Volume 4

1 March 2003

Number 1

The Taxonomic Report

FIXATION OF TYPE LOCALITY FOR LYCAENA ACMON WESTWOOD AND

CHARACTERIZATION OF THE SPECIES AND ITS DISTRIBUTION

PAUL A. OPLER 1

Department of Bioagricultural Sciences and Pest Management,

Colorado State University, Fort Collins, Colorado 80523

ABSTRACT. Lycaena acmon Westwood, 1852 is based on a painting and plate legend in Westwood and Hewitson’s Genera

of Diurnal Lepidoptera. The specimen illustrated was located in the British Museum Natural History and is the holotype by

monotypy. The accompanying plate legend gives “California” as the type locality. Because the butterfly is a member of a

complex of species, now considered in the genus Plebejus (Opler and Warren, 2003), that requires much systematic study, and

some confusion exists on the identity of L. acmon, it is necessary to fix a more specific type locality, to characterize the

species, and to present its synonymy. In addition, a number of taxa described or cited as P. acmon are listed, but which likely

represent other species.

Additional key words: Plebejus.

THE “ACMON COMPLEX”

Lycaena acmon Westwood, 1852, was the first of a number of names that refer to a cluster of

described taxa that occur in western North America. Several of these names have been associated as

infraspecific taxa with acmon by various authors, including Downey, 1951, Goodpasture, 1973a, Tilden,

1973, Scott, 1998, and Emmel et al., 1998b. In recent decades, the complex has been treated as being

comprised of three species — acmon Westwood, lupini Boisduval, and neurona Skinner (Goodpasture,

1973a, Tilden, 1973, Scott, 1998) with attendant subspecies. Previously, authors treated the non -neurona

members of the complex as ranging from one species — acmon (Downey, 1961) to as many as five

species (McDunnough, 1938).

Goodpasture (1973a) considered the genitalia of the complex members to be “bafflingly

complex,” but settled on a narrow definition of “lupini” as being comprised of a set of California

Eriogonum obligates, and “acmon” as being comprised of a widespread set of populations in western

North America which feed on both legumes and Eriogonum. More recently, Scott (1998) reinterpreted

Goodpasture’s findings about the group’s genitalia and applied “lupini” more broadly in the West, and

“acmon ” more narrowly.

In the last few years, several workers, e.g. Davenport (2003) and Opler (unpublished data), have

observed as many as three “non -neurona” entities of the complex occurring in sympatry or parapatry

without hints of intermediacy. Furthermore, as hinted at by Shields (1975), the many populations in the

“acmon” complex have evolved together with those of Euphilotes, and it seems that many acmon complex

populations are almost as closely associated with their Eriogonum hosts as are their convergently evolved

ecological brethren, the genus Euphilotes (Opler, unpublished). It is possible, when research has been

Research Associate, Department of Entomology, Smithsonian Institution, Washington, D.C. 20560

-Museum Associate, Essig Museum of Entomology, University of California, Berkeley, Calif. 94720

completed, that the acmon complex will turn out to be composed of a cluster of species, some currently

without names. In order to make the research more orderly it is necessary to fix the type locality of

Lycaena acmon Westwood, a pivotal species to the understanding of the complex.

THE TYPE OF LYCAENA ACMON WESTWOOD

The “description” of Lycaena acmon Westwood appears in the work Genera of Diurnal Lepidoptera

by Westwood and Hewitson (1852). Westwood is the author of the name and Hewitson painted the

representation of the specimen that appears in volume 2 on Plate 76 (Figure 3), dated 5 th December 1851.

The legend for the plate was published later in volume 1 (1852) (Figure 4). The specimen figured on the

plate was located in the Natural History Museum (London) (Figures 1 & 2) by Phillip Ackery and bears

several labels (Figure 5). In correspondence, Dr. Ackery indicates that his current judgment is that the

specimen in the collection is likely the specimen that was illustrated. Accordingly, I have sent a red label

which reads: “holotype, Lycaena acmon , det. Paul Opler 2003” to be placed on this specimen. 2

Figures 1-5. Fig. 1. Holotype, Lycaena acmon , dorsal. Fig. 2. Holotype, Lycaena acmon, ventral. Fig. 3. Lycaena acmon fig. 2,

Plate 76, Volume 2, Westwood & Hewitson, 1852. Fig. 4. Text from Volume 1, Westwood and Hewitson, 1852. Fig. 5. Labels

from Fig. 1 holotype in British Museum Natural History.

The holotype is a male and an individual of the generations seen in late spring, summer, and fall,

clearly that which is considered to be typical lowland Californian Plebejus acmon (Goodpasture, 1973a;

Tilden, 1973). Males of the first generation that fly as early as February represent overwintered

individuals, and are characterized by a more grayish appearance below, more blackish scaling above, and

a deeper shade of blue. Dorsally, the illustrated individual is a pale lilac blue above with a submarginal

salmon-pink aurora with what appear to be 6 contained black spots on the hindwing. Both wings have a

thin continuous solid black terminal line and white fringes. Ventrally, the wings have a pale whitish gray

ground with the black terminal line and white fringes repeated. There is a comma-shaped black mark at

the end of the forewing discal cell and a tiny black point at the inner edge of the forewing discal cell.

Additionally, the outer half of the fore wing has two rows of black spots. The postmedial is an outwardly

curved row of six spots; the 3 rd , 4 th , and 5 th from the costal margin are elongate and tilted more or less

along the long axis of the wing. The row lying between the submarginal row and a submarginal line of

smudged black dashes also contains six spots, but they follow the curvature of the wing, are somewhat

squarish and become gradually larger proceeding toward the inner margin. On the hindwing, there are a

2 Editor’s note. The editor submitted this situation to the advise of the ICZN Co mmi ssioners and other taxonomists via the

ICZN’s official internet list-serve. The consensus of that group was that this specimen is best viewed as the holotype by

monotypy and not a syntype. The Vol. 4 Code is more explicit and restrictive on typification. Holotypes are not to be assumed

(Rec. 73F) and when previous instances where this has occurred is detected, the inferred holotype automatically becomes a

lectotype (§ 74.6). In this case, the sole specimen illustrated is captioned as deposited in the British Museum and is thus

designated by monotypy as the holotype.

2

number of more or less equally sized roundish black spots as follows: (1) a postbasal costal spot, (2) a

medial postbasal spot, (3) an outwardly curved postmedial row of seven spots, the most inner of which is

a black dash, and (4) a dash on the inner margin and at right angles to it. The aurora is a bright reddish-

orange with four barely touching marks capped with black inwardly arching triangular or rounded marks.

The outward portion of each orange mark contains a black spot surrounded or almost completely

surrounded by iridescent silvery scintillant scales. Toward the anal angle is a contiguous patch of

apparently three merged orange spots sparsely capped inwardly with black and marginal outwardly by

black bars surrounded by scintillant scaling. Toward the anal angle lie two additional black spots with no

subtending orange spots. The dorsal surface of the thorax and abdomen appear blackish, and legs as well

as the ventral surface of abdomen are black covered with white scales. The genitalia of the figured

specimen have not been dissected. Genitalic drawings that probably represent typical P. acmon, but

which are not topotypical, are illustrated by Goodpasture (1973) and Tilden (1973).

FIXATION OF TYPE LOCALITY

At this point, it appears that typical P. acmon is relatively geographically invariable throughout

cismontane (west slope) California. The species is seasonally dimorphic with smaller, darker early

spring individuals, and sexually dimorphic with blue males and dark brown females (early spring females

have some blue scaling above basally). There is no indication where in California the holotype may have

been collected. It is possible that the specimen was collected in central or northern California by Lorquin,

although there is no direct evidence. I feel it will be useful to select a type locality in central California

where no related species occur. Furthermore, since the type localities of two subjective junior synonyms

of P. acmon Westwood ( Lycaena antaegon Boisduval, 1852 [the type of which may be part of the same

lot from which Lycaena acmon was described (Emmel et al., 1998a)] and Rusticus acmon cottlei Grinnell,

1916) are located at San Francisco, San Francisco County, California, it will leave little doubt as to the

taxonomic identity of acmon and its junior synonyms if the type locality were to be fixed at San

Francisco. Therefore the type locality of L. acmon Westwood is fixed here at San Francisco, San

Francisco County, California.

SYNONYMY OF PLEBEJUS ACMON WESTWOOD

The type locality of both junior synonyms is San Francisco. The type locality of Lycaena

antaegon was fixed by Emmel et al., 1998a. Rusticus acmon cottlei was described as a subspecies, but its

phenotype represents the spring form of the species in which the female has extensive blue scaling and

both sexes are small and have wings with wide black borders. These individuals occur at the same sites

where later generations have the typical P. acmon phenotype. Note that here and below, infrasubspecific

names are not included.

Plebejus acmon (Westwood, 1852), Genera of Diurnal Lepidoptera. Vol. 2, page 494.

Lycaena antaegon Jean Alphonse Boisduval, 1852. Lepidopteres de la Californie. Annales de la

Societe Entomologique de France 12: 295-296. [junior synonym]

Rusticus acmon cottlei Jordyce Grinnell jr., 1916. An unnamed butterfly from San Francisco.

Pomona Journal of Entomology and Zoology 8: 83. [junior synonym]

3

CHARACTERIZATION OF PLEBEJUS ACMON WESTWOOD

The concept of this species and its relatives have received a different treatment by almost every

author who has studied them (Clemence, 1909; Comstock, 1922; Downey, 1951; Goodpasture, 1973a;

Tilden, 1973; Scott, 1998; and Emmel et al., 1998b). All authors seem to agree on the identity of

nominotypical Plebejus acmon , but many have widely discordant concepts of related taxa. Goodpasture

(1973a) and subsequent authors have included at least one other entity as conspecific with P. acmon.

Goodpasture (1973a) thought that P. acmon was a widespread, variable western butterfly, but this

conclusion was suggested by Scott (1998) as incorrect, and I agree with his conclusion. Three additional

taxa have been described as conspecific with P. acmon, but their authors did not present any species-level

characters supporting this association (i.e. dos Passos, 1938; Goodpasture, 1973b; Emmel et al., 1998b).

In several cases these concepts were further confused by the authors themselves who applied the names

widely beyond their type localities and may have confused them with other taxa, at least some of which

are likely undescribed (Opler, unpublished).

Extensive sampling and observations of Plebejus acmon and its relatives in the field have yielded

much information of the butterflies’ behavior, host plants, and distributions. In California, Plebejus

acmon occurs sympatrically with a number of other taxa, e.g. Goodpasture, 1973a, page 481; Opler,

unpublished. In California, these include Lycaena chlorina (Skinner, 1902), Lycaena lupini (Boisduval,

1869), Rusticus monticola (Clemence, 1909), and Lycaena neurona (Skinner, 1902). In some instances as

in Kern County, California, P. acmon, P. monticola, and P. neurona may be found flying together at the

same site with no apparent intermediates. In cismontane California, all other taxa in the “ Plebejus acmon

complex” utilize perennial species of Eriogonum (Polygonaceae) as their larval hosts and all but the

bivoltine Plebejus neurona are strictly single-brooded. Elsewhere, if the butterfly that seems to represent

texanus is truly widespread in the Southwest, then it is also multivoltine and appears to feed on both

Eriogonum species as well as legumes (e.g. Goodpasture, 1973a, 1973b; Bailowitz and Brock, 1991).

In cismontane California, the populations of butterflies here considered to belong to nominotypic

Plebejus acmon are multivoltine and seem to be continuously brooded where host plant availability and

climate are amenable. In the lowlands, there are certainly at least three broods and possibly as many as

five or six. The butterflies are found everywhere where suitable hosts occur from sea level to as high as

8,000’ (2438 m). There are numerous observations of this butterfly’s use of both species of Eriogonum

and certain genera of legumes ( Lotus species, especially annual species such as L. heermani (G. Pratt, in

litt.), but also L. scoparius (Nutt.) Ottley) (Goodpasture, 1973a, and many others).

Populations of multivoltine butterflies with the nominotypic phenotype and which use either

Eriogonum species or legumes as larval hosts range from along the Pacific coast from as far south as the

Sierra San Pedro Martir of Baja California, Mexico north through cismontane California, through western

Oregon to at least as far as southwestern Washington. Additionally, butterflies which seem to fit this

general description have been seen in the Carson Range of western Nevada, possibly elsewhere in

northern Nevada, and on the east side of the Sierra Nevada in the Owens Valley and perhaps other sites of

transmontane eastern California (Opler, unpublished).

As discussed by Goodpasture (1973a) most populations of the P. acmon complex outside of this

distribution are single-brooded and limited to species of Eriogonum as their larval hosts. The phenotypes

of the butterflies in these populations differ in a number of characters from that of nominotypic P. acmon.

In addition, there are populations of the complex that occur in the Southwest, high plains, and northern

mainland Mexico that are likely multivoltine, and some are reported to utilize legumes as larval hosts.

Most of these latter populations fit the general concept of P. acmon texanus Goodpasture (1973b), but

many of these populations may or may not be conspecific with P. a. texanus itself, and texanus is not

herein considered as conspecific with P. acmon.

In accord with the above discussion, I limit application of the species P. acmon to the nominotypic

populations that occur within the above described geographic distribution. I conclude that all other

populations and the names applied to them are not conspecific with P. acmon. If populations of any other

4

description than given above are conspecific with P. acmon , their placement must await the results of

further morphological, behavioral, and genetic research, some of which is in its initial phases (Opler and

others, unpublished).

EXCLUDED ACMON COMPLEX TAXA

The following taxa [given in their original combinations], listed in chronological order by year of

description, are excluded from the concept of P. acmon. They represent both species-level and

infraspecific taxa. In agreement with previous authors, P. neuroma is considered distinct at the species-

level, but the taxonomic status and level of the remaining names is not evaluated here. The recent

practice of considering “acmon complex” taxa, except P. neuroma , to be subspecies of either P. acmon or

P. lupini is flawed because in several instances more than one non-P. acmon entity have been found

sympatrically or parapatrically without indication of intermediacy. Further research is necessary to clarify

just how many additional species-level taxa are represented in western North America.

