taxonID	type	description	language	source
039FB9086513FFAFFD674110FEDFB0FE.taxon	description	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 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 Fig. 1. A. diana male wings comparison. (Arkansas) is 29 mm, confirming that Appalachian males have 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 Cu 1, 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. 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 ”.	en	Pavulaan, Harry (2021): Subspecific designation of the U. S. A. Interior Highlands population of Argynnis (Speyeria) diana (Cramer, 1777) (Nymphalidae: Heliconiinae: Argynnini: Argynnina). The Taxonomic Report of the International Lepidoptera Survey 9 (9): 1-14
039FB9086515FFADFD2942E6FB3AB7E2.taxon	description	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 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 ”. 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, 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. Recentlyemerged 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).	en	Pavulaan, Harry (2021): Subspecific designation of the U. S. A. Interior Highlands population of Argynnis (Speyeria) diana (Cramer, 1777) (Nymphalidae: Heliconiinae: Argynnini: Argynnina). The Taxonomic Report of the International Lepidoptera Survey 9 (9): 1-14
039FB9086515FFADFD2942E6FB3AB7E2.taxon	etymology	Etymology: The subspecies name arkansana represents the primarily Arkansas portion of the subspecies range. In 2007, it was designated as the Arkansas State Butterfly.	en	Pavulaan, Harry (2021): Subspecific designation of the U. S. A. Interior Highlands population of Argynnis (Speyeria) diana (Cramer, 1777) (Nymphalidae: Heliconiinae: Argynnini: Argynnina). The Taxonomic Report of the International Lepidoptera Survey 9 (9): 1-14
039FB9086515FFADFD2942E6FB3AB7E2.taxon	materials_examined	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.	en	Pavulaan, Harry (2021): Subspecific designation of the U. S. A. Interior Highlands population of Argynnis (Speyeria) diana (Cramer, 1777) (Nymphalidae: Heliconiinae: Argynnini: Argynnina). The Taxonomic Report of the International Lepidoptera Survey 9 (9): 1-14
