The upper surface of the forewings of both males and females is deep orange with black marks and white spots along the outer wing margins. The underside is a mirror image of the upper surface. The undersurface of the hindwing is a rich brownish-black with white spots. The upper surface of the hindwings in both sexes has an iridescent blue-black background with two rows of spots. These spot rows are one of two characteristics used to visually differentiate females from males. The females have two rows of cream-colored spots in contrast to an orange outer row and a cream-colored inner row on the males' wings. The second characteristic is size difference, with regal fritillary females larger than males. Their bodies are brownish-black.
Regal fritillary eggs are conical with irregular ridges running from the crown to the base. They are cream colored when first deposited and appear to darken slightly over time. Larvae have been described as being either velvety black with yellowish-orange blotches or yellow to yellowish orange with black blotches. There are six rows of black bristled spines. The larvae have a black middorsal line, two black transverse lines behind the dorsal and subdorsal spines, yellowish middorsal and lateral stripes, dorsal spines silverish at the base, sublateral and lateral spines orange at the base. The head is black with orangish coloration on the top rear portion. (Holland, 1931; Kopper, et al., 2002; Maynard, 1891; Nielsen, 1999; Pyle, 1995; Scott, 1986)
Scott (1986) describes the pupae as light mottled brown with pink tinge, small black spots on the wings and thorax, transverse bands of yellow on the abdomen, and short, dorsal cones. (Scott, 1986)
Historically, the range for regal fritillaries was from Nova Scotia south to western North Carolina, and westward from northern Arkansas to eastern Colorado and Manitoba, Canada. Currently, there are only two known populations remaining east of the Mississippi River. The largest of these occurs in central Pennsylvania and a second, smaller one is found in West Virginia. (Barton, 1993; Pyle, 1995; Williams, 2002)
Regal fritillaries are considered prairie specialists and in the Midwest are known to occur in a variety of grassland habitats including moist tall-grass prairies, wet fields and meadows, virgin grasslands, old fields, and floodplain forest openings and edges. (Pyle, 1995; Nielsen, 1999; Scott, 1986; Swengel, 1996; Barton, 1996; Struttman, ; Swengel and Swengel, )
Eggs are laid in the late summer through early fall and hatch about three weeks later. These first instar larvae usually eat their egg chorion before immediately crawling into the leaf litter and entering into diapause, overwintering without feeding on plant material. In early spring the larvae begin feeding on violet leaves. They will go through five molts before entering the pupal stage, which lasts about 17 days. Larval development was followed in 86 captive larvae by Wagner et al. (1997). Each of the first five instars lasted from an average of 3.6 to 6.6 days. The final instar lasted an average of 17.1 days. Males begin emerging in early June with females following ten days to two weeks later. Opler and Krizec (1984) state that emergence continues through late August, however Barton (1996) found a few freshly emerged males in early September. Female emergence appears to end in mid-July. Few males are still flying by August, and females die off by mid-September. (Weed, 1926; Opler and Krizek, 1984; Holland, 1931; Kopper, et al., 2002; Barton, 1996; Wagner, et al., 1997)
Regal fritillaries are univoltine and begin mating soon after the females emerge in mid-June. Males spend most of their time patrolling for females and Kopper (2002) observed three types of flight behavior. The first type was the actual patrolling flight where the male flew upwind from 10 cm to 50 cm above the ground for distances of 5-30 m. There was a second "tack" flight (short, rapid) that was repeated one to several times followed by the third type where the male flew up into the air for 5-10 m and flew 20-50 m downwind in fast flights between 1 and 2 m above the ground. If no females were encountered, they resumed their upwind flight. Barton (1996) observed males flying low over vegetation then dropping down and presumably searching for females around the bases of blueberry bushes Vaccinium spp. and scrub oaks Quercus ilicifolia. When two males come into each others' path, they often will spiral up into the air then continue on their patrols. They have also been observed chasing other insects while patrolling, including monarchs Danaus plexippes, black swallowtails Papilio polyxenes asterias, great spangled fritillaries Speyeria cybele and aphrodite fritillaries Speyeria aphrodite ( B.J. Barton, personal observation, Kopper et al. 2001). (Barton, 1996; Kopper, et al., 2001a; Kopper, et al., 2002)