Lycaena lupini Boisduval, 1869

Lycaena neuroma Skinner, 1902

Lycaena chlorina Skinner, 1902

Rusticus monticola Clemence, 1909

Plebeius carolyna J.A. Comstock, 1922

Plebejus acmon lutzi dos Passos, 1938

Plebeius lupini spangelatus Burdick, 1942

Plebejus acmon texanus Goodpasture, 1973

Icaricia acmon dedeckera J.F. Emmel, T.C. Emmel, and Mattoon, 1998

Icaricia lupini argentata J.F. Emmel, T.C. Emmel, and Mattoon, 1998

Icaricia lupini alpicola J.F. Emmel, T.C. Emmel, and Mattoon, 1998

Icaricia lupini goodpasturei Austin, 1998

ACKNOWLEDGMENTS

I thank J.D. Lafontaine, Agriculture Canada, Ottawa, and Phillip Ackery, Natural History

Museum, London, U.K. for providing references and images related to the type of Lycaena acmon.

George T. Austin, Nevada State Museum, Las Vegas; Boris Kondratieff, Colorado State University, Fort

Collins; and Gordon Pratt, University of California, Riverside read the manuscript and provided many

comments and corrections which resulted in its improvement.

LITERATURE CITED

Bailowitz, R.A. & J.P. Brock. 1991. Butterflies of Southeastern Arizona. Sonoran Arthropod Studies,

Inc., Tucson, Arizona. 342 pp.

Clemence, V.L. 1909. Notes on the forms of Rusticus acmon (Db.-Hew.), occurring in the vicinity of

Pasadena, Calif. Canadian Entomologist 41: 38-39.

Comstock, J.A. 1922. Studies in Pacific Coast Lepidoptera. Continued. Notes on the acmon-neurona

group of lycaenids, with description of a new species. Bulletin of the Southern California Academy

of Sciences 21: 43-48.

Davenport, K. 2003. Butterflies of Kern and Tulare Counties, California. Butterflies of Sequoia-Kings

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

University, Ft. Collins, Colo., in press.

Downey, J.C. 1951. Lycaenidae. Pages 248-342 in: P.R. Ehrlich and A.H. Ehrlich. How to know the

butterflies. W.C. Brown and Co., Dubuque, Iowa.

Emmel, J.F., T.C. Emmel and S.O. Mattoon. 1998a. The types of California butterflies named by Jean

Alphonse Boisduval: designation of lectotypes and a neotype, and fixation of type localities. Pages 3-

76 in: Emmel, T.C., editor. Systematics of Western North American Butterflies. Mariposa Press,

Gainesville, Florida. 878 pp.

_. 1998b. New Polyommatinae subspecies of Lycaenidae from

California. Pages 171-200 in: Emmel, T.C., editor. Systematics of Western North American

Butterflies. Mariposa Press, Gainesville, Florida. 878 pp.

Goodpasture, C. 1973a. Biology and systematics of the Plebejus (Icaricica) acmon group (Lepidoptera:

Lycaenidae). I. Review of the group. Journal of the Kansas Entomological Society 46: 468-485.

_. 1973b. A new subspecies of Plebejus acmon (Lepidoptera: Lycaenidae). Pan-Pacific

Entomologist 49: 149-159.

McDunnough, J. 1938. Check list of the Lepidoptera of Canada and the United States of America.

Volume 1. Memoirs of the Southern California Academy of Sciences, Los Angeles, 272 pp.

Opler, P.A. and A.D. Warren. 2002. Butterflies of North America. 2. Scientific names list for butterfly

species of North American north of Mexico. Contributions of the C.P. Gillette Museum of Arthropod

Diversity, Colorado State University, Fort Collins, Colo. 79 pp.

Scott, J.A. 1998. New western North American butterflies. Papilio, New Series 11:1-12.

Shields, O.A. 1975. Studies on North American Philotes (Lycaenidae) IV. Taxonomic and biological

notes, and new subspecies. Bulletin of the Allyn Museum 28:1-36.

Tilden, J.W. 1973. Specific entities of the subgenus Icaricia Nabokov (Lycaenidae). Journal of Research

on the Lepidoptera 12: 11-20.

Westwood, J.O. and W.C. Hewitson. 1852. Genera of Diurnal Lepidoptera. Volume 1. Van Vorst,

London.

6

Volume 4

April 2003

Number 2

The Taxonomic

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

Report M

URVEY

BUTTERFLIES OF

THE NORTH OKANAGAN, BRITISH COLUMBIA, CANADA

DAVID L. THREATFUL

INTRODUCTION

The butterfly fauna of British Columbia has been described as part of larger geographic areas in

sundry books (e.g. Guppy and Shepard 2001, Layberry et al. 1998, Scott 1986). But considering the size (90

million hectares) and lengthy history of butterfly study in the province; the butterfly fauna of specific areas

has been poorly documented. Some areas that have received local treatments are the Peace River area

(Kondla et al. 1994), Pend-d’Oreille valley (Kondla 1999), Glacier and Mount Revelstoke National Parks

(Threatful 1982, 1989), Cherry Creek (Threatful 2000a), Red Mountain (Threatful 2000b), Chilcotin

grasslands (Fischer et al. 2000), Lillooet area (McDunnough 1927), Kootenay region (Dyar 1904),

Kootenay Lake (Schmidt 1996), Forbidden Plateau (Hardy 1954) and Shuswap (Buckell 1947). Previous

literature specific to the butterflies of the north Okanagan is limited to Downes (1918) and Threatful (2001).

Blackmore (1920, 1921) and Llewellyn-Jones (1951) mention some butterfly records from Vernon and

Armstrong. The information contained in this report is primarily from my field notes and specimens I

collected 1970 - 2002.1 visited more than 49 discrete sites on various dates and many sites were sampled

repeatedly over the years.

Supplemental information was garnered from the insect collection of the BC Ministry of Agriculture,

Fisheries and Food (BCMAFF) in Kelowna through the courtesy of Hugh Philip. The collection had been in

the office of the Provincial Entomologist in Vernon but when that office was closed the collection was

transferred to BCMAFF, Kelowna. Butterfly specimens in the BCMAFF collection were taken between

1903 and 1932, primarily by staff of the former Provincial and Federal entomology offices in Vernon. The

early entomologists who contributed to the BCMAFF collection were C. Bigland; N.L. Cutler; E.A.

Rendell; M.H. Ruhmann - resided in Vernon 1912-43, Provincial Entomologist 1935-42 (Riegert, 1991);

N.W. Ruhmann; R.C. Treheme - resided in Vernon 1914-22, Head, Field Crop Insect Laboratory,Vernon

1914-22 (Riegert, 1991); R.C. Woodward; and I. J. Ward - resided in Vernon 1912-47, Provincial

Entomologist 1943-47 (Riegert, 1991). From this collection I have been able to include 84 new records

(locations/dates) and three new species of butterflies collected in the north Okanagan. Personal collection

data was also provided by C.S. Guppy and N.G. Kondla. The Canadian Biodiversity Information Facility

was searched by N.G. Kondla for additional historical records.

The earliest known butterfly records for the study area are by J. Fletcher in 1899 in the vicinity of

Vernon and include specimens of Lycaena helloides and Cercyonis sthenele. One or more voucher

specimens are in the collection of the University of Guelph and the Canadian National Collection. A number

of additional people have collected butterflies in the study area over the years: E. Anderson, R.G. Bartman,

R.A. Cannings, R. Carcasson, J. Carr, J.K. Cooper, E. Fuller, A.N. Gartrell, J.L. Gordon, C. Guiget, G.A.

Hardy, G.E. Hutchings, S. Ife, J. K. Jacob, G.H. Lamder, J.R. Lewellyn-Jones, A.P. Mackie, R.P. Nelson, J.

Reichel, E.A. Rendell, G.B. Straley, A. Tate, E.P. Venables; J.B. Wallis.

The subspecies designations should be treated with caution because the taxonomic status of a

number of species in this area is only poorly known. Figure 1 shows the approximate location of the major

sites within the study area where butterflies have been observed or collected. The study area covers

approximately 1900 km 2 .

The common names for the butterflies conform to those provided in Guppy and Shepard (2001).

Zoological names are not drawn from one source but have been mostly researched individually to determine

the most reasonable taxonomic interpretation and resulting nomenclature. I follow Johnson (1992) for the

callophryne elfins and Balint and Johnson (1997) for the polyomatine blues. Common names for plants are

used throughout, with the corresponding botanical names provided on first use of the name.

Specimens of nearly all the butterfly taxa documented in this paper are in a representative collection

that I have donated to the north Okanagan Naturalists Club. The Vernon Museum, as custodian of the

collection, has provided specialized cabinets to house the collection for viewing and study.

NORTH OKANAGAN ECOSYSTEMS

In British Columbia two ecosystem classification systems widely in use - the biogeoclimatic

ecosystem classification and the ecoregion classification systems. The differences and similarities between

the two systems have been well described by Harding and McCullum (1994). For this brief overview of the

study area ecosystems, I have chosen to use a modified version of the biogeoclimatic classification

(Meidinger and Pojar 1991). The study area is on the Thompson Plateau of the north Okanagan Valley.

The ecosystems have characteristic mean annual precipitation regimes, elevation ranges, and

seasonal temperatures. These physical characteristics determine in part the presence of associated flora

including those species of plants used by butterflies for larval food and nectar sources for the adults. The

mean annual precipitation at low elevation Vernon is 30 - 40 cm; that at high elevation on Silver Star

Mountain is 40 - 50 cm (Farley, 1979). I refer to four elevation zones in the habitat descriptions and in the

species accounts: low elevation (400m - 450m); moderate elevation (450m - 1070m); moderately high

elevation (1070m - 1590m), and high elevation (1590m+).

Subalpine Forest

Subalpine Forest occurs at the highest elevations found in the north Okanagan. At these elevations,

Subalpine Fir (/ibies lasiocarpa ) and Engelmann Spruce (Picea engelmannii ) are the predominant trees

species with Whitebark Pine ( Pinus albicaulis) established on the drier sites, as on Terrace Mountain near

Fintry. Subalpine meadows, with a rich assortment of plant species whose flowers are both colourful and

major nectar sources, are characteristic of the higher elevation subalpine forests. Spotted saxifrage

C Saxifraga bronchialis) is present and stonecrop ( Sedum spp) is abundant on rocky sites. Ragwort

(Senecio spp), Indian paintbrush ( Castilleja spp), arctic lupine (Lupinus arcticus ), subalpine daisy

(Erigeron peregrinus), and Sitka valerian (Valeriana sitchensis) grow in the subalpine meadows, with

Indian Hellebore ( Veratrum viride) common along streambeds and seepage slopes. However, species vary

with locality and aspect. For example, Columbian monkshood ( Aconitum columbianum) is present on Silver

Star Mountain, but not on most other mountains in the area. Silver Star Mountain also has shrubs like blue-

3

berries ( Vaccinium spp) and white flowered rhododendron {Rhododendron alhiflorum). On Terrace

Mountain, at 1904 metres, whitebark pine grows in the rocky open subalpine habitat on south facing slopes,

while Engelmann spruce mixed with subalpine fir grows on the north facing slopes.

Four butterflies characteristic of the subalpine forests are the Opis Fritillary {Speyeria mormonia

opis ), the Hydaspe Fritillary {Speyeria hydaspe rhodope ), Vidler’s Alpine {Erebia v idler i), and the Lilac-

bordered Copper {Lycaena nivalis browni).

Mountain Forest

Mountain Forest covers extensive areas of the north Okanagan. It is an intermediate biogeoclimatic

zone between the subalpine forest above and the interior Douglas fir forest below. This ecosystem occupies

much of the Thompson Plateau. Highest elevations of this zone are often dominated by Engelmann spruce

and Engelmann X white spruce hybrids. Shrubs such as Labrador tea (L edum groenlandicum) appear

frequently in bogs, while soopallalie {Shepherdia canadensis ) is abundant on drier sites of semi-open

forests. Lodgepole pine {Pinus contorta) and Douglas fir {Pseudotsuga menziesii) are frequent fire-

successional species.

At lower elevations (ca. 600 m) on north facing slopes and at moderate elevations (ca. 1150 m) the

forest is highly transitional in nature. The forests here are mostly a mix of western larch {Larix

occidentalis), western red cedar ( Thuja plicata ), lodgepole pine and Douglas fir. Drier areas support

shrubs like raspberry {Rubus ssp) and thimbleberry {Rubus parviflorus). Fire weed is often very abundant

on logged and burned sites. On southwest facing slopes the interior Douglas fir forest occurs at higher

elevations than does mountain forest at lower elevations on north facing slopes.

Butterflies characteristic of the Mountain Forest include the Chryxus Arctic {Oeneis chryxus

chryxus ), the Pink-edged Sulphur {Colias interior ), the Persius Duskywing {Erynnis per sms'), Freija

Fritillary {Boloria freija ), Western Meadow Fritillary {Boloria epithore chermocki ), Cedar Hairstreak

{Mitoura rosneri ), Margined White {Pieris marginalis reicheli ), Stella’s Orange-tip {Anthocharis Stella ),

Roadside Skipper {Amblyscirtes vialis) and the Anna’s Blue {Plebejus anna).

Interior Douglas-fir Forest

The Interior Douglas-fir Forest covers a large area of the north Okanagan. In the north Okanagan it is

above the Ponderosa Pine forests, where it merges into the Mountain Forest. It also extends down to lower

elevations, to 400m, on the north-facing slopes of the Thompson Plateau, north and east of Kalamalka Lake.

