Volume 10 Number 6 18 September 2022

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The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

ISSN 2643-4776 (print) / ISSN 2643-4806 (online)

Subspecific designation for the Central Appalachian Mountains

population of Argynnis (Speyeria) atlantis (W. H. Edwards).

Harry Pavulaan

606 Hunton Place NE

Leesburg, Virginia, USA, 20176

ABSTRACT. The population of Argynnis atlantis of the central Appalachian Mountains, primarily on the Allegheny

Plateau of West Virginia, has long been recognized by entomologists in the eastern United States as distinct from

nominotypical populations of A. atlantis in the northeastern United States and eastern Canada. Larger size is often cited, but

actual comparison of wing markings has not been done to date. This study applies RGB color analysis on a series of adults

from both regions and looks at potential differences in wing markings which may distinguish them.

Additional key words: Isolate, Allegheny Plateau, Canadian Zone, RGB color code.

ZooBank registration: http://zoobank.org/CB8FCC4A-57DA-4555-AA9E-CAF59761947

INTRODUCTION

A difficulty with the identification of specimens in the genus Argynnis (Speyeria), commonly

known as “Fritillaries”, is that among many of the populations, especially those in the western portion of

the United States, correct determination of specimens to which species or subspecies they belong is often

problematic due to similarity in wing patterns, size and ground color. There is considerable character

overlap and phenotypic intergradation between species and subspecies, and populations frequently take on

the appearance of another species. Thus, several species and subspecies have been reclassified from their

originally described rank. Some examples of this close resemblance are subspecies of A. atlantis, A.

aphrodite, A. zerene, A. hesperis, and A. hydaspe (Howe, 1975; Scott, 1986), some of which resemble

related species. Additional “problem” pairs include A. e. egleis vs. A. m. mormonia (Davenport, pers.

comm.). The present study will not delve into these, as more involved research on the North American

Argynnis (Speyeria) will be needed, especially genomic analysis at the species and subspecies level.

The population of A. atlantis on the Allegheny Plateau of northeastern West Virginia, also known

as the Allegheny Mountains, has been a popular target species of eastern U.S.A. butterfly collectors. Both

the late Bill Grooms and Ron Gatrelle, as well as a host of other entomologists in this region, have shared

with me observations that specimens of the Allegheny Plateau population “differ” from nominotypical

populations found in New York, northern New England and eastern Canada, but exact wing characters

were often difficult to pinpoint. This study provides a more detailed comparison.

The Allegheny Plateau is known for its “northern” character, primarily consisting of Canadian and

Transition Zone habitat types at higher elevations, most pronounced toward the eastern edge along the

Allegheny Front. Other endemics with northern affinities are Colias interior carolae, Tharsalea

epixanthe nr-michiganensis and Chlosyne harrisii liggetti. Pterourus appalachiensis is most common in

this region, having evolved from ancient hybrid introgression between P. canadensis and P. glaucus.

METHODOLOGY

Specimens from the Allegheny Plateau of West Virginia were compared to specimens from

northern New England, New Brunswick, New York and Pennsylvania considered to be nominotypical.

The TL of A. atlantis is currently recognized as “Hunter, Green County, New York” (dos Passos, 1935),

though the original description (W. H. Edwards, 1862) covered a broad area: Catskill Mountains (New

York), White Mountains (New Hampshire), Williamstown (Masshusetts), Lake Winnipeg (Manitoba),

“Hudson’s Bay” (Canada) and “the north side of Lake Superior’. The original image of Argynnis atlantis

was published in a later work (Edwards, 1869-1897) and is illustrated below (Fig. 1).

Fig. 1. Original illustration of Argynnis atlantis (dorsum on left, venter on right) on plate ‘Argynnis 5’ from ‘The Butterflies of

North America’ (W.H. Edwards, 1868-1897).