When a female is encountered, males give chase flying a circular pattern behind them. Unreceptive females fly up to 35 m into the air, often with several males in pursuit, then drop rapidly into the vegetation (Barton 1996). Either the female effectively loses the males, or courtship rituals continue. Barton (1996) observed a successful pairing in the Pennsylvania population. A female was observed exhibiting oviposition behavior, and a male flew down and began following her, quivering his wings. She continued with the oviposition behavior for approximately 10 minutes before starting to quiver her wings at a faster rate than the male. The female crawled down into the vegetation and stopped the wing movement, again dipping her abdomen. The male positioned himself next to the female and slowly beat his wings against the female. They both took flight and landed a short distance away, where the male resumed the wing beats. Copulation then occurred and they paired for 1 hr and 43 min. At one point the pair took flight with the male carrying the female. The female was seen exhibiting ovipositing behavior prior to this event, indicating she may have previously mated. This is in contrast to the findings of Kopper et al. (2001) who state that females mate only once. (Barton, 1994; Kopper, et al., 2001b)
Regal fritillaries produce one brood per year (univoltine). Mating begins soon after females emerge and continues through early July. Females enter reproductive diapause near the end of July and become active again in late August when they begin ovipositing. Kopper et al. (2001) studied the reproductive characteristics of females in a northeast Kansas population specifically looking at reproductive diapause. By monitoring hemolymph juvenile hormone titers, ovarian development, and fat body utilization, they determined that females mate only once. Oogenesis (the formation of eggs) and fat body depletion began in late August/early September, indicating the females undergo reproductive diapause in mid-August. Oviposition begins soon after. When females are ready to oviposit, they will drop into the vegetation and walk around probing with their abdomen until the appropriate site is selected. They will then deposit the egg and fly off to find another site. (Kopper, et al., 2001b; Barton, 1996)
There is no parental care exhibited by regal fritillaries.
The longest life span recorded in the wild from mark-release-recapture studies in Pennsylvania was at least 69 days (sex unknown) (Barton 1996). A female is known to have lived in captivity for four months (Barton 1993). As the butterflies age, they experience wing wear which eventually hinders their ability to fly. This makes them vulnerable to predators and also makes travel to nectar sources difficult. Their lifespan is most likely limited by the deterioration of their bodies, and in the case of females, loss of nectar resources due to changes in the season. (Barton, 1993; Barton, 1996)
Regal fritillaries are active within their home range, nectaring and engaging in reproductive activities. They are daytime fliers and will often take cover in the shade during extremely hot weather (Barton, personal communication).
Adults of this species can travel considerable distances, but the majority stay close to their birth sites. During a 1993 mark-release-recapture study in Pennsylvania, the average distance traveled by both sexes was 2.14 miles (3.4 km) from their initial points of capture (19% of recaptured males and 5% of recaptured females actually traveled). The longest distance traveled by a male was 6.3 linear miles (10.1 km), and 4.3 linear miles (6.9 km) by a female. (Barton, 1994; Barton, 1993)
Regal fritillaries use vision to detect other animals and the world around them. They appear to be sensitive to vibrations as well. Barton (personal communcation) observed regals flushing from vegetation immediately following a loud explosion from artillery. We have no information on the chemical senses of this species, but since other species of Speyeria communicate with pheromones, adults of this species probably do too. Larval fritillaries can taste chemicals that allow them to choose their food plants. The only observed tactile communication was a male beating his wings slowly against a female's body during courtship. (Barton, 1994; Scott, 1986)
