Volume 9 Number 9 1 November 2021

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

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

Subspecific designation of the U.S.A. Interior Highlands population of

Argynnis (Speyeria) diana (Cramer, 1777)

(Nymphalidae: Heliconiinae: Argynnini: Argynnina)

Harry Pavulaan

606 Hunton Place NE

Leesburg, VA 20176 USA

ABSTRACT. Subspecific designation is designated for the North American Interior Highlands population of Argynnis diana,

based on four factors: mtDNA haplotype differences from nominotypical A. diana of the Appalachian Mountains; wing shape

difference in the males between both regions; wing size of the adults; and tendency for females of the Interior Highlands to

show tan coloration in the submarginal row of rectangular spots of the subapical region of the dorsal forewings.

Additional key words: Interior Highlands, Ozark Region, Ouachita Mountains, Appalachian Mountains, wing shape,

dispersal, range collapse, range contraction.

ZooBank registration: urn:lsid:zoobank.org:pub:EB9A7D46-79D7-4280-9548-3654F924530A

INTRODUCTION

The Interior Highlands is a combined physiographic region comprised of several geologic

provinces across the states of Missouri, Arkansas and Oklahoma, consisting of the Salem Plateau, St.

Francois Mountains, Springfield Plateau, and Boston Mountains, collectively known as the Ozark Plateau

or Ozark Region. The Ouachita Mountains, technically south of the Ozarks, form their own geologic

province, and together form the Interior Highlands, with floral and faunal links to the Appalachian

Mountain region of the eastern United States. Among the butterflies, Celastrina nigra, Celastrina

neglectamajor, Calephelis borealis and Argynnis diana each share Interior Highlands and Appalachian

Mountain affinities, with their distributions concentrated in each of the two regions, and with scant

records in the intervening region.

A. diana is the largest member of the North American subgenus Speyeria. It is also the most

sexually dimorphic species of the subgenus. The historical range of A. diana extended broadly from the

Atlantic Coastal Plain west to Oklahoma, and from central areas of Georgia, Alabama and Mississippi

north to the southern edge of the Great Lakes (Fig. 9). There has been some documented contraction of

its range, particularly in lowland habitats. It has become extirpated where it was originally discovered in

coastal Virginia; and in the Mississippi Valley and lower Ohio River Valley lowland regions. Factors

causing this range contraction are believed to be: habitat loss (woodland converted to farmland,

widespread logging, changes in land use, residential encroachment into the habitat, wetland draining);

habitat fragmentation; floral competition against host Viola species; pesticides (specifically aerial

application of Bacillus thuringiensis for Lymantria dispar (Gypsy Moth) control); uncontrolled deer

browsing; and fire suppression resulting in profound ecological changes (Rudolph et al., 2006). All have

negative impacts on diana populations in different areas, thus the species is of conservation concern.

Global warming is presented as a predicted factor for continued range contraction in the future (Wells,

2014; Wells & Tonkyn, 2013; Wells et al., 2015; Wells & Tonkyn, 2018), as evidenced by the butterfly

occupying higher elevation habitats in greater numbers over the studied time period, especially in the

southern Appalachians. Interestingly, Wells (2014) and Wells & Tonkyn (2018) noted that populations of

S. diana in the southern Appalachian Mountains and in Oklahoma and Arkansas have expanded in recent

decades. Moran & Baldridge (2002) concluded: “... A. diana does not appear to be in immediate risk of

extirpation in Arkansas, although monitoring of existing populations is warranted.” Habitat restoration

efforts, employing prescribed burning of forest habitat in the Ouachita Mountains and in western North

Carolina have been documented to help restore the habitat to pre-European conditions by reducing the

forest canopy, eliminating mid-story growth, and restoration of dense herbaceous ground cover (Rudolph

et al., 2006; Campbell, et. al., 2007), thus helping to increase diana abundance. Presently, A. diana is not

protected at any local, state or federal level, other than protection within parklands.