Douglas-fir is the predominant tree species. In some localities and at higher elevations western larch {Larix

occidentals) blends from this ecosystem into that of the Mountain Forest. Douglas maple {Acer glabrum)

grows at low elevations in the openings on north-facing slopes and black cottonwood {Populus balsamifera

trichocarpa) occurs along sunny riparian areas. In open sunny clearings occur shrubs such as soopollalie,

ocean spray {Holodiscus discolor ), and redstem ceanothus {Ceanothus sanguineus). In sunny openings in

the forest occur flowering plants such as heart-leaved arnica {Arnica cor difolia), tiger lily {Lilium

columbianum), red columbine {Aquilegia formosum ), and Solomon’s -seal {Smilacina spp). Pinegrass

{Calamagrostis rubescens) and rough-leaved ricegrass {Oryzopsis asperifolia) occur in grassy openings

with red Indian paintbrush {Castilleja miniata). Stonecrop is common in sunny breaks on south-facing

slopes. On dry sites of south facing lopes adjacent to Douglas-fir forests are grasslands characterized by

bluebunch wheatgrass {Agropyron spicatum) and rough fescue {Festuca scabrella) growing with big

sagebrush {Artemisia tridentata). Big sagebrush can be locally abundant where soil conditions permit.

Flowering plants are similar to those of the Ponderosa Pine ecosystem grasslands, and include silky lupine

{Lupinus sericeus ), arrow-leaved balsamroot {Balsamorhiza sagittata ), and creamy buckwheat

{Eriogonum heracleoides).

Butterflies that occur in the dry grasslands include the Nevada Skipper (Hesperia nevada ), the

Desert Marble ( Euchloe lotto), and the Rocky Mountain Apollo ( Parnassius smintheus magnus).

Butterflies in the sunny forest openings include the Northern Cloudywing ( Thorybes pylades) and Lorquin’s

Admiral ( Limenitis lorquini itelkae).

Ponderosa Pine Forest and Grassland

Forests dominated by ponderosa pine ( Firms ponderosa) occupy the valley bottoms from Okanagan

and Kalamalka Lakes northward through Armstrong. There it gives way in a transitional zone to the Interior

Douglas-fir forests and the wetter western red cedar and western larch forests. Plants of the dry south¬

facing slopes are mostly shade intolerant species including balsamroot, saskatoon ( Amelanchier alnifolia ),

smooth sumac ( Rhus glabra ), mock orange ( Philadelphus lewisii) and chokecherry ( Prunus virginiana).

These species usually grow among the rocky outcrops with ponderosa pine on steeper terrain. In some areas

bluebunch wheatgrass is still present among open stands of pine but is being replaced by two grasses; rough

fescue on the dry to moist sites and Idaho fescue ( Festuca idahoensis) on cooler, north-facing slopes.

Dry grasslands occur at the periphery of the Ponderosa Pine ecosystem, and are dominated by

bluebunch wheatgrass. Qher plants include silky lupine, creamy buckwheat, balsamroot, and sagebrush

buttercup ( Ranunculus glaberrimus). Where soil conditions are favorable big sagebrush is present.

Tarragon ( Artemisia dracunculus) is abundant on the open slopes. Black hawthorn ( Crataegus douglasii)

is common in the moister draws of the grasslands. Lake borders and riparian areas often have stands of

black cottonwood.

Butterflies that fly in the open areas include the Great Basin Woodnymph ( Cercyonis sthenele

sineocellata ), the Western Pine Elfin ( Incisalia eryphon). The Oreas Anglewing (. Polygonia oreas

threatfuli) occurs mainly in riparian habitats in dry Douglas fir and Ponderosa Pine Forest where Ribes

spp. are present. Butterflies found in the dry grasslands include Sheridan’s Hairstreak ( Callophrys

sheridanii neoperplexa ), the Arrowhead Blue (Glaucopsyche piasus toxeuma ), the Oregon Swallowtail

( Papilio bairdii oregonius), and the Common Sootywing (Pholisora catullus).

Abundance

The five terms used to describe abundance are as follows:

1. Common: A species usually encountered in numbers of >10 each day during its flight period.

2. Uncommon: A species encountered in small numbers <10 each day during its flight period.

3. Rare: A species encountered only on a few days during its flight period.

4. Extremely rare: A species not seen in most years with few records for any given location.

5. Local: A species known only from restricted localities and habitats. It may be further designated as

common or rare depending upon the circumstances.

These terms are all relative. They may only apply in an average year. A species population may be

considerably reduced during wet or cold years and in some cases habitat destruction may decimate a

population.

Flight Periods

Flight periods are fairly constant in seasonable years, as evidenced by the synchronous dates for

both the historic and present day collections. Early in the butterfly season, overwintering species only fly on

sunny warm days of late February and March. During inclement weather there is no butterfly activity.

Weather can delay or advance emergence by two to three weeks.

SPECIES ACCOUNTS

Specific flight dates are given only for the historic records from BCMAFF to provide a published

record and a few other records of note. Abbreviations for institutional record credits are: Canadian

National Collection of Insects and Arthropods (CNC), Royal British Columbia Museum (RBCM), Royal

Ontario Museum (ROM). Records from the Canadian Biodiversity Information Facility

(http://www.cbif.gc.ca/home_e.php) which repeat my own or other local records are not repeated here

since they are easily retrievable.

Skippers, Family Hesperiidae

N orthern Cloudy wing - Thorybes pylades (Scudder)

Records: Vernon; Cosens Creek, northeast side of Kalamalka Lake 730m. Habitat: flies in open sunny clearings,

roadsides, and edges of Douglas-fir forests at moderate elevations. It can be found on moist ground, usually in small

numbers. Absent from open dry grasslands. Flight period: May to early July.

Dreamy Duskywing - Erynnis icelus (Scudder and Burgess)

Records: Vernon; Bella Vista 595m; Ellison Ridge 600 m. Habitat: moderate elevations, in grassy areas at forest

edges and in gullies, often where there are willows and aspen groves. Absent from open, dry grasslands. Flight

period: mid-May to early July.

Pacuvius Duskywing - Erynnis pacuvius lilius (Dyar)

Records: Spionkop Hill on Ellison Ridge 750m Habitat: absent from the open, dry grasslands. It flies in open dry,

rocky areas with snowbrush ( Ceanothus velutinus ) in Ponderosa pine forests located at moderate elevations. Flight

period: late May through June. Note: In BC this is generally viewed as a rare and local species but it is likely more

widespread and abundant than records suggest due to the ease with which it can be overlooked while flying with other

Erynnis species.

Persius Duskywing - Erynnis per sins (Scudder)

Records: Vernon; Vernon Hill, east of Vernon, 1160m; King Edward Lake, southeast of Vernon, 1340m; Cosens

Bay, Kalamalka Lake Prov. Park 400m; Aberdeen Lake road; Goose Lake; Shorts Creek canyon; Ellison Ridge;

Silver Star Mtn. (ROM). Habitat: flies in a wide range of habitats at low to moderately high elevations. In the spring,

low elevation sites include the open grasslands as well as openings at forest edges, roadsides, clearings, and damp

ground at lakeshores. At moderate elevation, such as Vernon Hill it flies in lupine-rich grasslands adjacent to stands of

Douglas-fir. At moderately high elevations, such as the King Edward Lake area, it frequents open fen edges. Flight

period: May to early July.

Two-banded Checkered Skipper - Pyrgus ruralis (Boisduval)

Records: Vernon Hill, 1220m; Kalamalka Lake 550m; Aberdeen Lake road 1250m; Silver Star Mtn. road 1150m;

Armstrong (CNC) Habitat: flies near forest edges, clearings and roadsides at moderate to moderately high elevations;

no records for lower elevations or open dry grasslands. Flight period: late April through June.

Checkered Skipper - Pyrgus communis communis (Grote)

Records: known only from a few historical records by N.W. & M.H. Ruhmann and R.C. Woodward at Vernon in

1918. Habitat: no habitat information exists with the historical specimens; lack of subsequent records suggests the

species may no longer be present in the study area. Flight period: The historical records are dated July 7, August 13

and August 19.

Common Sootywing - Pholisora catullus (Fabricius)

Records: West side of Kalamalka Lake, 400m; Kalamalka Lake Prov. Park 456m. Habitat: low, open, dry areas of

the grasslands, and along lakeshores, roadsides and trails. Flight period: at least two generations per year; from April

through September.

6

- Car ,

le open dry grasslands of th

rary presence in the study area is q

e: In contrast to the north Okanagan it can be common in th

is yet due to insufficient late season

■ado Idaho (W.H. E

V area and it is absent from the dry

and the other from a ski area parking lot. Flight period: Probably late May throu;

viewed as a rare species in BC. In 2001 I found it common in a spring flight and a lai

Records: Lowef Shorts Creek valley,^80 m; Aberdeen Lake road, 1250 m. Habita

dry grassy meadow, while the Aberdeen Lake road site is a wet grassy area. Flight pi

Tawny-edged Skipper - Polites themistocles turneri (Freeman)

Records: Vernon, 28-VI-1903 (collector unknown); Bella Vista area 700m; Goose Lake 500m; Aberdeen Lake road

1250m; Armstrong (RBCM). Habitat: flies at moderate elevations over the wet grassy margins of small ponds and in

open grassy areas in stands of aspen. Absent from the open dry grasslands. Flight period: mid-June into early July.

Woodland Skipper - Ochlodes sylvanoides (Boisduval)

Records: Vernon, 15-VIU-1932, 16-VUI-1932 (collector unknown); Goose Lake 500m; Kalamalka Lake Prov. Park

400m; Silver Star Mtn. 1650m; Lavington 535m. Habitat: very common in the open dry grasslands at low to

moderate elevations. It visits damp ground in large numbers and often nectars on knapweed (Centaurea spp) and

thistles ( Cirsium spp). Flight period: July through September. Note: This species is the most abundant skipper

during the summer and early autumn.

Roadside Skipper - Amblyscirtes vialis (W.H.Edwards)

Records: Vernon, 20-V-1904, 26-V-1905, VI-1907, 30-VI-1906 (collector unknown); 14-VI-1919 (M.H. Ruhmann);

Silver Star Mtn. 1150m; Ellison Ridge 620m. Habitat: flies along forest roads, trails, forest clearings, and riparian

areas at moderate elevations. Absent from the open dry grasslands of the north Okanagan. Flight period: mid-May to

early July.

Swallowtails and Parnassians, Family Papilionidae

Rocky Mountain Apollo - Parnassius smintheus magnus (Wright)

Records: Vernon VI-1905 (collector unknown); Vernon, 10-VI-1918 (M.H. Ruhmann); Vernon Hill 1160m; Terrace

Mtn. 1905m; Silver Star Mtn. 1890m; Shorts Creek canyon 1500m; Aberdeen Lake road 1100m; Okanagan Landing

(RBCM). Habitat: flies at moderate to high elevation sites with rocky ridges, and open rocky grassy slopes with its

larval food plant, stonecrop. Flight period: mid-June through August, depending on elevation. Note: Most of the

research for Guppy (1986) was completed on Vernon Hill and Silver Star Mountain.

Baird’s Swallowtail - Papilio bairdii oregonius W.H. Edwards

Records: Vernon 1-VI-1909, 5-VHI-1909, 29-IV-1922 (M.H. Ruhmann); Goose Lake area 500m; Vernon Hill 1065

to 1250m; Kalamalka Lake Prov. Park 457m; Bella Vista area 790m; Ellison Ridge 600 to 900m; west side of

Kalamalka Lake 400m. Habitat: flies over rocky, open areas and knolls in the dry grasslands at low to moderate

elevations. Often seen on damp ground and during summer nectaring on thistles. Males occur in numbers on rocky

hilltops above Goose Lake. Flight period: late April through September, with two generations per year. Note: Often

treated as a subspecies of the European species P. machaon. Pyle (2002) treats oregonius as a full species.

Anise Swallowtail - Papilio zelicaon Lucas

Records: Vernon, 12-VII-1924 (E.A.Rendell); Bluenose Mtn. 1220m; one specimen was seen at Cosens Bay,

Kalamalka Lake Prov. Park 400m; Silver Star Mtn. 1890m; Bella Vista area 790m; Aberdeen Lake road 1250m;

Bluenose Mtn. 1213m; Shorts Creek canyon 1500m. Habitat: an uncommon species, flying at low to high elevations

in open forest edges, subalpine meadows, mountain summits, and riparian areas. It frequents damp ground along

forest roadsides. Males hilltop. In the spring it is often seen nectaring on common dandelion ( Taraxacum officinale ).

Flight period: late April through early August, depending on elevation.

Canadian Tiger Swallowtail - Papilio canadensis Rothschild and Jordan

Records: Goose Lake area 500 m; One specimen observed at Kalamalka Lake Prov. Park 400m; Ellison Ridge

600m; Silver Star Mtn. 1150m; Shorts Creek canyon 1130m; Aberdeen Lake road 1250 m. Habitat: flies in open

forest edges, riparian areas and clearings. It gathers in large numbers on damp ground. It has many nectar sources

including spreading dogbane ( Apocynum androsaemifolium) and tiger lily. Flight period: late May through early July.

However, the one specimen seen in late September at Cosens Bay was in perfect condition.

Western Tiger Swallowtail - Papilio rutulus rutulus Lucas

Records: Ellison Ridge 580 to 650m; Kalamalka Lake Prov. Park 400m; Shorts Creek Canyon 580m. Habitat: flies

in open sunny places, in similar habitat as P. canadensis with which it is reported to hybridize. Both species and the

hybrid can be seen on wet ground along the shoreline at Cosens Bay. It has many nectar sources including mock-

orange and american vetch. Flight period: late May through early July.

Pale Swallowtail - Papilio eurymedon (Lucas)

Records: Vernon 31-V-1907 (collector not known); Ellison Ridge 750 to 914m; Bluenose Mtn. 1215m; Shorts Creek

canyon 580 m; Kalamalka Lake Provincial Park 400m; Bluenose Mtn. 1213m; Aberdeen Lake road 1250m. Habitat:

low to moderately high elevations in the cooler Douglas-fir areas where it inhabits open forest edges in association

with redstem ceanothus. It frequents moist areas along lake shorelines. Males hilltop. Various nectar sources include

red columbine and orange honeysuckle. Flight period: late May to early July, although the specimen observed at

Cosens Bay in September was in perfect condition.