Regal fritillary larvae are violet feeders and reported to use Viola pedatifida, V. papilionacea, V. lanceolata, V. pedata, and V. pratincola. In Pennsylvania, larvae are believed to feed only on V. sagittata (Barton 1996). They are considered diurnal herbivores. (Barton, 1993; Barton, 1994; Barton, 1996; Kopper, et al., 2001a; Kopper, et al., 2002; Scott, 1986; Weed, 1926)
Adult regal fritillaries have a long proboscis that is used for inserting into flowers to obtain nectar. They feed on a variety of nectar plants, most commonly butterflyweed Asclepias tuberosa, ironweed Vernonia baldwinii, purple coneflower Echinacaea augustafolia, blazing star Liatrus spp., thistles Cirsium spp., and other milkweeds Asclepias spp. Other recorded nectar sources include Deptford pink Dianthus armeria, mountain mint Pycnanthemum spp., Joe-pye weed Eupatorium perfoliatum, boneset E. maculatum, goldenrod Solidago spp., Calyophus serulatus, Carduus nutans, Schrankia nuttalli, Liatris squarrosa, L. aspera, yarrow Achillea millefolium, dogbane Apocynum cannabinum, Centaurea maculosa, Chrysanthemum leucanthemum, and black-eyed Susaun Rudbeckia serotina. (Barton, 1993; Barton, 1994; Barton, 1996; Kopper, et al., 2002; Scott, 1986; Weed, 1926)
There is no information available on anti-predator adaptations by regal fritillaries. Adults are susceptable to predation by birds, spiders, and possibly robber flies. Known predators include the eastern kingbird Tyranus tyranus, crab spiders (family Thomisidae), and black widow spiders Latrodectus mactans. (Barton, 1996)
Regal fritillaries serve as hosts for parasitic wasps and flies, and as food for birds and spiders. They may also serve as pollinators.
There are no known adverse affects of regal fritillaries on humans.
Regal fritillaries attract ecotourists who are interested in viewing this rare species, thus bringing economic benefit to the nearby communities. The conservation efforts of groups preserving the prairie habitats where regal fritillaries occur provide places for people to visit to enjoy a snapshot of what the midwest once looked like. Research activities have enhanced our knowledge of prairie ecosystems.
Regal fritillaries are not currently on the IUCN Red List, CITES, or the US Endangered Species list; however, they are listed as endangered, threatened, or extirpated by several state governments. They are listed as endangered in the state of Michigan. In the eastern U.S. where the species is most imperiled, efforts have been underway by the Pennsylvania Air National Guard and various private organizations to protect the habitat of the population in Pennsylvania. Other conservation groups such as The Nature Conservancy are actively protecting many known sites (Barton, personal communication).
Regal fritillaries have been known by several common names, including regal silverspot butterfly, regal silverwing, and silver Argynne. Scientific names have changed as well and are as follows: Papilio idalia (Drury) 1773, Argynnis astarte (Fisher) 1858, A. ashtaroth (Fisher) 1859, A. infumata (Oberthur) 1912, A. dolli (Gunder) 1927, and A. pallida (Eisner) 1942. (Maynard, 1891; Miller, 1992)
Williams (2002) believes that ecological differentiation may exist between Midwestern populations and those in the east. The larvae in the Pennsylvania population eat Viola sagittata, a plant not used anywhere else in their range. Habitat differences exist as well, with mesic conditions found in the east and xeric in the west. Williams speculates that glaciers extending into southern Indiana and Ohio during the Pleistocene may have divided regals into eastern and western populations. The results of mtDNA sequencing analysis suggest the Pennsylvania population is a distinct lineage, and that they may have separated from the midwestern population about 400,000 years ago. (Williams, 2002)
Matthew Wund (editor), University of Michigan-Ann Arbor.
Barb Barton (author), Special Contributors.
The Nature Conservancy. "Regal Fritillary at Fort Indiantown Gap, Pennsylvania" (On-line ). Accessed 02/18/03 at http://nature.org/wherewework/northamerica/states/pennsylvania/preserves/art824.html.
Barton, B. 1993. 1993 Field study of the regal fritillary <<Speyeria idalia>> at ... (site name deleted). Report to the Department of Defense: unpublished.
Barton, B. 1996. Final report on the Regal Fritillary 1993-1995...(site name deleted). Report to the Department of Defense: unpublished.
Barton, B. 1994. The status report of the regal fritillary at ... (site name deleted). Report to the Department of Defense.