However, it begs one to consider that many of the records plotted within the “collapsed” central

portion of the species’ range during human record keeping were merely historical strays outside of the

species’ Appalachian and Interior Highlands strongholds. For example, there have been recent strays of

unknown origin, reliably reported from places as far north as Chatham, ON. (BAMONA, 2011) and as far

east as Jamestown, R.I. (D. Albaugh, pers. corr., 1999). A continuous range across the eastern United

States certainly predates the most recent glacial maximum. Wells et al. (2015) estimated that the split

between Appalachian and Interior Highlands populations occurred at least 20,000 years ago, thus enough

time to develop differences in DNA, and long predates human activity. Thus, it is premature to conclude

that human activity or global warming are responsible for the perceived collapse of the species’ range in

the Mississippi and Ohio River Valley regions. Hovanitz (1963) sums this up nicely: “This species...is a

relatively rare species due to the limited areas in which it lives. There is no evidence that at any time in

the past the species was any less local than it is now” but notes that deforestation [in Virginia] contributed

to its demise there. He further states: “The species exists in parts of North America away from the most

heavily inhabited regions and therefore collectors have not had as much experience with it...”” This would

certainly skew the historical record of its distribution and abundance!

Considering that A. diana has long occupied mainly the Appalachian and Interior Highlands

regions, it has been questioned whether Interior Highlands A. diana is represented at subspecies-level,

differentiated from its Appalachian counterpart. A recent study by Wells et al. (2018) indicated that

males of the Interior Highlands and Appalachian populations of A. diana differ in forewing shape and the

authors posed a hypothesis for this. Females in that study showed a different alignment of wing shape,

reflective of high vs. low elevation populations rather than regional. Independent investigation by myself

(2016, unpubl.) found no consistent difference in dorsal or ventral wing markings of the males from either

region. Individual variation among males overlapped completely, range wide. Likewise, females from

both regions displayed no consistent differences with the exception of the color of the postmedian marks

in the apical portion of the dorsal forewing, and wing size, that might be considered weakly subspecific.

COMPARISON OF INTERIOR HIGHLANDS AND APPALACHIAN A. DIANA

Wells et al. (2018) conducted a thorough geometric morphometric analysis of the two A. diana

regional populations and found significant regional differences in male forewing and hindwing shape, and

concluded that wing shape of adults can be used as a character to measure population-level differences.

Males from the Appalachian Mountains were determined to have narrower and more angular wings,

believed to support high dispersal behavior, vs. Interior Highlands males which have rounder wings,

supporting low dispersal behavior. The authors also found significant differences in female forewing

shape, not regionally, but between high and low elevation populations, where high elevation females

possessed narrower forewings than those from low elevations. Unfortunately, they did not specify in

which region(s) this female forewing character occurred or was most prominent [it is my interpretation

that this means that Appalachian Mountain females qualify as “high elevation” vs. “low elevation”

females found in the remainder of diana’s range, including the Interior Highlands region].

My own detailed analysis of adult morphology from both

regions shows virtually no consistent differences in wing marks in

the males. The range of variation in both Interior Highlands and

Appalachian males, including coloration and extent of all dorsal

and ventral markings, appears to fully overlap range wide.

Preliminary measurements of my collected series of males from

both Arkansas (n=18) and Virginia (n=16) similarly show little

difference in wing shape, with both series showing a high degree

of variability [certainly not at the scale of investigation of the

Wells et al. (2018) study, but worthy of note, and supports the

argument that large series are often needed to make reliable

conclusions]. However, two males were selected, one from each

region, each with similar forewing length, and measurements were

made, which corroborate the findings of Wells et al. (2018). The

red-outlined silhouette (ex Needmore, AR) is overlain with the

blue-outlined silhouette (ex Longdale Furnace, VA) (Fig. 1). Both

specimens have identical forewing length (measurement A) at 43

mm. Measurement B (Virginia) is 26 mm, and measurement C

(Arkansas) is 29 mm, confirming that Appalachian males have

Fig. 1. A. diana male wings comparison.

narrower forewings than their Interior Highlands counterparts. The hindwing of the Arkansas specimen is

measurably larger than the Virginia specimen, measuring 35 mm (measurement E) along vein Cui,

whereas the Virginia specimen measures 33 mm (measurement D). A visual representation of the male

wing shape difference is shown below (Fig. 2) [same specimens measured for Fig. 1]. Difference in

forewing shape is most evident. Overall, Arkansas males (n=24) averaged 45.8 mm forewing length,

while Appalachian males (n=58) averaged 43.7 mm (2.1 mm smaller). Arkansas females (n=12) averaged

55.2 mm while Appalachian females (n=21) averaged 51.5 mm (3.7 mm smaller). A study by Showalter

& Drees (1980), using an unspecified sample size of Appalachian diana indicated forewing lengths

averaging 44.1 mm for males and 52.3 mm for females, each slightly larger in size than the present

sample.