15 female and 29 male specimens (all fresh-condition) from West Virginia and 17 female and 18

male specimens of nominotypical atlantis (similarly fresh-condition) from New York, New England and

immediately adjacent areas of New Brunswick were analyzed using the Color Grab™ cellphone

application (www.loomatix.com), version 3.9.2, to establish exacting RGB color codes under “daylight”

fluorescent lighting, in combination with the Colblindor™ application (www.color-blindness.com/color-

name-hue/) to produce refined color swatches rather than giving generalized color descriptions as is

traditional with taxon descriptions. Determining exact RGB color code of the examined series is

important, as specimens fade over long periods of time and their RGB color codes will likely drift. Four

areas of the wings were measured (Fig. 2): (1) outer dorsal ground color of the forewings in cell Ms; (2)

inner dorsal ground color of the forewings near the base of cell CuAg; (3) interior ventral ground color of

the forewings in cell CuA1; (4) inner ventral ground color of the hindwings at the base of cell Sc+R1

between the adjacent silver spots within that cell. The RGB values for each series, separately for males

and females, was averaged and the results are shown in Figs. 3 & 4. While not critical to the description,

the color nomenclature from the Colblindor™ application is available in Appendix A for those interested

in exacting color names. [It is important for future researchers to note that the RGB color codes of any

examined series may not perfectly match the results of the current study due to differences in lighting

used and freshness of specimens. Online viewing of this study and hardcopy printing may also distort the

colors. A textbook example lies with the original illustration of atlantis when viewed and copied off

www.biodiversitylibrary.org; a very dark version of the male is depicted (Fig. 1). The important thing to

remember is to be consistent with ALL specimens analyzed in any comparative study employing

lighting. |

The specimens used for RGB color analysis in this study were also measured for size. The leading

edge of the forewings were measured from the base of the wing at the thorax to the farthest point outward,

to the apex. These measurements are shown in Figs. 3 & 4. Howard Grisham, Tom Kral and Ricky

Patterson provided additional extensive imagery of West Virginia specimens and Tom Kral provided

extensive imagery of nominotypical atlantis from McKeen County, Pennsylvania collected by William

Houtz approximately 100 miles west of the Hunter, New York TL of atlantis.

Fig. 2: Wing areas measured for RGB color analysis and forewing length in

this study. Numbers correspond to charts in Figs. 3 & 4. “A” represents exact

area of forewing length measurement. Dorsal view (left), ventral view (right).

216, 145, 27 92, 57, 30 190, 124, 57 71, 44, 35 27-31

Argynnis atlantis West Virginia - ~ RGB color averages and FW length

216,131,314 110, 69, 44 190, 129, 53 81, 47, 35 27-31

Argynnis atlantis atlantis - 3! RGB color averages and FW length rs

226, 145, 37 123, 77, 44 207, 152,82 | 109, 73, 57

Fig. 4: RGB color analysis of four male characters and forewing length.

DISCUSSION

Though both dorsal and ventral wing markings are variable in both central Appalachian and

nominotypical populations, this study finds no consistent differences in wing markings except in the

females which have reduced black spotting dorsally and a larger, “brighter” appearance in the central

Appalachians than females in the nominotypical region. Three of the measured female characters (2, 3

and 4) showed no appreciable difference in colors, but the outer ground color showed measurable (and

visually noticeable) difference. West Virginia females average larger than nominotypical females (Figs. 3

& 5) and are occasionally of very large size, some difficult to differentiate from sympatric A. aphrodite

females when viewed (or photographed) from just the dorsal side. Fig. 2 displays the largest specimen

examined in this study, with a forewing length of 35 mm. Thus, the holotype of the new subspecies is

based on the female phenotype.

Males of both nominotypical and Appalachian populations showed no appreciable difference in

wing markings, which is variable in both populations. But from the four analyzed characters, the males

did show a slight difference in average dorsal ground color, and the forewing length measurements

showed West Virginia males to average slightly larger (Figs. 4 & 6).

DESCRIPTION, TYPE SPECIMENS AND DISTRIBUTION

Argynnis (Speyeria) atlantis brittanae - new subspecies

ZooBank registration: urn:lsid:zoobank.org:act:7675E05 1-CESA-4C1D-BA47-C5AE3067DC11

Phenotypic description: Males are like nominotypical atlantis (Fig. 6) but average a scant 1.09

mm larger in forewing length. Measured length of male forewings in the measured series (n=29) is 27-31

mm, averaging 29.92 mm. Color swatches generated using RGB color codes (Fig. 4) show a slight

average difference in the shade of dorsal orange-brown ground color in the study series. West Virginia

populations (subspecies brittanae) males tend to have a subtle “warmer” dorsal reddish orange-brown

ground color and the dorsal wing margins are less black as in nominotypical atlantis. Nominotypical

populations of atlantis have a tanner orange-brown ground color and very dark margins. Variability in

any specimen series will show considerable overlap. Large series of both subspecies are necessary for

ascertaining an average RGB using a color analysis tool.