Two-tailed Swallowtail - Papilio multicaudatus pusillus Austin and Emmel

Records: Vernon, 13-VI-1927 (I.J. Ward); Goose Lake area 500m; Kalamalka Lake Prov. Park 400m; Ellison Ridge

580m. Habitat: forest edges, riparian areas, clearings, and open sunny places. It frequents moist ground and takes

moisture along lake shores. At bw to moderate elevations it often flies high above its larval food plant, chokecherry.

Nectar sources include thistles. Flight period: late May through August. Note: Individuals of this species are the

largest of the swallowtails in western Canada.

Whites and Sulphurs, Family Pieridae

Pine White - Neophasia menapia tau (Scudder)

Records: Silver Star Mtn. 1150m and 1650m; Shorts Creek canyon 1524m; Lavington (CNC); Oyama (RBCM).

Habitat: most often seen at open forest edges including those in the subalpine, roadsides, and clearings. It takes

nectar from asters. The larval food plant is ponderosa pine as well as other conifer species within its range. In some

years the adults are abundant around these host trees. Flight period: mid-July to mid-September.

Becker’s White - Pontia beckerii (W.H. Edwards)

Records: Goose Lake 500m, Coldstream 500m (R.P. Nelson). Habitat: It is rare at moderate elevations. Flight

period: July.

Spring White - Pontia sisymbrii flavitincta (Comstock)

Records: Vernon, l-V-1920 (M.H. Ruhmann); Kalamalka Lake Prov. Park 455m; Vernon Hill 1189m; Shorts Creek

canyon 1067m; Winfield area 500m; Bella Vista area 750m; Kalamalka Lake 490m; Ellison Ridge 670 m. Habitat:

low to moderate elevations in dry open grasslands, rocky ridges near open forests, and on wet ground along roadsides.

Flight period: mid-April to May. Note: Local populations of the Spring White fluctuate yearly from common to rare.

Western White - Pontia occidentalis occidentalis (Reakirt)

Records: Vernon, VI-1905 (not identified); Vernon, 16-IX-1932 (C. Bigland); Goose Lake 500m; Kalamalka Lake

400m; Shorts Creek Canyon 1524m; Terrace Mtn. 1904m; Aberdeen Lake road 1250m; Silver Star Mtn. 1890m;

Bella Vista area 750m; Ellison Ridge 914m; Winfield (CNC). Habitat: It occurs from low to high elevations in open

dry grasslands, open forest edges, roadsides, agricultural lands, and on mountain summits. The spring generation is

local and rare. The summer generation flies in alfalfa fields during the late summer and early fall along with Colias

eurytheme and C. philodice. It nectars from asters as well as alfalfa. Flight period: mid-April to September, or early

October in some years, with usually two generations per year. Note: Adults of the first generation in the spring are

smaller and have darker veination on the ventral hind-wing than those of the summer generation.

Cabbage White - Pieris rapae (Linneaus)

Records: Vernon, IX-1914 (M.H. Ruhmann); Vernon, IX-1917 (M.H. Ruhmann); Vernon, 6-VIII-1920 (N.L.

Cutler); Goose Lake 500m; Ellison Ridge 580m; Kalamalka Lake 456m; Aberdeen Lake road 1250m; Armstrong area

480m; Silver Star Mtn. 1650m. Habitat: This species was introduced from Europe into Quebec about 1860 and

rapidly extended its range throughout North America. It occurs in all habitats, particularly agricultural lands and in

9

10

Hedgerow Hairstreak - Satyrium saepium okanaganum (McDunnough)

Records: upper Shorts Creek Canyon 1463 to 1575m. Habitat: Found locally in the study area at moderate

elevations in association with Ceanothus velutinus. Flight Period: late June to late August. Notes: I have found it to

be abundant in some years. Nectaring has been observed on Sedurn lanceolatum.

Immaculate Green Hairstreak - Callophrys affinis washingtonia Clench.

Records: Goose Lake area 500m; Kalamalka Lake Prov. Park 456m; Kalamalka Lake, west side, 400m; Ellison

Ridge 600 to 700m; Bella Vista area 790m. Habitat: open dry Ponderosa pine forests and rocky knolls and ridges in

dry grasslands. It gathers on moist ground in these sites. It is rare and local at low elevations in the grasslands, and is

uncommon at moderate elevations. Flight period: mid-May to June.

Sheridan’s Hairstreak - Callophrys sheridanii neoperplexa Barnes and Benjamin

Records: Kalamalka Lake Prov. Park 460m; Goose Lake 500m; Ellison Ridge 580 to 730m; Bella Vista area 610m;

Okanagan Landing (CNC). Habitat: dry open grasslands often near rocky places and trails where it perches upon low

vegetation. Adults infrequently take moisture on damp ground and sometimes nectar on fern-leaved desert parsley

(Lomatium dissectum ). Flight period: mid-March to mid-May although Ihe peak flight is normally early April into

mid April. It is one of the first species to appear in the spring. Note: There are differing views on the correct

subspecies name for this area. Guppy and Shepard (2001) and Layberry et al. (1998) use newcomeri, while Pyle

(2002) limits the distribution of newcomeri to areas south of British Columbia. Further work on this is needed.

Thicket Hairstreak - Mitoura spinetorum (Hewitson)

Records: Ellison Ridge, west of Winfield, 750m; Westside Road, Okanagan Lake, nr. Nahum, 410m; Shorts Creek

canyon 1524m. Habitat: In the north Okanagan it is extremely rare and spotty. It usually occurs at moderate

elevations in open dry transition forests of Douglas-fir and ponderosa pine/lodgepole pine and along forest roads. It

takes nectar from snowbrush, a species of pussytoes ( Antennaria ) and dandelions. Flight period: mid-April into early

July. Note: Some authors place this species under a supergenus concept of Callophrys, while others use a microgenus

concept of Loranthomitoura. I take a middle of the road approach.

Cedar Hairstreak - Mitoura rosneri Johnson

Records: observed along Cosens Creek, 600m northeast side of Kalamalka Lake. Habitat: moderate elevations,

along roadsides and forest edges. Its habitat must include its larval food plant, western red cedar. It is not found in the

open dry forests and grasslands of the north Okanagan. Flight period: mid-May through June. Note: Some authors

treat this species as being conspecific with M. nelsoni but there is no published evidence to support this view.

Juniper Hairstreak - Mitoura barryi Johnson.

Records: Bluenose Mtn. 1150m; Kalamalka Lake 490m; Jade Bay, Kalamalka Lake Prov. Park 400m; Shorts Creek

canyon 1067m; Armstrong (CNC). Habitat: On the west side of Kalamalka Lake it is often seen in small numbers in

rocky dry grasslands, nectaring on sources close to their larval food plant, Rocky Mountain Juniper. This subspecies

flies locally at low elevations in our area, and is rare at moderately high elevations. The single adult recorded at Jade

Bay was on damp ground. Flight period: early April to May. Note: This butterfly has been placed as a subspecies of

M. siva or M. grynea by some authors, however, there is no published evidence to support these interpretations.

Brown Elfin - Deciduphagus iroides (Boisduval)

Records: Silver Star Mtn. road 550m; Bluenose Mtn. 1150m; Winfield area 500m; Bella Vista area 550m; Kalamalka

Lake 490m; Shorts Creek canyon 1128m; Ellison Ridge 580m; Armstrong (RBCM). Habitat: uncommon at

moderate elevations, where it flies in open forest edges and clearings and visits damp ground along forest roads. It

occurs in numbers near the summit of Bluenose Mountain, from mid- to late-May, perching on Common Juniper.

Flight period: mid-April to early June.

Moss’ Elfin - Deciduphagus mossii schryveri Cross

Records: Vernon, 2-VI-1905 (collector unknown); Kalamalka Lake Prov. Park 456m; Ellison Ridge 580 to 730m;

Shorts Creek canyon 1158m; Bella Vista area 790m; Okanagan Landing (CNC). Habitat: local and sometimes

common at moderate elevations; inhabits the dry grasslands in rocky areas where its larval food plant, lance-leaved

12

stonecrop grows. It sometimes takes nectar from sagebrush buttercup, and occasionally visits damp ground. Similar to

C. sheridanii which it flies with. It often lands on dry rocky ground and perches upon low vegetation in the

grasslands. Flight period: late March through early May. This species, along with C. sheridanii , is the first to appear

in the spring other than those species whose adults overwinter.

Hoary Elfin - Deciduphagus polios obscurus (Ferris)

Records: Armstrong, 30-V-1920 (M.H. Ruhmann); Vernon Hill 1160m; Kalamalka Lake 550m; Bluenose Mtn.

1050m; Armstrong (RBCM). Habitat: occurs at a moderately high elevation on Vernon Hill. In the open, dry

Douglas-fir transition forest it flies low to the ground where its larval food plant Kinnikinnick grows. It is not found at

low elevations in the dry grasslands. Flight period: late April into early June.

Western Pine Elfin - Incisalia eryphon (Boisduval)

Records: Kalamalka Lake Prov. Park 500m; Shorts Creek canyon 1463m; Ellison Ridge 600m; Winfield area 500m;

Silver Star Mtn. 1150m. Habitat: uncommon but occurs near open dry Ponderosa pine forests at low to moderate

elevations, and visits moist ground along open forest roads. Adults often perch on the needles of regenerating

ponderosa pine that are 150-180 cm high. Flight period: Late April to June.

Gray Hairstreak - Strymon melinus setonia McDunnough

Records: Goose Lake 500m; Kalamalka Lake Provincial Park 456m; Kalamalka Lake 550m; Ellison Ridge 600m;

Shorts Creek canyon 580m. Habitat: flies at low to moderate elevations in the dry grasslands and rocky ridges on the

west side of Kalamalka Lake. It also occurs in agriculturally disturbed areas and along trails in open dry forests. From

mid- to late-September this species, along with H. Colorado idaho and O. sylvanoides, often takes nectar from

common rabbit brush. Flight period: mid-April to early September, with two generations per year.

Western Tailed Blue - Cupido amyntula (Boisduval)

Records: Vernon, 13-IV-1907 (collector not identified); Vernon, 14-VI-1919 (M.H. Ruhmann); Kalamalka Lake

Prov. Park 400m; Ellison Ridge 600m; Shorts Creek canyon 1127m; Vernon Hill 1219m; Armstrong (RBCM).

Habitat: absent from the dry open grasslands, and is uncommon at low elevations in the open forest edges and

riparian areas of Douglas-fir forests. It oviposits on peavine and vetch ( Vicia ). Flight period: late April through June.

Note: This butterfly has in North America been traditionally placed in the genus Everes but due to extremely minor

differences between Cupido and Everes, I treat Everes as a junior subjective synonym of the older name Cupido, as

has been done by some other recent authors.

Western Spring Azure - Celastrina echo (W.H. Edwards)

Records: Kalamalka Lake Prov. Park 455m; Ellison Ridge 580 to 730m; Shorts Creek canyon 1127m; Goose Lake

500m; Bella Vista area 610m; Aberdeen Lake road 1250m; Silver Star Mtn. road 1150m; Winfield area 500m;

Vernon Hill 1189m. Habitat: flies in open forest edges, clearings, and riparian areas, and often gathers in large

numbers on damp ground along forest roads and lakeshores. It is not found in dry open grasslands. Flight period: late

March to mid-May. Note: This nominal taxon needs further study and taxonomic review in southern British

Columbia. Preliminary review by N. Kondla reveals that the phenotypes present in this study area and other interior

locations are not compliant with the nominal taxon echo, differ from the coastal populations of southwestern British

Columbia and are a mix of the nominal taxa nigrescens and bakeri.

Square-Spotted Blue - Euphilotes battoides glaucon (W.H. Edwards)

Records: Goose Lake 500m; Kalamalka Lake Prov. Park 400m; Shorts Creek canyon 580 to 1087m; Ellison Ridge

580 to 610m; Bella Vista area 600m; Vernon Hill 1189m; Winfield (CNC). Habitat: open dry grasslands where its

larval food plant and nectar source, buckwheat, is present. It occurs at low to moderate elevations, often on moist

ground. Flight period: late May to June.

Arrowhead Blue - Glaucopsyche piasus toxeuma F.M. Brown

Records: Goose Lake 500m; Kalamalka Lake Prov. Park 456m; Ellison Ridge 600 to 730m; Armstrong (RBCM).

Habitat: rare, and occurs in open dry grasslands at low to moderate elevations where its larval food plant, lupine, is

present. Adults are sometimes seen on moist ground. Flight period: late April through May.

13

Silvery Blue - Glaucopsyche lygdamus Columbia (Skinner)

Records: Vernon, 20-V-1904 (collector unknown); Vernon, 6-V-1929 (I.J. Ward); Goose Lake 500m; Ellison Ridge

580 to 730m; Kalamalka Lake Prov. Park 400m; Shorts Creek canyon 580 to 1524m; Bella Vista area 595m;

Aberdeen Lake road 1250m; Silver Star Mtn. 1150 to 1650m; Winfield area 500m; Armstong area 480m; King

Edward Lake road 1350m. Habitat: common in openings in forest edges, subalpine meadows, and subalpine forest

clearings. Seasonally, it can be very common in dry open grasslands. It can be observed at low to high elevations

throughout the north Okanagan; including at high elevations on Silver Star Mountain. It is far more common and

widespread than the Arrowhead Blue. It is commonly seen flying near its larval host plant, lupine, in spring. It

sometimes gathers in large numbers on damp ground. Flight period: mid-April to July.

Anna’s Blue - Plebejus anna (W.H. Edwards)

Records: Silver Star Mtn. 1150 to 1650m; Aberdeen Lake road 1250m; Shorts Creek canyon 1524m; Terrace Mtn.