Fig. 2. A. diana male, nr. Needmore, Scott Co., AR., 6/12/2016 (left image).

A. diana male, nr. Longdale Furnace, Alleghany Co., VA 6/28/2017 (right

image). Narrower, more elongated forewing in Appalachian male is evident.

While the Wells et al. (2018) study found a consistent wing shape difference between males of

both regions, they did not study differences in wing markings. Analysis of male wing marks from both

regions in the author’s personal collection, various printed literature sources (Table 1), and a selection of

196 images [clearly-focused perpendicular views of fresh individuals, not sun-backlit and no shadows on

wings] posted to the internet via butterfliesandmoths.org, butterfliesofamerica.com, and iNaturalist.org

(Table 2) corroborates no consistent differences in dorsal or ventral wing marks in males beyond a wide

range of individual variation matched in both regions.

Additionally, the Wells et al. (2018) study determined that females expressed a different degree of

wing shape development based on elevation, where low elevation females have wider forewings than

those from high elevations, reflective of dispersal and host-locating habits. The authors also did not

identify any differences in wing marks among females from both regions, for their study. However, my

own detailed analysis of female wing mark characters, utilizing the same sources listed above (Table 2),

did find that there is one character that might be considered useful in differentiating some females from

both regions. Females from the Appalachian populations generally displayed consistently whitish-blue

coloration in the submarginal band of rectangular marks in the subapical area of the forewing (dorsal

side), while females from the Interior Highlands populations showed a tendency toward tan to whitish-tan

coloration of these rectangular marks. This tendency for tan coloration in the subapical area of the

forewing is not seen in all Interior Highlands females examined, but is rarely encountered in Appalachian

females. [One problem encountered is the quality of published and web-sourced images. Lighting and

focus often distorts the true color of wing marks. ]

An earlier study (Dunford, 2007), which constructed a phylogeny of Speyeria based on 653

characters of the mitochondrial gene COI, placed Arkansas diana separately from West Virginia diana.

The two sampled Arkansas diana were identified with 657 COI base pairs, whereas the two West Virginia

diana were separately identified with 643 and 657 COI base pairs. Several phylograms of the Speyeria

(Dunford, 2007: Figures 3-8 through 3-11) were constructed for strict consensus trees, all consistently

showing the two Arkansas samples grouping together, separated from the two West Virginia samples.

More recently, the Wells (2014) and Wells et al. (2015) studies found significant differences in

mtDNA haplotypes between Appalachian and Interior Highlands populations, with eastern populations

showing high levels of genetic diversity and Interior Highlands populations with less genetic diversity.

Different haplotypes were found to dominate the Appalachian and Interior Highlands populations (Wells,

2014; Wells et al., 2015), with ‘haplotype 1’ dominating the Interior Highlands population and ‘haplotype

2’ dominating the Appalachian population. Wells et al. (2018) concluded that morphological and

mitochondrial DNA differences between the two regional populations “may warrant subspecies

designation”’.

ARGYNNIS (SPEYERIA) DIANA ARKANSANA —- NEW SUBSPECIES

ZooBank registration: urn:]sid:zoobank.org:act:3 CDESE6E-F9F9-4CEB-BEEC-22CAA67D94B2

Description: The primary difference between Interior Highlands and Appalachian populations of A.

diana is in the rounder shape of the male forewings of the Interior Highlands (Wells et al., 2018), whereas

Appalachian male forewings were found to be “narrower and more angular’ than Interior Highlands

males (Figs. 1 & 2). The authors also found that male hindwings from the Appalachian population were

narrower than those from the Interior Highlands. Also, there is a tendency for females of the Interior

Highlands population (Fig. 7) to display tan coloration in the submarginal row of marks in the subapical

area of the forewing (dorsal side) rather than the whitish-blue coloration found in Appalachian females

(Fig. 8). In my analysis, male forewings of the Interior Highlands region averaged 2.1 mm longer than

Appalachian males, with a sampled range of 42-48 mm in the Interior Highlands and a range of 39-48 mm

in the Appalachian region. The forewings of Interior Highlands females averaged 3.7 mm longer than

Appalachian females, with a sampled range of 54-57 mm in the Interior Highlands and a range of 48-56

mm in the Appalachian region.