Females of the West Virginia populations are of a stronger shade of reddish orange-brown than

nominotypical atlantis females (Fig. 5) which appear to be paler and have a slight yellow component.

While males of both atlantis and brittanae have similar, variable dorsal wing pattern, the females differ.

Nominotypical atlantis females have a more developed pattern of black markings and darker wing

margins, while brittanae females have a tendency for reduced markings and less dark margins by

comparison, most noticeably on the hindwings. This gives the females a “brighter” appearance.

The measured females (n=15) average 3.17 mm larger than nominotypical atlantis females in

forewing length, ranging from 30-35 mm, averaging 32.42 mm. [Some larger females in West Virginia

approach the sympatric A. aphrodite aphrodite phenotype (Fig. 2), differentiated mainly by more

extensive dark wing margins, and the more expansive tan postmedian band on the ventral side of the

hindwings, between the outer two rows of silver spots, whereas in aphrodite the inner brown ground color

of the ventral side of the hindwings encroaches and partially fills in this tan band.] Interestingly, the

series of nominotypical atlantis females and males were of about the same size; the female forewing

length averaging only .42 mm larger than the males. By contrast, subspecies brittanae females were

markedly larger than the males, averaging 2.5 mm larger.

Fig. 5. Comparison of females. Ssp. brittanae (above), W.V., Fig. 6. Comparison of males. Ssp. brittanae (above), W.V.,

Randolph County, Spruce Knob Lake, 25 June, 2009. Ssp. Randolph County, Spruce Knob Lake, 15 June, 2014. Ssp.

atlantis (below), N.Y., Ulster Co., Ellenville, 9 July, 1992. atlantis (below), N.Y., Ulster Co., Ellenville, 9 July, 1992.

Distribution: The range is basically confined to the Canadian Zone of the central Appalachian

Mountains (Allegheny Plateau region). Allen (1997) notes that West Virginia populations are disjunct

from northern populations. It is primarily an isolate but some contact with ssp. atlantis may occur in

southwestern Pennsylvania. County records are as follows: Maryland: Garrett. Virginia: Highland, with

an unconfirmed historical sight records in Bath Co. (Clark & Clark, 1951) and Rappahannock Co. (Opler,

1995). West Virginia: Barbour*, Grant*, Greenbrier, Hampshire*, Hardy*, Marion*, Mineral*,

Monongalia*, Pendleton, Pocahontas, Preston, Randolph, Tucker, Webster. Populations in southwestern

Pennsylvania on the Allegheny Plateau require confirmation of assignment to brittanae. The butterfly is

best known from Spruce Knob, Canaan Valley and Dolly Sods Wilderness in West Virginia. Counties

indicated with (*) indicate historic non-breeding (stray) records (Allen, 1997; Olcott, pers. comm.).

Interestingly, the West Virginia Butterfly Survey (Olcott, pers. comm.) shows the active, current range

very restricted, primarily limited to a relatively small area on the Allegheny Plateau. Past records outside

the active range indicates historic strays to neighboring counties or perhaps a slightly greater historic

range now contracted.

Habitat: Frequently found in high elevation fields, meadows, fens, bogs, pastures and other open

areas in association with Canadian Zone and upper Transition Zone forests. They are easily found

nectaring on a broad variety of flowers along roadsides in these habitats and through forest.

Hosts: A. atlantis is known to utilize Violets (Viola sp.) range wide but the exact species on the

Allegheny Plateau has not been determined.

Flight Period: Single brood. Records span June 8 — October 2, with peak flight in late June and

early July. Immature stages of this subspecies have not been studied.