1600m; King Edward Lake road 1350m; Vernon Hill 1189m. Habitat: It occurs at high elevations in open dry

subalpine forests and on drier gravelled sites, and along roadsides with lupines, its larval food plant. Adults sometimes

gather in large numbers on moist ground. Flight period: mid-July through August. Note: Guppy and Shepard (2001)

place study area butterflies as Lycaeides idas scudderi [sic]. The correct spelling of this taxon is scudderii.

Subsequent work by Guppy, Kondla and C. Schmidt (unpublished data) reveals that scudderii is not an appropriate

name to use for study area butterflies. Although most populations of this butterfly in the study area are not

phenotypically compliant with P. anna ricei, I place them with anna as a species at this time because I have found

them laying eggs on lupines but not blueberries. Butterflies at Shorts Creek canyon include both classic P. anna ricei

phenotypes and also P. ‘idas ’ atrapraetextus phenotypes. Aberdeen Lake area butterflies are phenotypically more like

atrapraetextus than they are to the boreal forest taxon scudderii. Butterflies on Silver Star Mtn. are most like anna in

series. Additional research on the taxonomy of these butterflies is needed.

Melissa’s Blue - Plebejus melissa (W.H. Edwards)

Records: Vernon, V-1906 (collector unknown); Vernon, 16-VII-1917, 24-VH-1917 (R.C. Treheme); Goose Lake

500m; Kalamalka Lake Prov. Park 400m; Bella Vista area 600 to 790m; Ellison Ridge 580 to 600m; Kalamalka Lake

400m. Habitat: It flies in open, dry grasslands and roadsides in ponderosa pine and Douglas-fir forests. Adults

frequent moist ground and use a variety of nectar sources. Lupine is its probable larval food plant. Flight Period:

mid-May through September, with at least two generations per year.

Greenish Blue - Aricia saepiolus arnica (W.H. Edwards)

Records: Vernon Hill 1220m; Silver Star Mtn. 1150 to 1650m; Aberdeen Lake road 1250m; Shorts Creek canyon

1219m; Kalamalka Lake Prov. Park 400m; King Edward Lake road 1350m; Armstrong (RBCM). Habitat: it

frequents moist areas in open forest edges, bogs, grassy fields, meadows, and roadsides, but also flies in drier sites at

moderate elevations. It is uncommon at high elevations on Silver Star Mountain. It is not found in the dry open

grasslands and open forests at lower elevations of the north Okanagan. Adults take nectar at the introduced perennial

white clover ( Trifolium repens), its larval food plant. Flight period: June to August, depending on elevation.

Boisduval’s Blue - Aricia icarioides (Boisduval)

Records: Vernon, 14-VI-1917 (R.C. Treherne); Goose Lake 500m; Kalamalka Lake Prov. Park 400m; Shorts Creek

canyon 1158m; Ellison Ridge 600m; Bella Vista area 595m; Vernon Hill 1189m; Silver Star Mtn. 1150m; Armstrong

area 480m; Coldstream (RBCM). Habitat: common in open dry grasslands at low to moderately high elevations,

where there is lupine, its larval food plant. It often gathers in large numbers on moist ground. Flight period: late May

to July. Note: Study area populations have historically been assigned to ssp pembina but this needs to be reconsidered

because they are different in size and appearance from the southern Alberta populations which have also been

historically assigned to the same ssp.

Lutz’s Blue - Aricia lupini lutzi (dos Passos)

Records: Goose Lake 500m; Kalamalka Lake Provincial Park 400m; Shorts Creek canyon 1524 m; Ellison Ridge

580m; Vernon Hill 1189m; Bella Vista area 610m; Winfield (CNC). Habitat: low to moderate elevations in open dry

grasslands where there is buckwheat, its larval food plant and nectar source. It frequents moist ground, sometimes in

large numbers. Flight period: late May to July. Note: This butterfly has been variably placed in the literature at the

14

Brushfoots, Family Nymphalidae

ar Mtn. Road 1150 to 1650m; Shorts Creek c<

.f open dry Douglas-fir ft

n 1158m; Silver Star Mtn. 1150 to 1

is of this group of g

d by Guppy & Shepard (2001) o

(CNC).

d to the open dry Interior Douglas-ftr fo

m and local in the p<

it open grasslands in the dry ponderosa pine and Ir

5t yet found in the dry gi

n open dry grasslands and a

e it overlaps the range of S. 4

>r B. myrina is supported by tl

i 1370 m; one specimen seen ii

I-, 17-VI-1905, 16-VI-1906 (collector i

Kalamalka Lake Prov. Park 400m;

m; Armstrong (CNC). Habitat: at lc

tt (1994), use of the m

r. Park 400m; west side of K

n BC. In Kalamalka Lake Provincial Park its habitat is n

te is an unusually high el

st side of Kalamalka Lake 4

•d (2001) use a

jctor unknown); Vernon, ll-VI-1917 (R.C. Treherne); Vernon, 26-V-1924 (M.H.

hmann); Ellison Ridge 600 m; Kalamalka Lake Prov. Park 400m; Shorts Creek canyon 1069m; Goose Lake area

3 to 535m; Bella Vista area 610m; Armstrong area 480m; Vernon Hill 1127m; Aberdeen Lake road 1250m;

eshores and open dry grasslands and rocky ridges, to open subalpine forests with meadows, ridges and rockslides.

Flight period: It lies from late April to early September depending on elevation. Note: Local populatio

assigned to ssp hopfingeri by Guppy and Shepard (2001) but they generally do not comply well with th

d in Y Lake area, west of G

al is not found in open dry forests and gi

i, 5-VII-1918 (M.H. Ruhmann); Vernoi

m, 16-VI-1924 (E.A. Rendell); Ellison

ts Creek canyon 579m; Goose Lake 5<

13-VI-1919 (R.C. T

.n Wood Nymph - Cercyonispegala ariane (Boisduval)

i: Vernon 15-VIII-1932 (C. Bigland); Goose Lake 500m; Kalamalka Lake Prov. Park 457m; Shorts Creek

y high elevations. On th

20

country at low elevations where showy milkweed, its larval food plant, is present. Adults take nectar from many

species of flowers. The Goose Lake specimen was collected while it was flying around a clump of showy milkweed.

Flight period: Migrates into the north Okanagan by late June. The summer brood probably leaves the area by hte

August or early September. Note: It has only been found as a few individuals in this study area.

DISCUSSION

To date I have been able to confirm the presence of 106 butterfly species in the north Okanagan

study area as defined in this paper. Additional species that have been found nearby and which may yet be

found in this study area include: Pieris oleracea , Satyrium californicum , Speyeria aphrodite , and Boloria

bellona. Guppy and Shepard (2001) show Hesperia comma manitoha , Boloria bellona, Euphydryas

editha and Cupido comyntas as present in the study area but I have not been able to verify these records. It

is unlikely that alpine specialists will turn up in the study area due to lack of alpine habitats. Fieldwork

over the past 100 years shows that Ihe study area is one of the premiere areas in Canada for species level

butterfly diversity. More than one third of all known Canadian butterfly species have been found in this

relatively small area.

There are no endangered butterflies in the study area. Guppy et al. (1994) and Kondla et al. (2000)

have reviewed the butterflies of conservation concern in British Columbia. Guppy and Kondla (2000)

assigned provincial conservation status ranks of S3-Sensitive to the following study area taxa: Pyrgus

communis , Polites sabuleti , Callophrys affinis , Lycaena nivalis , Danaus plexippus. These are species that

are not believed to be at risk of extirpation or extinction, but that may require special attention or protection

to prevent them from becoming at risk.

The study area has vast areas of natural and semi-natural habitats that will sustain robust butterfly

populations into the foreseeable future. The habitats and hence species most likely to be adversely affected

by increasing human population are the low elevation species. The supply and quality of low elevation

habitats will likely continue to decrease through continued urbanization and agriculture. Forestry is

practiced on a sustainable harvest basis and is not believed to be a significant factor in butterfly abundance

at the landscape scale. A greater impact on butterfly populations in predominantly forested areas is that of

grassland loss by forest encroachment and loss of host plants in open canopy forests through forest ingrowth

and resultant shading out of host plants that cannot survive under a continuous tree canopy.

ACKNOWLEDGEMENTS

Identification of the north Okanagan material was confirmed in part by: Crispin S. Guppy, Norbert

G. Kondla and Jon H. Shepard. Guppy and Kondla shared their collection information for this area and

Kondla also searched databases for museum records and the literature for relevant papers. Creation of the

digital file of this paper was begun by P. Jones, furthered by Guppy and completed by Kondla. Kondla also

digitized the photos used in this paper and produced the map. Guppy and Kondla also provided peer review

of draft manuscripts. Masthead photos of the C. affinis and C. sheridanii are by Kondla. This project

would have been impossible without their help, expertise and encouragement. It was greatly appreciated.

Special thanks go to the late Robert P. Nelson of Vernon and the late James Grant of Lavington. Both

of these knowledgeable men showed me a number of collecting localities and generously provided records

of several species they had collected in the north Okanagan.

I acknowledge the assistance of the following for their participation in the project: Ms. Terry Hurst,

Vernon; Bob McVickar, Sicamous; Ron Candy, Curator of the City of Vernon Museum; Gordon Bloom and

Peter Mayfield, volunteers at the Museum; and Leonard Threatful, Revelstoke.

21

REFERENCES

BALINT, Z. AND K. JOHNSON. 1997. Reformation of the Polyommatus section with a taxonomic and

biogeographic overview. Neue Entomologische Nachrichten 40:1-68.

BIRD, C.D., G.J. HILCHIE, N.G. KONDLA, E.M. PIKE, AND F.A.H. SPERLING. 1995. Alberta Butterflies.

Edmonton: The Provincial Museum of Alberta. 349 pp.

BLACKMORE, E.H. 1920. The Lycaeninae of British Columbia. Proceedings of the Entomological Society of British

Columbia 14:5-11.

_. 1921. The genus Argynnis in British Columbia. Proceedings of the Entomological Society of

British Columbia 16:27-31.

BUCKELL, E.R. 1947. A list of the Lepidoptera collected in the Shuswap Lake district of British Columbia by Dr.

W.R. Buckell. Proceedings of the Entomological Society of British Columbia 43:11-21.

CLARK, A.H. 1941. Notes on the American respresentatives of the butterfly genus Argynnis. Journal of the

Washington Academy of Sciences 31(9):381-384.

DOWNES, W. 1918. Notes on the Lepidoptera of the northern Okanagan. Proceedings of the Entomological Society

of British Columbia 10:11-13.

DYAR, H.G. 1904. The Lepidoptera of the Kootenai district of British Columbia. Proceedings of the United States

National Museum 27(1376):779-938.

FARLEY, A.L. 1979. Atlas of British Columbia. UBC Press. 136pp.

FISCHER, A.I., J.H. SHEPARD AND C.S. GUPPY. 2000. Butterflies and moths of the Chilcotin district, British

Columbia. Report for Ministry of Environment, Lands and Parks; Government of British Columbia. 66 pp.

GUPPY, C.S. 1986. The adaptive significance of alpine melanism in the butterfly Parnassius phoebus F.

(Lepidoptera: Papilionidae). Oecologia 70(2):205-213.

GUPPY, C.S. AND N.G. KONDLA. 2000. Status of the butterflies and skippers of British Columbia for the National

Accord for the Protection of Species at Risk. Prepared for Conservation Data Centre, Ministry of Environment,

Lands and Parks. 87 pp. + MS Excell Spreadsheet.

GUPPY, C.S., J.H. SHEPARD AND N.G. KONDLA. 1994. Butterflies and skippers of conservation concern in

British Columbia. Canadian Field-Naturalist 108(1): 31-40.

GUPPY, C.S., AND J.H. SHEPARD. 2001. Butterflies of British Columbia. Vancouver, BC: UBC Press 414pp.

HARDING, L.E., AND E MCCULLUM. 1994. Overview of Ecosystem Diversity. Chpt. 18. Pp.227-243. In:

Biodiversity in British Columbia: Our Changing Environment. Eds.: Harding, L.E., and E. McCullum.

Environment Canada, Canadian Wildlife Service. Ottawa. Ministry of Supply and Services. 426pp.

HARDY, G.A. 1954. [Rhopalocera]. Pages B51-B52 in The Natural History of the Forbidden Plateau Area,

Vancouver Island, British Columbia. Report Provincial Museum Natural History and Anthropology (British

Columbia) 1954:B24-B63.

JOHNSON, K. 1992. The palearctic “elfin” butterflies (Lycaenidae, Theclinae). Neue Entomologische Nachrichten

29:1-141.

KOHLER, S. 1977. Revision of North American Boloria selene (Nymphalidae) with description of a new subspecies.

Journal of the Lepidopterists’ Society 31(4):243-268.

KONDLA, N.G. 1999. Pend-d’Oreille Butterfly Survey. Living Landscapes Program report for Royal BC Museum

and Columbia Basin Trust. 38 pp. ( http://www.livingbasin.com/cbasin/butterflv/index.html )

_. 2001. Clarification of and comments on northern Speyeria hydaspe subspecies

(Lepidoptera:Nymphalidae). Taxonomic Report 3(1): 1-5.

KONDLA, N.G. AND C.S. GUPPY. 2002. Nomenclatural correctness of Phyciodes pratensis vs Phyciodes

pulchellus (Lepidoptera: Nymphalidae). Journal of the Lepidopterists’ Society 56:171-172.

KONDLA, N.G., E.M. PIKE AND F.A.H. SPERLING. 1994. Butterflies of the Peace River region of Alberta and

British Columbia. Blue Jay 52:71-90.

KONDLA, N.G., C.S. GUPPY AND J.H. SHEPARD. 2000. Butterflies of conservation interest in Alberta, British

Columbia, and Yukon. Pp. 95-100 in Darling, L.M. (ed.). Proceedings of a Conference on the Biology and

Management of Species and Habitats at Risk. Volume 1. BC Ministry of Environment, Lands and Parks and

University College of the Caribou. 490 pp. (http://owlnut.rr.ualberta.ca/~barb/butsatrisk3.html )

22

23

Figure 4. Kalamalka Lake Provincial Park.