Habitat: Carlton & Nobles (1996) compiled information from various sources and listed the habitat

choices for the Interior Highlands variously as: hardwood/pine forest (especially edge habitats); second

growth pine hardwood forest; even-aged pine stands; mature upland hardwood forest; “a mosaic of

severely disturbed pine and second growth mixed forest in various stages of succession with a dense

understory of woody vines, shrubs and small trees; and tallgrass prairie/patchy forest with dense

undergrowth. A. diana was also recorded in prairie habitats in southwest Arkansas, small prairie

remnants in mountainous northwest Arkansas, and in wetland habitats in central Arkansas (Moran &

Baldridge, 2002). The authors’ survey indicated that diana was more widespread throughout the

mountainous region than previously thought. They also found that diana thrived in moderately disturbed

habitat such as second growth forest and pastureland and the butterflies thrive where the habitat is

frequently burned. In the Interior Highlands region, the butterfly was also reported in pine-dominated

forests (Rudolph, et. al., 2006), especially consisting of Pinus echinata, with sparse midstories and an

understory of Schizachrium spp. grasses and abundant nectar sources (Fig. 3). They reported the species

is also found in Quercus/Carya-dominated forest such as found on Mt. Magazine in Arkansas (Fig. 4).

Spencer (2014) summarizes the habitat in Arkansas as “open moist (mesic) forests, prairies and

wetlands”.

Fig. 3. Pine-dominated lowland habitat west Fig. 4. Upland habitat atop Mt. Magazine, Logan County, AR.

of Needmore, AR.

Larval hosts: Viola pedata, Viola riviniana (Spencer, 2011). Other Viola species are suspect, but no

others have been recorded in the Interior Highlands. Several Viola species recorded in the Appalachians.

Nectar sources: A. diana is highly dependent on high-quality nectar sources that can enhance the

species’ reproductive abilities (Wells & Smith, 2013). Moran & Baldridge (2002) recorded the following:

Cephalanthus occidentalis, Echinacea purpurea, Echinacea pallida, Silphium laciniatum, Satureia

arkansana, Pycnanthemum albescens, and Rubus sp. Primary nectar sources recorded by Rudolph et al.

(2006) added the following: Asclepias tuberosa, Cirsium carolinianum, Cirsium discolor, Liatris elegans,

Monarda fistulosa, Porteranthus stipulatus, and Pycnanthemum tenuifolium. Secondary nectar sources

recorded during their study were: Rhexia sp., Scutellaria ovata, Erigeron strigosus, Bidens aristosa,

Eupatorium fistulosum, Solidago rugosa, Helianthus divaricatus, Vernonia gigantea and Vernonia

baldwinii. Spencer (2014) listed Coreopsis sp., Vernonia sp., and garden cultivars such as Lantana

camara, Pentas lanceolata and Buddleia sp. Various internet-sourced images confirmed the following:

Asclepias syriaca, Asclepias exaltata, Daucus carota, Eupatorium perfoliatum and Cornus sp. Recently-

emerged adult males were also observed to imbibe from non-nectar sources such as animal feces,

regurgitated plant material (animal vomit), carrion, damp soil, dusty road surfaces, even human sweat

(Rudolph et al., 2006).

Etymology: The subspecies name arkansana represents the primarily Arkansas portion of the subspecies

range. In 2007, it was designated as the Arkansas State Butterfly.

Holotype, allotype and paratypes: A female originating in the Ouachita Mountains region is selected as

the holotype of Argynnis (Speyeria) diana arkansana (Fig. 5). The type locality is: County Road 30, 0.64

miles west of U.S. Route 71, Needmore, Scott County, Arkansas. Date is June 12, 2016. The holotype,

allotype male (Fig. 6) and several paratype males collected at the TL are deposited in the McGuire Center

for Lepidoptera and Biodiversity, Gainesville, FL. and a pair is retained by the author.

Fig. 5. Holotype female, 6/12/2016, Needmore, AR. Dorsal view (left), ventral view (right).

Fig. 6. Allotype (paratype) male, 6/12/2016, Needmore, AR. Dorsal view (left), ventral view (right).

