cybele leto (Behr) , exhibit sexual dimorphism with males being bright orange and females being nearly white. Older literature, as well as contemporary works, treat ‘ leto ’ as a distinct species ( Holland 1931; Edwards 1864b; Scudder 1875; Howe 1975; Kondla 2004 ). Eastern and western populations reportedly intergrade or show mixed wing characteristics where they meet in Alberta and Montana ( Glassberg 2001a) . Speyeria cybele bear silver spots on the ventral hindwings, but these spots are reduced compared to other Speyeria species. The ventral discal area is typically brown and the submarginal band is wide and yellowish in color. Males have prominent sex scaling on along forewing veins. The eyes on living adults are yellow-green ( Glassberg 2001a). Female genitalia in S. cybele differ from most other Speyeria by having a secondary bursal sac, closely allying S. cybele with S. diana and S. aphrodite . The male genitalic armature bears a hooked uncus ( Figure 4B ), similar to those in S. aphrodite , S. diana , S. idalia , and S. nokomis , which differs from the relatively unhooked, less excavate uncus of the remaining species of Speyeria ( Figure 4C ). Eggs are light yellow when first deposited and turn pale brown after 3-4 days. Duration from oviposition to larval eclosion is reportedly 12-17 days ( Edwards 1880) or 22-23 days indoors ( Ross and Henk 2004). Mature larvae are approximately 51 mm in length, are typically chocolate-brown on the ventral surface, and bear dorsal black spines that are red-yellow to orange at the base. There is also a row of gray spots located dorsally. The larval head capsule is orange above and black below. Pupae are mottled dark brown, occasionally with reddish-orange over the wing cases. The anterior abdominal tubercles are usually black or black and yellow in color. Duration of the pupal stage is 16 to 20 days in eastern S. cybele ( Edwards 1880) .

Range. Forms of S. cybele occur from the east coast to the west coast in the United States and Canada, south to northeastern California, New Mexico, and eastward to central Arkansas (reportedly common in Clay, Greene, and Craighead Counties in northern Arkansas ( Lavers 2006) and the northern portions of Georgia, Louisiana, and Mississippi. The range of nominate S. cybele includes much of the eastern United States, where it is still considered common. It was once considered common in areas such as Staten Island, but was reportedly rare in the early 1970’s ( Shapiro and Shapiro 1973). Records for S. cybele exist as far south as Mississippi (Lafayette County-Mather 1966) and Florida ( Kimball 1965; Heppner 2003). Some S. cybele forms may be declining in western North America because of habitat changes such as the loss of habitat ( Opler and Wright 1999). Howe (1975) reported a decline in eastern Kansas S. cybele and noted considerable fluctuations in its numbers from one season to the next. It has recently been reported in Cowley County, Kansas and McCurtain County, Oklahoma (Tuttle 2007) and populations should continue to be monitored in these regions.

Life history. Habitat includes Transition to Canadian Zone in moist deciduous woods and moist meadows, conifer forest openings, aspen-lined streams or glades, valleys, prairies, and along roadsides. Females mate immediately after emerging in May and June but do not commence oviposition until August or September, strongly suggesting reproductive diapause ( Sims 1984). Eggs are typically laid singly near dead or dying Viola spp. and unfed first instar larvae hibernate; however, Scudder (1889) noted that eggs are also laid upon the leaves and stalks of the host plant. First instars commence feeding the following spring. Adults are swift fliers and males patrol all day while seeking females; females carry males while mating. Males typically frequent flower heads more often than females; the females remain hidden and rarely venture out into the open. Ross (2002, 2004) noted that dead or decoy adult S. cybele placed on nectar sources attracted additional S. cybele individuals as well as other butterfly species. Flight period is mid-June through mid-September.

Larval host plants. Viola rotundifolia , V. paplionacea , V. palustris , V. adunca , V. adunca variation bellidifolia, V. sororia , V. canadensis ( Scott 1986b; Swengel 1997; Robinson et al. 2002; Heppner 2003). Western S. c. leto also reportedly feeds on V. glabella , V. nuttalli , and V. sempervirens ( Warren 2005) ; also V. adunca (Jonathan Pelham, pers. comm.).

Adult food sources. Butterfly milkweed, Asclepias exaltata , common milkweed, Buddleia , ironweed, thistles, dogbane, knapweed, vetches, red clover, purple coneflower, Joe-Pye weed, and black-eyed Susan, also occasionally feed on dung ( Howe 1975; Scott 1986b; Broyles and Wyatt 1991; Gochfeld and Burger 1997; Opler and Malikul 1998; Ross 1998 ; Foote 2002; Ross 2002 ; Douglas and Douglas 2005). Rudolph et al. (2006) listed several plant species as primary nectar sources in Arkansas, including Asclepias tuberosa , Monarda fistulosa , Cirisium carolinianum , Echinacea purpurea , Carduus nutans , and Liatris squarrosa .

Figures

Figure 4. Male genitalic armature. A) Speyeria idalia (locality unknown). B) Speyeria cybele (Nebraska). C) Speyeria hesperis (aedeagus removed) (Idaho). D) Digitus located on left valva, Speyeria edwardsii (Nebraska). 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.