25

21(l):24-26.

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

Catch us on the web at: www.tils-ttr.org

27

Volume 4

1 April 2003

Number 2

Erratum. 5 July 2003

"Butterflies of the North Okanagan, British Columbia, Canada"

by David L. Threatful

The International Lepidoptera Survey

The Taxonomic Report

Page 14, paragraph 2: Anna’s Blue - Plebujus anna

Flight period.

Was: mid-July through August.

Should be: Mid-July into mid-September

Page 15, paragraph 1: Arctic Blue - Agriades rusticus megalo

Flight period.

Was: mid-June through early September, depending on elevation.

Should be: mid-June through July in the North Okanagan.

Page 15, paragraph 4: Zephyr Anglewing - Polygonia gracilis zephyrus

Records.

Was: Silver Star Mtn. 1150... Ellison Ridge 600m.

Should be: Westside of Kalamalka Lake 400m.

Page 19, paragraph 4: Anicia Checker-spot - Euphydryas anicia

Habitat.

Was: ... It is absent from the su m mits of ...

Should be: ... It appears to be absent from the summits of ...

Flight period.

Was: It flies from late April to early September depending on elevation.

Should be: It flies from late April into mid-June depending on elevation in the North

Okanagan.

Volume 4

5 July 2003

Number 3

The Taxonomic Report W

A SUBSPECIFIC ASSESSMENT OF THE GENUS HESPERIA (HESPERIINAE)

IN EASTERN NORTH AMERICA

(PART I: THE SOUTH)

New Subspecies of Hesperia metea, Hesperia sassacus and Hesperia meskei

RONALD R. GATRELLE 1

126 Wells road, Goose Creek, South Carolina 29445

with sections by

ALEX GRKOVICH 1

4 Valley Circle, Peabody, Massachusetts, 01960

MARC C. MINNO

600 NW 35 th Terrace, Gainesville, Florida 32607

ABSTRACT. Hesperia meskeipinocayo Gatrelle and Minno is described from Big Pine Key, Monroe Co., Florida.

This subspecies is thought to be nearly extinct. Its mainland status is uncertain. It is characterized by bi-colored ventral

hindwings with more prominent ventral hindwing spots. Hesperia metea intermedia Gatrelle is described from Pickens Co.,

South Carolina. Its range is from northwestern South Carolina west through Mississippi and probably into Missouri. Males

have the least dorsal fulvous of the metea subspecies; ventrally they are well marked. Females are similar to subspecies

licinus. Hesperia sassacus nantahala Gatrelle and Grkovich is described from Clay Co., North Carolina. It is often boldly

spotted on the ventral hind wing especially in females. The ventral ground color is often dark mottled brown in both sexes

rather than yellowish as in the nominate subspecies. Its known range is the southern Appalachian mountains of western North

Carolina, but likely extends from northern Georgia to West Virginia.

INTRODUCTION

The southern United States is an historically under studied region of the country relative to its

butterflies and skippers. Only Florida has received adequate attention from colonial times to the present.

Yet, even in Florida, there are still a few taxa in need of further research. One such species is Hesperia

meskei (W.H. Edwards, 1877). The populations of this skipper on the lower Florida Keys, Monroe County,

have long been suggested by various writers and researchers as a likely undescribed subspecies. Yet, it has

remained undescribed until this publication.

North of Florida, the deep South from Louisiana to North Carolina has had few resident

lepidopterists. The few who have lived in this area and built notable collections have not been working

butterfly taxonomists. Further, only a limited number of areas within this region have been well collected

and much of the region is still unsurveyed. Even in some well collected areas, little taxonomic scrutiny has

been locally applied. One well collected but under studied region is the area around Clemson University in

northwestern South Carolina. For decades it has been known that Hesperia metea Scudder, 1864 is an

inhabitant of this area. This taxon is examined in this paper and determined to be an undescribed

subspecies ranging from South Carolina through Mississippi and probably into Missouri.

1 Staff Researcher, The International Lepidoptera Survey, Goose Creek, South Carolina.

Perhaps the least studied area in the South is western North Carolina. This seems unusual as much

historical collecting was done in the Smoky Mountains National Park and also in the vicinity of the

Biological Station, Highlands, North Carolina. In the 1990s, the senior author began to survey the

butterflies and skippers of western North Carolina in a wider and thorough manner via contract work he

preformed for the US Forest Service and The International Lepidoptera Survey (TILS). Gatrelle has

previously described two taxa new to science from this region. A third is now added as Hesperia sassacus

Harris, 1862 has been found to occur widely in this region and to constitute an undescribed subspecies.

The description herein of three new Hesperia subspecies brings the total number of known Hesperia

taxa in the southern United States to eight. Ibur of these have been described just in the last decade, and

thus, the number of described Hesperia taxa in this region doubled at the turn of the twenty first century.

This serves to illustrate how taxonomically understudied the South has been. Other undescribed taxa are

presently known from the southern United States at both the subspecies and species level in various

families. Historically, the South seems to have been viewed as faunally ubiquitous - especially by those in

other regions. But the region is composed of many varied faunal zones and specialized niches produced by

its rich geologic and climatic history.

All: left side, dorsal; right side, ventral. Figs. 1-2: d holotype & 9 allotype, Hesperia meskei pinocayo (data in text). Figs.

3-4: d & 9 , H. metea licinus, 29 March 1962, Tyler S.P., Smith Co., TX. Fig. 5: H. m. meskei, 1 June 1975, Freestone

Co., TX. Fig. 6: d, H m. straton, 3 October 1995, Horry Co., SC. Figs. 7-8: d holotype & 9 allotype, H. metea intermedia

(data in text). Figs. 9-10: d & 9 topotypes, H. sassacus sassacus, 6 June 1975, Sherbom, MA. Figs. 11-12: d & 9

topotypes, H. m. metea, 17 May 1955, Willimantic, CT. Fig. 13: 9, H. m. metea, 29 April 2003, Harnett Co., NC. Figs. 14-

15: d holotype & 9 allotype, H. sassacus nantahala (data in text). Figs. 16 & 18: d & 9 paratypes, H. s. nantahala, 27 May

2003, Clay Co., NC. Fig. 17: d,Atalopedes campestris Huron (Same data as 16). Scale: 1.3 actual size. Photos: Joe Mueller.

Hesperia sassacus

Gatrelle and Grkovich

Until the 1990s, Hesperia sassacus was considered a rarity south of Virginia (Harris 1972, Howe

1975, Pyle 1981, Opler & Krizek 1984). But with the recent advent and boom in butterfly watching,

several colonies of this skipper have now been found in western North Carolina in Ashe, Watauga, Avery,

Mitchell, Burke, Yancey, Madison, Haywood, Macon, and Clay counties (Harry LeGrand pers. comm.)

Thus, it is now considered a not uncommon taxon in western North Carolina.

Sassacus has yet to be recorded from Georgia, but it is surely there as North Carolina specimens

have been collected within one mile of the Georgia line (Scaly area in Macon, Co. by James Adams). The

northeastern Georgia counties should be examined closely for the presence of this species. The only

historical Georgia record is attributed to the coastal region (Harris 1972), but this is certainly an

identification error. Gatrelle has located two highly probable sites but has not been able to survey these at

the proper time of year. They are the Popcorn Overlook serpentine barrens on Hwy. 76 east of Clayton and

the area around Sky Valley off Hwy. 246, both in Rabun County.

It has been known since the early 1940s that individuals of the then southernmost populations in

Virginia were often quite dark and similar to the Canadian “form manitoboides ” (A.W. Lindsey 1942,

Clark & Clark 1951, Forbes 1960). Forbes even referred to the populations in the mountains of western

Virginia as, “a richly colored race, approaching manitoboides .” Forbes used the term race for subspecies.

Thus, our current presentation of the southernmost populations as a dark new subspecies can not be called a

“new discovery.” We are merely certifying that which has previously been suggested by others - that the

southern mountain populations of sassacus are phenotypically and subspecifically distinct from nominate

sassacus of the New England region.

The authors are uniquely experienced with this species in the field and are thus positioned to make

an accurate subspecific assessment of this taxon. Gatrelle is familiar with the populations at the extreme

southern end of its range and has collected the type series. Grkovich resides in the type locality of nominate

sassacus and is thus familiar with the variation of the nominate taxon. He is also familiar with the

Appalachian subspecies via his observations and collecting of it in southeastern West Virginia.

Hesperia sassacus nantahala Gatrelle and Grkovich, New Subspecies

Diagnosis and description. The nantahala photos (Figs. 14-16 & 18) well illustrate the distinctiveness of this

new subspecies from nominate sassacus (Figs. 9-10). Dorsally: males of the two subspecies are patterned much alike but

nantahala is a slightly deeper orange and has the fulvous areas slightly more restricted; in females, the fulvous is much more

restricted in nantahala with 23% of the paratypes being form pocahontas (Fig. 18), 46% form alfaratta and only 31%

moderately to fully fulvous as is usual in nominate sassacus females. Ventrally: topotypical sassacus males are almost

always light yellow fulvous on the hind wing with the hesperia spot band only slightly or moderately different from the

ground color, few specimens have well defined spots on a darker ground; in nantahala males, the ventral hind wing ground is

quite variable, often dark brown (Fig. 16) and usually medium brown but occasionally light orange yellow (but not as light as

in typical sassacus ), against this dark ground the hesperia spot band is broad and more defined than in sassacus ; in nantahala

females, 95% of females have medium to rich brown hind wing ground with the cream colored hesperia spot band

pronounced, so much so that some female nantahala can easily be mistaken for the species Hesperia leonardus Harris,

1862.

Types. Holotype d (Fig. 14): NORTH CAROLINA: Clay County, Hwy. 64 at Buck Creek, 3400 ft., 27 May 2003.

Allotype 9 (Fig. 15): NORTH CAROLINA: Clay County, Buck Creek, For. Rd. 6269, 27 May 2003. Paratypes : 23 dd,

13 9 9: All NORTH CAROLINA: Clay County: various sites in and around Buck Creek: 1 d, 16 May 2002; 4 d d, 1 9,24 May

2002; Id, 19, 25 May 2002: Id, 10 May 2003; 13 d d, 9 9 9,27 May 2003; 2d d, 1 9,2 June 2003. Macon County: Jones

Knob area, 4000 ft.: 1 9, 25 May 2002; 1 d, 27 May 2003. All leg. R. Gatrelle. The holotype and all paratypes are currently

in the TILS Museum of the Hemispheres’ collection, Goose Creek, South Carolina. Some paratypes will eventually be

deposited in other institutions. (Dark paratypes, 27 May 03, Clay Co., figures 16 & 18.)

Etymology. Nantahala is taken directly from Nantahala National Forest, western NC and is used as a noun.

3

Remarks. Based on specimens Grkovich has observed and collected in Greenbrier Co., West Virginia, we consider

the range of nantahala to extend north into southern West Virginia. The region and range of intermediate populations north

and east of southern West Virginia needs to be determined. The colonies of nantahala in the southern Appalachians are

numerous but generally isolated from each other. As with all southeastern Hesperia (MacNeill 1964), this is an old taxon

and thus its phenotype is genetically stable within these micro colonies which have doubtless been isolated from each other

for hundreds to thousands of years. This taxon is found in open meadows, even highway right-of-ways (type locality). But it

is also resident deep in open hardwood forest along trails in the same environs as Poanes hobomok monofacies Gatrelle,

2002 and Phyciodes batesii maconensis Gatrelle, 1998. Nantahala is a fairly large Hesperia with an average expanse from

outer fore wing margin to margin in spread males of 30mm and in females 34mm (type series). In Massachusetts sassacus,

the average was 26mm in males and 30mm in females (n=10 each). We have not observed oviposition. Many specimens of

both sexes of nantahala strongly resemble Atalopedes campestris (Boisduval, 1852) (Sachem Skipper) on the ventral

surface (Fig. 17). This resemblance is to such a degree in many individuals that a positive sight determination between the

two from this surface alone is not possible. A series of eleven sassacus from Grand Isle, Vermont were dark beneath as in

nantahala, but above they were typical sassacus. These types of individuals should be referred to as form manitoboides and

may be frequently encountered in eastern Canada and the adjacent areas of the United States.

Hesperia metea

Gatrelle

The known southern colonies of Hesperia metea Scudder, 1864 are moderate in number. Most

southern specimens have been collected in Mississippi and Georgia. In North Carolina, it has now been

documented in widely scattered colonies in 17 counties by the many butterfly watchers in that state with

most records centered in the south central Sandhills region. Unfortunately, vouchers are lacking for most of

these sites. I am aware of only one regional site for this skipper in South Carolina even though it is

recorded for five counties. The records for Pickens and Oconee counties are confirmed. There are several

scattered colonies in that region in and around Clemson.

There is very little taxonomic information in the literature concerning the deep South populations of

metea. Even in Harris’ Butterflies of Georgia , most of his text is devoted to accounts of the species in

Pennsylvania and New Jersey. Opler & Krizek 1984 only state, “Populations in Arkansas and Texas

(subspecies licinus) are almost immaculate above and below, and some individuals elsewhere in the Deep

South are nearly so.” C. Don MacNeill in Howe 1975 provides the most, and most pertinent, information

on southern metea. Re the subspecies licinus (Edwards, 1871) he states, “This subspecies is known only

from Texas and Arkansas. Specimens from Georgia and Mississippi, however, show tendencies toward

this subspecies and a careful analysis might well demonstrate a large zone of intergradation between the

two east and north of Texas.”