Holotype, allotype, paratypes: Holotype (female) (Figs. 7 & 9): West Virginia, Randolph

County, Spruce Knob Lake, 25 June, 2009. Collected by Harry Pavulaan. Allotype (male) (Figs. 8 &

10): West Virginia, Randolph County, Spruce Knob Lake, 15 June, 2014. Collected by Harry Pavulaan.

Both holotype and allotype to be deposited in the McGuire Center for Lepidoptera and Biodiversity,

Gainesville Florida. Paratypes: 28 male and 14 female specimens from Pendleton, Randolph, and Tucker

Counties, W.V. retained in the collection of Harry Pavulaan, to be disseminated at a future date. 15 male

and 7 female specimens from Pendleton and Tucker Counties, W.V. in the collection of Howard Grisham.

31 male and 11 female specimens from Pendleton and Tucker Counties, W.V., collected by Howard

Grisham, John Hyatt and Tom Allen, in the collection of Tom Kral. 14 male and 8 female specimens

from Pendleton County, W.V. in the collection of Ricky Patterson.

Etymology: I name this subspecies after my daughter Brittany. The latinized equivalent is

“brittanae”. I propose the common name “Allegheny Fritillary” due to its small range in that region.

Fig. 7. Holotype, W.V., Randolph County, Spruce Knob Fig. 8. Allotype, W.V., Randolph County, Spruce Knob

Lake, 25 June, 2009, female, dorsum. Coll. Harry Pavulaan. Lake, 15 June, 2014, male, dorsum. Coll. Harry Pavulaan.

Fig. 9. Holotype, W.V., Randolph County, Spruce Knob Fig. 10. Allotype, W.V., Randolph County, Spruce Knob

Lake, 25 June, 2009, female, venter. Coll. Harry Pavulaan. Lake, 15 June, 2014, male, venter. Coll. Harry Pavulaan.

ACKNOWLEDGMENTS

Thanks are expressed to Tom Kral for review of the manuscript. Thanks also to Ricky Patterson,

Howard Grisham and Tom Kral for extensive specimen imagery and data, also to Sue Olcott (West

Virginia Butterfly Survey) for survey data and feedback on common name. Tom Kral provided extensive

imagery and data for specimens collected by William Houtz in McKeen County, Pennsylvania. Cris

Guppy and Ken Davenport provided additional helpful comments.

LITERATURE CITED

Allen, T. J. 1997. The Butterflies of West Virginia and Their Caterpillars. University of Pittsburgh

Press, Pittsburgh, PA.: xii + 388 pp.

Clark, A. H. & L. F. Clark. 1951. The Butterflies of Virginia. Smithsonian Miscellaneous Collections

116 (7): v + 239 pp.

dos Passos, C. 1935. Some butterflies of southern Newfoundland with descriptions of new subspecies

(Lepidoptera: Rhopalocera). Canadian Entomologist 67(4): 82-88.

Edwards, W. H. 1862. Descriptions of certain species of DIURNAL LEPIDOPTERA found within the

limits of the United States and British America — No. 2. Proceedings of the Academy of Natural

Sciences of Philadelphia 14(1/2): 54-58.

Edwards, W. H. 1868-1897. The Butterflies of North America. The American Entomological Society,

Philadelphia, Pennsylvania: 11+ 52 pp. + 50 pl.

Howe, W.H. 1975. The Butterflies of North America. Doubleday & Company, Inc., Garden City, N.Y.:

Xili + 633 pp. + 97 pl.

Opler, P. A. 1995. Distribution of the Butterflies (Papilionoidea and Hesperioidea) of the eastern United

States. Contributions of the C.P. Gillette Museum of Arthropod Diversity, Colorado State

University. Lepidoptera of North America 2: 167 pp.

Scott, J. A. 1986. The Butterflies of North America. A Natural History and Field Guide. Stanford

University Press, Stanford, CA.: xiii + 583 pp. + 64 pl.

APPENDIX

226, 145, 37

216, 145, 27

ri!

216, 131, 31

;

207, 152, 82

207, 125, 17

190, 129, 33

190, 124, 37

186, 111, 35

123, 77, 44

110, 69, 44

109, 73, 57

102, 60, 24

92,37, 30

81,47, 35

71, 44, 35

o2, 29, 26

Appendix A. Color Nomenclature from Colblindor™ for corresponding RGB color codes used in this study.

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