Fig. 7. A. diana arkansana female displaying tan markings Fig. 8. A. diana diana female displaying blue markings in

in FW subapical area. Mount Magazine State Park, Logan FW subapical area. Cleburne Co., AL, August 23, 2021.

Co., AR, June 26, 2004. Photo courtesy © Bill Bouton. Photo courtesy © Sara Bright.

Fig. 9. Extant range of Argynnis diana, showing the current range of ssp. diana (blue) and ssp. arkansana

(tan). Historic range, pre-1960 (extirpated and strays) shown in red.

CONCLUSION

The following subspecific arrangement is hereby designated. Alignment under the genus Argynnis and

subgenus Speyeria follows Simonsen (2006), Simonsen, et. al. (2006), and Zhang, et. al. (2020):

Argynnis (Speyeria) diana diana (Cramer, 1777), TL: Jamestown, James City County, VA.

Argynnis (Speyeria) diana arkansana Pavulaan 2021, TL: County Road 30, 0.64 miles west of

U.S. Route 71, Needmore, Scott County, AR.

ACKNOWLEDGMENTS

Thanks to Steve Spomer for review of the manuscript, discussions, wing measurements and leads

to additional literature references; David M. Wright for providing additional leads to literature and for

proofreading the manuscript; Bill Dempwolf, Ricky Patterson, Jeff Slotten, and Mark Walker for

additional wing measurements. Thanks also to Sara Bright and Bill Bouton for permission to reproduce

the live photo images. A posthumous acknowledgement goes to Ron Gatrelle for previous discussions on

taxonomic matters concerning A. diana.

LITERATURE CITED

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

Pittsburgh, PA.: xii + 388 pp.

Bouseman, J. K. & J. G. Sternburg. 2001. Field Guide to Butterflies of Illinois. Illinois Natural History

Survey, Champaign, IL., Manual 9: xii + 264 pp.

Campbell, J. W., J. L. Hanula & T. A. Waldrop. 2007. Observations of Speyeria diana (Diana Fritillary)

utilizing forested areas in North Carolina that have been mechanically thinned and burned.

Southeastern Naturalist 6(1): 179-182. https://www.srs.fs.usda.gov/pubs/ja/ja_campbell002.pdf

Carlton, C. E. & L. S. Nobles. Distribution of Speyeria diana (Lepidoptera: Nymphalidae) in the

Highlands of Arkansas, Missouri and Oklahoma, with comments on conservation. Entomological

News 107(4): 213-219. https://www.biodiversitylibrary.org/page/270049 1#page/233/mode/lup

Cech, R. & G. Tudor. 2005. Butterflies of the East Coast, an Observer’s Guide. Princeton University

Press, Princeton, N.J.: xii + 345 pp.

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

116(7): v + 239 pp. https://archive.org/details/cbarchive_ 107998 _ thebutterfliesofvirginial 862

Dole, J. M., W. B. Gerard & J. M. Nelson. 2004. Butterflies of Oklahoma, Kansas & North Texas.

University of Oklahoma Press, Norman, OK.: xiv + 282 pp.

Dunford, J.C. 2007. The genus Speyeria and the Speyeria atlantis/Speyeria hesperis complex: species

and subspecies accounts, systematics, and biogeography (Lepidoptera: Nymphalidae). PHD

dissertation. University of Florida, Gainesville, FL.: 245 pp.

Glassberg, J. 1999. Butterflies Through Binoculars, the East. A Field Guide to the Butterflies of Eastern

North America. Oxford University Press, New York, N.Y.: x + 242 pp. + 71 pl.

Harris, L. H. Jr. 1972. Butterflies of Georgia. University of Oklahoma Press, Norman, OK.: xxii + 326

pp.

Heitzman, J. R. & J. E. Heitzman. 1987. Butterflies and Moths of Missouri. Missouri Department of

Conservation, Jefferson City, MO.: viii + 385 pp.

Howe, W. H. (ed.). 1975. The Butterflies of North America. Doubleday & Company, Garden City,

N.Y.: xiii + 633 pp. + 97 pp.

Hovanitz, W. 1963. Geographical distribution and variation of the genus Argynnis III. Argynnis diana.

Journal of Research on the Lepidoptera 1(3): 201-208.

https://www.biodiversitylibrary.org/item/225295#page/22 1/mode/lup

Howell, W. M. & V. Charny. 2010. Butterflies of Alabama. Pearson Learning Solutions, Boston, MA.:

vil + 509 pp.