Until two years ago, I considered the southern populations of metea to be subspecies licinus and

ranging from Texas to South Carolina. There is no doubt that the southern populations, as alluded to by the

above authors, are not the nominate subspecies and that in many ways they most resemble subspecies

licinus. However, the phenotype of the populations from Missouri to South Carolina does not indicate a

“blend zone”. Rather, the populations over this broad area are homogenous in appearance and intra-region

variation. I now hold that this is a unique subspecies and here propose it as such. I have chosen the name

intermedia not because it blends into the other subspecies, but because its characters fall between them.

Evolutionally, I consider this “new” subspecies to be the taxon closest to the parent of both metea

metea and metea licinus. As the southeastern segregate, it is certainly older than metea metea as much of

that subspecies’ range lies in areas that were glaciated a mere 20,000 years ago. At some time, intermedia¬

like parents are hypothesized to have been isolated in Texas or Mexico giving rise to licinus. In modem

times, the ranges of these three subspecies have expanded and moved closer to each other, but I hypothesize

that they have been isolated evolutionally from each other for some time. There is evidence of this at the

east and west ends of the range of intermedia.

In the east, nominate metea’s range extends south into the southern Sandhills of North Carolina (Fig.

13) and perhaps into the northern part of that region in South Carolina. It is a relatively short distance to

upstate South Carolina where intermedia occurs. There are no known “blend zone” (actually, post glacial

suture zone) colonies between North Carolina metea and South Carolina intermedia. In the west, the same

situation exists with a relatively short distance between Missouri intermedia and Arkansas licinus with no

known “blend zone” colonies between the two. The area to check in the west would be Remington’s (1968)

Louisiana-East Texas Suture Zone F.

While metea metea extends south into the midlands of North Carolina, intermedia extends north into

the Mountains of North Carolina and perhaps, West Virginia (Allen 1997, figures) and southern Ohio (Iftner

et al 1992, figures). Metea metea has a narrow east west range from New England into Wisconsin. True

metea is a light colored taxon with bold VHW spots and veins. In describing this new subspecies, the range

of metea is much reduced by defining its phenotype more specifically. I have not seen Kentucky or

Tennessee specimens, but I expect them to all fall within the newly recognized subspecies.

MacNeill (1964) states that the genus Hesperia is oldest in the southeastern US. It would seem odd

indeed for the oldest taxon within metea to not be recognized as a distinct subspecies.

Hesperia metea intermedia Gatrelle, New Subspecies

Diagnosis and description. The intermedia photos (Figs. 7-8) well illustrate this new subspecies. Dorsally:

males of intermedia are darker than males of either metea or licinus with males of the latter two being much alike. Females

of intermedia are dark blackish brown as in licinus but with (restricted) light forewing spots; females of metea are usually

bright fulvous brown with bold white spotting. Ventrally: males of topotypical metea have the hind wing hesperia spot band

well developed and often with white “cobweb” veining; in licinus the wins are not white and the spots greatly reduced; in

intermedia males the spots are usually well developed but very little white veining. Females of intermedia have a small but

bright hesperia spot band and no white veining w hil e in licinus the spots are often absent; in metea females they are very well

developed and often with extensive white veining.

Types. Holotype d* (Fig. 7): SOUTH CAROLINA: Pickens County, vise. Lake Issaqueena, 20 May 1993. Allotype

9 (Fig. 8): SOUTH CAROLINA: Pickens County, vise Lake Issaqueena, 21 April 1993. Paratypes: 33dd, 69 9: SOUTH

CAROLINA: Pickens County: three sites vicinity Lake Issaqueena: 9dd, 19, 20 April 1993; 3dd, 19, 21 April 1993:

3d d, 29 9, 21 April 1994; 3d d, 15 April 2003. (All leg R. Gatrelle). MISSISSIPPI: Lee County: Tombigbee State Park:

1 d, 13 April 1975; 2 d d, 7 April 1977; Tishomingo County: 1 9, Mt. Woodall; 1 9,5 mi. w. Belmont; 1 d, Burnsville, all 16

April 1994; lidd, 2 mi. sw Bloddy Springs, 18 April 2002. (All leg. R. Patterson). The holotype and all South Carolina

paratypes are currently in the TILS Museum of the Hemispheres’ collection, Goose Creek, South Carolina. All Mississippi

paratypes in personal collection of R. Patterson. Some paratypes will be deposited in other institutions.

Etymology. Intermedia indicates that this taxon is phenotypically between.

Remarks. In this study, specimens of H. metea licinus were examined from: ARKANSAS: Carroll Co. (7d d, 1 9);

OKLAHOMA: Cleveland Co. (1 d, 6 9 9 ) (det. McGuire & Freeman); TEXAS: Leon Co. (1 d, 1 9); Smith Co. (3 d d, 2 9 9);

Tarrant Co. (Id ); Wise Co. (Id, 29 9). 4 pair were examined from Benton Co., MISSOURI and determined to be

intermedia. This area is on the Ozark Plateau which has faunal affinity with the southern Appalachian region. Only 16

specimens of intermedia were examined from GEORGIA, but these were from a wide area and provided a good sample. GA

locations were: Greene Co., Cobb Co., Bartow Co., McDuffie Co., Douglas Co., and DeKalb area. The Georgia material was

more variable than the South Carolina or Mississippi series but all individuals fell within the concept of intermedia. 52

specimens of nominate metea were examined from MASSACHUSETTS, CONNECTICUT, NEW JERSEY, RHODE ISLAND,

PENNSYLVANIA, and MICHIGAN. All specimens examined were in the Florida State Collection of Arthropods,

Gainesville, Florida or the private collection of the author. The holotype and allotype specimens of intermedia were

selected because they were “average”. There are individuals of both sexes that are both lighter and darker.

A major question was the southeastern limi t of metea metea’s range. No North Carolina Sandhills specimens were

located for loan, and while photos are available, they are risky for taxonomic assessment as they can be skewed by camera

settings, lighting, or electronically “enhanced” on internet web sites. Thus, I made a late trip in 2003 to the North Carolina

southern midlands to locate and examine individuals in the field and collect any as necessary. I located two males and got a

good look at them before they eluded me. Two females were collected (Fig. 13). This sample was tiny, but nonetheless

revealed that they are best referred to subspecies metea. Their phenotype differed from New England individuals (and are

larger,) but I do not feel they can be called “transitional” to intermedia. I certainly do not include them in intermedia.

Hesperia meskei

Gatrelle and Minno

The populations of Hesperia meskei on the lower Florida Keys already have a common name, the

Rockland Grass Skipper. This attests to its having been well known for many years as a unique entity. One

of the most recent treatments of this segregate was by Minno and Emmel (1993) in Butterflies of the

Florida Keys. In the species account, they state that the Keys population seems to represent an unnamed

subspecies. Why this segregate has never been described is not fully known, however, one reason is that

various lepidopterists have talked of doing so over the last 40 years and others held off from describing it

out of professional courtesy. But none have done so. Now, not only have the decades passed by, but this

taxon has nearly passed into extinction. The authors believe it is imperative that this unique and now very

rare insect receive formal status as a taxonomic taxon to better assure that it is afforded any and all

protection that it may require. This official status can also help in promoting searches for it on the mainland

and reintroduction efforts if needed.

The Keys subspecies is very different in all characters from nominate meskei which is endemic to

Texas. True meskei is a very light and bright taxon (Fig. 5). The Rockland Grass Skipper is closest to

subspecies straton (W.H. Edwards, 1881) (Fig. 6) which is the subspecies of the southeastern United States

including mainland Florida where its type locality is Indian River, Florida. All of the specimens of straton

we have examined exhibit a fairly consistent phenotype over its rather large range. Clearly, pinocayo has

evolved a unique phenotype. But more significantly, this subspecies is biologically unique as it does not

diapause but is continually brooded. It has not been recorded in February, July or August but this reflects

larval development (broods) not true diapause (which is usually induced by day length).

Hesperia meskei pinocayo Gatrelle and Minno, New Subspecies

Diagnosis and description. Figures 1-2 well illustrate this new subspecies. Dorsally: males of pinocayo vary

from fulvous as in males of straton and meskei to much less fulvous. Females of pinocayo, straton and meskei are much

alike dorsally. Ventrally: males and females of pinocayo are unique in the color and pattern of the ground and hesperia spot

band of the hind wings; from CU2 to the inner margin the ground is dark ocher to rust brown and from CU2 to the anal margin

(vanal area) light rusty yellow; the hesperia spots are more prominent than in straton and meskei meskei, which in the latter

are usually not visible being the same color as the bright ground; meskei meskei is unique in that the VFW is usually heavily

light fulvous (Fig. 5) while in straton (Fig.6) and pinocayo it is dark to varying degrees.

Types. Holotype d* (Fig. 1): FLORIDA: Monroe County, Big Pine Key, 22 November 1979, leg. W.L. Adair.

Allotype 9 (Fig. 2): FLORIDA: Big Pine Key, 22 April 1973, leg. R.A. Anderson. Paratypes : 20 c? d, 14 9 9: All FLORIDA:

Dade Co.: Id, 19, Navy Wells Preserve, 26 August 1980, leg L.C. Koehn. Monroe Co., Big Pine Key: Id, 26 November

1978, Id, 15 May 1979, leg C.M. Stevens; 19, 11 March 1984, leg. M.C. Minno; 6dd, 19, 20 March 1979, 19, 6

November 1981, leg. R.W. Boscoe; 39 9-, 10 December 1972; Id, 28 January, 2dd, 11 March, 19, 15 March, 3dd, 1 9,

18 March, Id,29 9,25 March, 19,5 April, Id, 22 April, 1 d, 13 May, 1 d, 16 September, all 1973, all leg. R.A. Anderson;

1 d,l 9, 24 November 1979, leg. Jeff Slotten; 1 9, 24 May 1981, leg. M.G. Douglas. The holotype and 10 paratypes are in

the FSCA, Gainesville, Florida. The allotype and 7 paratypes are in the TILS collection, Goose Creek, South Carolina.

Remaining paratypes as follows: Min no collection: 1; Koehn collection: 2; Slotten collection: 2; Anderson collection: 12.

Etymology. Pinocayo is derived from Spanish and is used by us as a noun meaning Pine Key.

Remarks. More research is needed to determine the biogeographic evolution of pinocayo. We do not expect this

taxon to be found to be a species, but acknowledge this possibility. DNA and breeding studies may reveal sufficient distance

and/or incompatibility of a degree that such a status is warranted. Minno has examined the genitalia and found it very similar,

if not the same as, subspecies straton. Our greatest question is the presence and range of this species on the mainland. The

specimens collected by Koehn in Dade County at Navy Wells means it may be (or once was) found in the Miami area.

Gatrelle has often wondered why “ meskei ” has not been found in southwestern Florida in the area just west of the

Fakahatchee Strand as the habitat looks perfect. Conservation is of immediate concern. The taxon is within the Key Deer

Refuge on Big Pine Key and is therefore already afforded habitat and disturbance protection. However, the number of

individuals have now declined so greatly that some type of breeding program seems called for soon if not immediately.

6

ACKNOWLEDGEMENTS

We thank those who loaned us essential specimens of the various taxa in these studies: Andy Anderson (FL),

Jeff Slotten (FL), Leroy Koehn (KY), David Hoag (Vermont), Dale Clark (Texas). We thank the DPEFSCA

Gainesville, FL and TILS/MOTH Goose Creek, SC for access to their diverse collections. We thank Joe Mueller for

his fine photography of specimens.

LITERATURE CITED

ALLEN, T.J. 1997. The Butterflies of West Virginia and Their Caterpillars. Pittsburgh, PA: University of Pittsburgh

Press xii + 388 pp.

CLARK, A.H., and L.F. CLARK. 1951. The Butterflies of Virginia. Smithsonian Misc. Coll. 116 (7): vii + 239 pp.

FORBES, W.T.M. 1960. Lepidoptera of New York and Neighboring States, part 4. Cornell Univ. Agr. Exp. Sta.

Memoir 371, 188 pp.

HARRIS, L., Jr. 1972. Butterflies of Georgia. Univ. of Okla. press, Norman OK. 326 pp.

HOWE, W.H. 1975. The Butterflies of North America. Doubleday & Co., Inc. New York, NY. 633 pp.

IFTNER, D.C., J.H. SHUEY & J.V. CALHOUN. 1992. Butterflies and Skippers of Ohio. Ohio St. Univ. (in co-op

with) Ohio Dept, of Nat. Res. Div. of Wildlife and Ohio Lepid. Columbus, Oh. 214 pp.

LINDSEY, A.W. 1942. A Preliminary Revision of Hesperia. Denison Univ. Bull. Jour. Sci. Laboratories, Vol. 37,

pp. 1-50, 6 pis.

MacNEILL, C.D. 1964. The Skippers of the Genus Hesperia in Western North America, With Special Reference to

California. Univ. of Cal. Press, Berkeley/Los Angeles, CA. 230 pp.

_. 1975 in: HOWE, W.H. The Butterflies of North America. Doubleday & Co., Inc. New York, NY.

pg. 475.

MINNO, M.C. & T.C. EMMEL. 1993. Butterflies of the Florida Keys. Dept, of Zoology, Univ. of FL, Mariposa

Press. Gainesville, FL 168 pp.

OPLER, P.A. & G.O. KRIZEK. 1984. Butterflies East of the Great Plains, An Illustrated Natural History. Johns

Hopkins Univ. Press, Baltimore, MD. 294 pp.

PYLE, R.M. 1981 (1992). The Audubon Society Field Guide to North American Butterflies. Alfred A. Knopf, NY.

pp. 924.

REMINGTON, C.L. 1968. Suture-zones of Hybrid Interaction Between Recently Joined Biotas. Evol. Biology, Vol.

2 (8), 325-413.

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

Donations are needed to support and further our efforts to discover and protect butterflies worldwide. All

donations are US tax deductible. Please help generously.

Donations should be mailed to: Treasurer, 126 Wells Rd., Goose Creek, SC 29445.

Checks should be made payable to: TILS. Please indicate if you need an individual receipt.