Glassberg, J. 1999. Butterflies Through Binoculars - The East. New York City, New York, USA: Oxford University Press.
Holland, W. 1931. The Butterfly Book. Garden City, NY: Doubleday, Doran, and Company.
Iftner, D., J. Shuey, J. Calhoun. 1992. Butterflies and Skippers of Ohio. Columbus, OH: Ohio Biological Bulletin Volume 9 No. 1.
Kopper, B., R. Charlton, D. Margolie. 2000. Oviposition and site selection by the regal fritillary, *Speyeria idalia*, as affected by proximity of violet host plants. Journal of Insect Behavior, 13/5: 651-655.
Kopper, B., D. Margolies, R. Charlton. 2002. Life history notes on the regal fritillary, <<Speyeria idalia>> (Drury) (Lepidoptera: Nymphalidae), in Kansas tallgrass prairie. Jounal of the Kansas Entomological Society, 74/3: 172-177.
Kopper, B., D. Margolies, R. Charlton. 2001. Notes on the behavior of <<Speyeria idalia>> (Drury) (Nymphalidae) larvae with implications that they are diurnal foragers. Journal of the Lepidopterists' Society, 54/3: 96-97.
Kopper, B., S. Shu, R. Charlton, S. Ramaswamy. 2001. Evidence for reproductive diapause in the fritillary <<Speyeria idalia>> (Lepidopter: Nymphalidae). Annals of the Entomological Society of America, 94/3: 427-432.
Maynard, C. 1891. Manual of North American Butterflies. Boston MA: DeWolf, Fiske, and Co.
Miller, J. 1992. The Common Names of North American Butterflies. Washington DC: Smithsonian Institution Press.
Miller, L., F. Brown. 1981. Catalogue/Checklist of the Butterflies of America North of Mexico - Memoir No. 2 of the Lepidopterists' Society. Lepidopterists' Society.
Nielsen, M. 1999. Michigan Butterflies and Skippers: A Field Guide and Reference. East Lansing, MI: Michigan State University Extension.
Opler, P., G. Krizek. 1984. Butterflies East of the Great Plains: An Illustrated Natural History. John Hopkins University Press.
Pyle, M. 1995. National Audubon Society Field Guide to North American Butterflies. NY: Alfred A Knopf.
Scott, J. 1986. Butterflies of North America: A Natural History and Field Guide. Stanford, CA: Stanford University Press.
Singer, M., D. Vasco, C. Parmesan, C. Thomas, D. Ng. 1992. Distinguishing between 'preference' and 'motivation' in food choice: An example from insect oviposition. Animal Behavior, 44: 463-471.
Struttman, J. "Butterflies of New Jersey: Regal Fritillary" (On-line ). Butterflies of North America. Accessed 02/18/03 at http://www.npwrc.usgs.gov/resource/distr/lepid/bflyusa/usa/80.htm.
Swengel, A. 1996. Effects of fire and hay management on abundance of prairie butterflies. Biological Conservation, 76: 73-85.
Swengel, A., S. Swengel. "The Tall-grass Prairie Butterfly Community" (On-line ). Accessed 02/18/03 at http://biology.usgs.gov/s+t/noframe/f071.htm.
Wagner, D., M. Wallace, J. Boettner, J. Elkinton. 1997. Status update and life history studies on the regal fritillary (Lepidoptera: Nymphalidae) in Ecology and Conservation of Grasslands and Heathlands of Northeastern North America. Lincoln: Massachusetts Audubon.
Weed, C. 1926. Butterflies. Doubleday, Page, and Company.
Williams, B. 2002. Conservation, genetics, extinction, and taxonomic status: a case history of the regal fritillary. Conservation Biology, 16/1: 148-157.
Williams, B. "Evolutionary/Conservation Genetics of the Regal Fritillary <<Speyeria idalia>>" (On-line ). Accessed 02/18/03 at http://www.life.uiuc.edu/paige/williams/idalia.html.
Williams, B. "Regal Fritillaries in a Tailspin" (On-line ). North American Butterfly Association. Accessed 01/28/03 at http://www.naba.org/chapters/nabasep/99_W_article.HTM.