Iftner, D. C., J. A. Shuey & J. V. Calhoun. 1992. Butterflies and Skippers of Ohio. Bulletin of the Ohio

Biological Survey, New Series, 9(1): xii + 212 pp.

Klots, A. B. 1951. A Field Guide to the Butterflies of Eastern North America. The Peterson Field Guide

Series. Houghton Mifflin Company, Boston, MA.: xvi + 349 pp. + 40 pl.

Monroe, J. L. & D. M. Wright. 2017. Butterflies of Pennsylvania, a field guide. University of Pittsburgh

Press, Pittsburgh, PA.: xiii + 304 pp.

Moran, M. D. & C. D. Baldridge. 2002. Distribution of the Diana Fritillary, Speyeria diana

(Nymphalidae) in Arkansas, with notes on nectar plant and habitat preference. Journal of the

Lepidopterists’ Society 56(3): 162-165.

https:/Amages.peabody.yale.edu/lepsoc/jls/2000s/2002/2002-56(3)162-Moran.pdf

Opler, P. A. & G. O. Krizek. 1984. Butterflies East of the Great Plains. The Johns Hopkins University

Press, Baltimore, PA.: xvii + 294 pp. + 54 pl.

Opler, P. A. & V. Malikul. 1998. A Field Guide to Eastern Butterflies. The Peterson Field Guide Series.

Houghton Mifflin Company, Boston, MA.: xvii + 486 pp.

Rudolph, D. C., C. A. Ely, R. R. Schaefer, J. H. Williamson & R. E. Thill. 2006. The Diana Fritillary

(Speyeria diana) and Great Spangled Fritillary (S. cybele): dependence on fire in the Ouachita

Mountains of Arkansas. Journal of the Lepidopterists’ Society 60(4): 218-226.

https://www.srs.fs.usda.gov/pubs/ja/ja_rudolph0O22.pdf

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.

Showalter, A. H. & B. M. Drees. 1980. Bilateral gynandromorphic Speyeria diana (Nymphalidae).

Journal of the Lepidopterists’ Society 34(4): 340-344.

https:/Amages.peabody.yale.edu/lepsoc/jls/1980s/1980/1980-34(4)340-Showalter.pdf

Simonsen, T. J. 2006. Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on

the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini).

Biological Journal of the Linnean Society 89(4): 627-673.

https://academic.oup.com/biolinnean/article/89/4/627/269165 1 ?login=true

Simonsen, T. J., N. Wahlberg, A. V. Z. Brower & R. de Jong. 2006. Morphology, molecules and

fritillaries: approaching a stable phylogeny for Argynnini (Lepidoptera: Nymphalidae). Insect

Systematics and Evolution 37: 405-418. http:/Awww.nymphalidae.net/Simonsenetal2006.pdf

Smith, C. R. & E. A. Domingue. 2019. Butterflies and Moths of the Smokies. Great Smoky Mountains

Association, Gatlinburg, TN.: 301 pp.

Spencer, L. A. 2006. Arkansas Butterflies and Moths. Second Edition. Ozark Society Foundation.

Little Rock, Arkansas: xiv + 314 pp.

Spencer, L. A. 2011. The Diana Fritillary, Arkansas’s State Butterfly. Ozark Society Foundation.

Little Rock, Arkansas: 24 pp.

Wells, C.N. 2014. Range collapse, genetic differentiation, and climate change: an ecological history of

the Diana Fritillary, Speyeria diana and projections for its future. All Dissertations, Clemson

University TigerPrints 1382: 120 pp. https://onlinelibrary.wiley.com/doi/epdf/10.111 1/icad. 12059

Wells, C. N., P. B. Marko & D. W. Tonkyn. 2015. The phylogeographic history of the threatened Diana

fritillary, Speyeria diana (Lepidoptera: Nymphalidae): with implications for conservation.

Conservation Genetics 16(3): 703-716. https://doi.org/10.1007/s10592-014-0694-9

Wells, C. N., A. Munn & C. Woodworth. 2018. Geomorphic morphometric differences between

populations of Speyeria diana (Lepidoptera: Nymphalidae). Florida Entomologist 101(2): 195-

202. https://doi.org/10.1653/024.101.0207

Wells, C.N. & E. A. Smith. 2013. Observations of resource use by the threatened Diana Fritillary

butterfly (Speyeria diana) in the Southern Appalachian Mountains, USA. Journal of Insects

Vol. 2013, Article I.D. 130694: 4 pp. https://downloads.hindawi.com/archive/2013/130694.pdf

http://dx.do1.org/10.1155/2013/130694

Wells, C. N. & D. W. Tonkyn. 2013. Range collapse in the Diana fritillary, Speyeria diana

(Nymphalidae). Insect Conservation and Diversity 7(4): 365-380.

https://onlinelibrary.wiley.com/doi/epdf/10.11 1 1l/icad.12059

Wells, C. N. & D. W. Tonkyn. 2018. Changes in the geographic distribution of the Diana fritillary

(Speyeria diana: Nymphalidae) under forecasted predictions of climate change. Insects 9(94):

1-20. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6164860/pdf/insects-09-00094. pdf

Zhang, J., Q. Cong, J. Shen, P. A. Opler & N. V. Grishin. 2020. Genomic evidence suggests further

changes of butterfly names. The Taxonomic Report 8(7): 1-41.

https://archive.org/details/t-report-8-7-further-changes-in-butterfly-names

APPENDIX - SOURCES OF IMAGES

With the presence of the COVID-19 pandemic, institutional collections were closed to the general

public, even institutional staff, throughout the compilation of this report. Published imagery in printed

literature was sourced (Table 1), when locational information was provided. With the aid of web-based

imagery, much visual analysis was able to be accomplished by examining available imagery posted to

three major websites: butterfliesandmoths.org, butterfliesofamerica.com, and iNaturalist.org, though

exacting measurements were not possible (Table 2). Additional web-sourced imagery was available but

not referenced for this study.

WOV., Raleigh Co., Prince |

WO, Raleigh Co., nr. Grandview State Park

WV. Monroe Co., Jefferson Nat. Forest, Potts Creek

WOW, Wyoming Co., Pinnacle Creek

vA., siratscnnagia

OK., Le Flore Co., Cucumber Creek

TH., Washington Co., Cherokee Nat. Forest

“western AC."

GA, Fannin a., Cooper Crack M (0) F(D)

Heiteman & Heitzman, 1987 M (0), F (OM)

GA., Fannin Co.

AC. Swain Co., Cherokee Reservation

Howell & Charney, 2010 AL., Tallapoosa Co., nr. Chatasotka Creek

Iftner, etal. , 1992

Klots, 1951 AR., Lincoln Co., Tarry

ALC. Transylvania Co., Lake Toxaway

Monroe & Wright, 2017 VA, Montgomery Co., Poverty Hollow MIDN). F(D/AV)

Opler & Krizek, 1984 VA., Bath Co. M (DN), F (DV)

Opler & Malikul, 1998

Shull, 1987 vA, _ Montgomery C5, Powerty Hollow MA (D/V), F (DV)

faa Montgomery Co., Blacksburg M (0), FV)

Smith & Domingue, 2019 NLC., Swain Co., Great Smoky Mountains Nat. Park M (D), F(D)

ai AR., Logan Co., Mt. Magazine State Park M(SD/¥}, F(3D/V)

Spencer, 2014 M (D/M), F (D/V)

Table 1. Summary of images examined for this study in published (print) literature.

[butterfliesofamericacom | [AR..HempsteadCs.. Rick Evans Grandview Prairie WMA [MID]

buttertliesofamericacom | [AR.LoganCo.Mt Magazine StatePark [MIDI FID)

butterfliesofamericacom [| [GA.FaninCo.CooperCreek MIDI FID)

butterfesotamericacom [| [OK CherokeeCo.Cherokee WMA MIDI)

butterfliesofamericacom [| [OK LeFloreCo.QuachitaNationalForest IMIVLFID)

butterfliesofamericacom | (OK. TulsaCo.. Tulsa

fbuttertiesotamericacom [| [TM BlountCo.. Townsend I

[butterfliesofamericacom | [WA MontgomenyCo..PowertyHollow MID]

Table 2. Summary of online image sources examined for this study (continued below).

Table 2. (cont.)

fnsturslstorg _———~—~«*zSTISET

inaturalistorg __———_—~dTTORe36

inaturalstorg __——~«*zSTST TET

o4Tsi04

Table 2. (cont.)

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