Catch us on the web at: www.tils-ttr.org

Volume 4

15 November 2003

Number 4

A TAXONOMIC REVIEW OF CHLOSYNEISMERIA

WITH DESCRIPTION OF A NEW ISMERIA SUBSPECIES FROM

THE SOUTHERN APPALACHIAN MOUNTAINS

RONALD R. GATRELLE h 2

126 Wells Road, Goose Creek, South Carolina 29445-3413

ABSTRACT. The taxon Boisduval and Le Conte described in 1833 as Melitaea ismeria is examined as a historical

entity and confirmed as a sister subspecies of the taxon Chlosyne ismeria nycteis. The ismeria populations in the southern

Appalachian mountains are described as a new subspecies: Chlosyne ismeria obsoleta. In facies, this new subspecies is most

similar to the western subspecies Chlosyne ismeria drusius in that it is dark and contrasting in its dorsal markings. It is

known to range from the mountain region of extreme northern Georgia north in the Appalachians to near southwestern

Virginia. It may range into southern West Virginia.

Additional key words: Appalachian peninsula, Ohio, nycteis type locality.

DETERMINING MELITAEA ISMERIA

HISTORICAL OVERVIEW

In 1833, Boisduval and Le Conte described Melitaea ismeria in a detailed text supplemented by an

accompanying illustration based on a small 10 Vix b^painting by John Abbot. The illustrations of the

ismeria imago in Boisduval and Le Conte have always been assumed, since publication, to be poor copies

of the Abbot originals. This was assumed because the whereabouts of the original had been unknown to

taxonomists until very recently (Cowan 1969, Calhoun 2003). Yet, from 1833 to today, these illustrations

have been the focal point of all researchers in trying to determine the zoological identity of this species

while the detailed written description has been virtually ignored.

Brown (1974) established what the taxon ismeria was not (not gorgone male Hubner, 1810), and

Gatrelle (1998) established what it was (a conspecific with nycteis Doubleday, [1847]). However,

Calhoun (2003) revisited this name and contradicted the conclusions of both Brown and Gatrelle, and

returned this whole matter to instability. Calhoun did not present or examine the original description of

ismeria nor did he give or weigh any information on the nycteis population at the ismeria type locality that

Gatrelle (1998) determined and established by neotypification to be Boisduval and Le Conte’s ismeria. It is

thus essential to herein present and weigh all evidence pertinent to ismeria’s taxonomic determination.

Since the Abbot paintings have been the historical focal point, it is appropriate to begin with an examination

of these.

1 Staff Researcher, The International Lepidoptera Survey, Goose Creek, South Carolina

2 Research Associate, Florida State Collection of Arthropods, Gainesville, Florida.

The Abbot original was among a folio of 148 paintings of the above small size format (Cowan,

1969). Calhoun (2003) states that the folio actually contains 149 paintings, of which 105 are attributable to

Abbot. These paintings have a history of being “lost” and then “found”. This history is detailed in several

publications and I will not repeat it again here. The best references are Cowan (1969), Brown (1974) and

Calhoun (2003). Today, these paintings reside at the University of South Carolina, Columbia, SC in the

Thomas Cooper Library. They are available for viewing on the Internet at:

http://www.sc.edu/library/spcoll/abbot/default.html.

As stated, the published OD illustrations of the ismeria imago have always been assumed to be poor

copies of their Abbot originals. This assumption existed for two reasons. First, the people of that era with

a serious interest in natural history were accustomed to having lifelike paintings in the finer scientific

books. The Boisduval and Le Conte ismeria illustrations are rather unrealistic looking. Second, the adult

figures (from time of publication to today) don’t precisely match any known species. However, as can now

be seen by comparing the original on the above website with the published copy (Fig. A), the two are

virtually identical. Thus, we now know that both are equally poor, or equally accurate, renditions of the

intended species. This sameness is significant in the taxonomic assessment of the name ismeria. This is

because we now see that the published illustrations are sufficient as they are the same as their original.

In his paper on ismeria , Calhoun (2003) shows three “original” Abbot paintings that are surely

different renditions of the same basic template (his figures 4, 5, & 6). They are: one from the Natural

History Museum, London painted between 1790 / 1816, one from the Turnbull Library, Wellington, New

Zealand painted between 1816/1818, and the Boisduval and Le Conte original of ismeria commissioned in

1813 (but perhaps painted a bit later) and deposited as above at the University of South Carolina.

Two things become immediately apparent when considering these paintings. First, all the adults

figured are similar to, yet unlike, any one specific taxon known from the eastern US. Second, in looking at

them together, the NHM and Turnbull copies look much alike, but the Boisduval and Le Conte original (and

copy) differs in subtle but noticeable elements 3 . Since all of these were painted by Abbot himself, and

apparently all about the same period, the question naturally arises as to why the one from which the

published ismeria plate was duplicated is an obvious departure from the other two. Further, since they are

all from his own hand, it would be total conjecture as to which is the most accurate representation of the

living model. It is just as likely that the original utilized by Boisduval and Le Conte is the most

representative. In fact, it is reasonable speculation that since Abbot knew this rendition was for Le Conte

rather than simple commercial sale, he may have given more attention to accuracy then with the others. It

would be illogical to think he gave less attention to this copy commissioned by and for Le Conte. The point

is that it is completely subjective opinion on anyone’s part as to which version someone might consider to

look more realistic - like the living model.

Abbot, like other artists of his day, utilized master templates from which they produced “original”

copies. It was from such a master template that the above “original Abbot’s” (and doubtless others) were

copied and produced. These master templates, which would have resided with Abbot, are apparently

unknown. What we do know is that the ismeria paintings used by Boisduval and Le Conte (the original and

the published copy) are clearly different from the other two renditions that Abbot painted. Thus, these two

“versions” are the only ones that have technical relativity to the taxonomic taxon ismeria. Brown was the

first to establish this technical point in his 1974 paper.

We thus have two obvious problems with these paintings. 1) All the adults figured are similar to,

yet unlike, any one specific taxon known from the eastern US, and 2) there are subtle but noticeable

differences between the Boisduval and Le Conte ismeria paintings and the others. There is a simple, and I

think obvious, explanation for these artistic discrepancies - they are composites of two taxa.

3 The ismeria original has the legs, abdomen and wing shape altered from Abbot’s master template. This is evidenced by the

significant differences in these areas from Abbot’s earlier paintings, and erased sketch marks (figs. 4 - 6 in Calhoun, 2003).

How this occurred, by whom and why is speculative.

All previous researches have tried to fit the OD adult ismeria illustrations to a single taxon. This

was the logical thing to do as the painting was presented as a single species. As the decades passed and the

fauna of coastal Georgia began to be understood, it was assumed that only one species of Chlosyne

occurred in that region. That species being Chlosyne gorgone (Hiibner, 1810). This assumption eventually

came to be presented as fact. Eventually, various workers just began to refer to the above NHM painting

(and the ismeria plate) as “gorgone” (e.g. Doubleday; see Calhoun, 2003) as if it was a fact that this is what

it represented. This assumption was the primary topic F.M Brown (1974) dealt with and unequivocally

rejected. Brown concluded that the Boisduval and Le Conte plate was neither gorgone nor directly

associated with the NHM Abbot painting which Brown was fully aware of and assessed. Calhoun (2003)

revived the same misidentifications, assumptions and persons addressed by Brown, and throughout his

paper utilizes the name gorgone in application to all of these paintings as if it was fact.

However, it is now known that not one but two species of Chlosyne occur in Burke County, Georgia

(Gatrelle, 1998). The first species of Chlosyne discovered in Burke Co., GA was not gorgone , but the

taxon then known as Chlosyne nycteis (Doubleday, [1847]). This taxon was found there in 1989. Then, in

1993 gorgone was rediscovered there also (Gatrelle, 1998). This discovery of two fairly similar species

of the same genus in Burke county in conjunction with the abstract elements found in all of these original

Abbot paintings dictates that the most logical assessment of these distorted artistic imago renditions is that

they are a composite of these two sympatric Burke County Chlosyne - C. ismeria (=nycteis) and C.

gorgone (Hubner, 1810). This is evidenced by the fact that the adult paintings incorporate characters that

are clearly possessed by one but not the other of these two species. It is also significant that the “nycteis”

that occurs in Burke County is phenotypically (subspecifically) different from northeastern and Midwestern

nycteis. This coastal Georgia and South Carolina subspecies is one that no one had ever seen between its

description in 1833 as ismeria and its rediscovery in 1989.

If indeed these paintings of the adult morphs are to any degree composites of these two Chlosyne

species, then they are unsuited in and of themselves, by their unscientific nature, for use in definitive

taxonomic diagnosis and determination (i.e. lectotypification or diagnostic depictions). Unfortunately, these

painting have been virtually the sole focus of workers down through the last century and an half. If argument

is want to be made to try and establish the true identity of the taxon ismeria solely, or primarily, from these

paintings, then taxonomists would be doomed to a never ending subjective debate over its taxonomic

identity.

In my 1998 paper I dealt very little with any historical aspects and simply referred the reader to

Brown (1974) as I agreed with his historical research completely relative to ismeria not being conspecific

with gorgone. In hindsight, I see I was remiss in not offering more historical information and just moving

ahead with defining ismeria by my field and biological data. Even there, I could have presented much more

information on the immatures from my rearing efforts. I could have provided detailed data on the habitats of

both ismeria and gorgone in Burke County. Since I thought the ismeria = gorgone issue was a dead one

(per Brown), I focused mainly on the basic taxonomic issues and resolving any remaining possible

confusion via the necessary neotypification of both ismeria and nominotypical gorgone from their mutual

type locality. My realization this year that an undescribed subspecies of ismeria exists in the southern

Appalachian Mountains provides the opportunity to revisit these issues, and in timely fashion in light of the

recently published paper by Calhoun.

Fortunately, there is a fairly large body of factual evidence in the original description of ismeria,

the notes of Abbot, the historical works of Cowan and Calhoun, and actual biological and field data by

which we may know with a great deal of certainty to which taxon the epithet ismeria was applied by

Boisduval and Le Conte. The definitive data whereby we may assess which living organism Boisduval,

and especially Le Conte, intended is not found within the various paintings of Abbot - or the altered

original used for the ismeria publication.

3

VERIFIABLE FACTS

Fact one. John Abbot was not a taxonomist and never authored any taxonomic act or described any

species. He was a marketer in specimens, art, and data. He did this for money. The only name Abbot ever

applied to the entity in these paintings was the common name Crop Wort Fritillary.

Fact two. The name gorgone was authored by Hubner in 1810 from specimens probably purchased

from one of Abbot’s European agents. While Hubner provided no verbal description, the paintings Hubner

did of the specimens in his possession are so accurate as to leave no doubt of the taxon he had in hand (Fig.

4). Abbot had nothing to do with this taxonomic act.

Fact three. Boisduval and Le Conte authored ismeria 23 years after Hubner’s gorgone in 1833 and

also from specimens. This is evidenced by the detailed and accurate textual description they alone penned.

(A description that apparently none of the historical researchers of ismeria have ever quoted or even

considered until this present publication.) This description would have been impossible without

specimens, because their verbal description differs definitively from the accompanying Abbot painting in

several diagnostic characters, especially on the ventral hind wing. This description had nothing to do with

Abbot, and he had nothing to do with the taxonomic act of authoring this name.

There are those who will dispute this as a fact. This is because it has commonly been assumed that

Boisduval had no specimens. But it is not a fact that they had none, or had not seen any. It is just an

entrenched assumption. Cowan (1969) stated on page 126: “In that year [1828] J.E. Le Conte went to

Paris with a large collection of North American insects, to study other material and with the idea of

writing a book ” This book became the 1833 volume coauthored with Boisduval in which ismeria was

described verbally and in detail. Calhoun (2003) stated on page 212: “Abbot may have collected natural

history specimens in South Carolina (Sanders & Anderson 1999), but the reference to ‘Carolina’ in the

original description of M. ismeria likely came from J.E. Le Conte, who traveled more widely in the

Southeastern United States [than Abbot]” Le Conte also often visited for periods on the family Plantation

south of Savannah, Georgia.

How would Le Conte know ismeria occurred in South Carolina if he had personally never

encountered it there? Obviously, from his locality citation, he had. As a collector with a scientific interest

to describe new taxa, it would be wild speculation to assume he never collected any of this taxon he

encountered, and if collected, that he would not have taken them among his “large collection” of American

insects to Paris with him. If not fact, it is highly probable Le Conte had specimens, and if not, he certainly

(for a fact) had a memory of what he encountered and meant as ismeria. This was Le Conte’s concept not

Abbot’s. His and Boisduval’s verbal description of the OD underside of ismeria (see below) with basal

white spots on a tawny ground with pupiled marginal tawny semi-circles (precisely as in coastal “nycteis”)

is far different than the almost black and white stripped nominotypical subspecies described by Hubner as

gorgone. It is also preposterous to think that if what Le Conte encountered personally in the Carolinas, and

surely discussed with Abbot in their Georgia visits, was Hubner’s 1810 gorgone, that Le Conte would not

have recognized it as such and/or described it again as a new species in 1833.

I have found ismeria (=nycteis) in both upper coastal Georgia and South Carolina (Fig. 8). Thus

establishing that Le Conte could have indeed found it in my home state of South Carolina. Both of my sites

are near colonial road crossings of the Savannah River and thus may be the same areas Le Conte also found

it. Le Conte mentioned the Carolinas first and then Georgia as the region to which he attributed it. Abbot’s

range notation says he found it only in Burke County. I agree with Calhoun, the attribution of this taxon to

Carolina was Le Conte’s doing. This necessitates he had personal experience with it there and thus: 1)

factually saw it in nature, 2) with high probability collected it and 3) with equally high probability had

specimens among his “large collection of American insects”. This is why he described the ventral

hindwing of ismeria as having basal white spots on a tawny background and marginal tawny half circles

with dark pupils - which is totally unlike the dark brown and basally striped gorgone venter (Figs. 3c & 6).

Fact four. Both ismeria and gorgone in Burke County are found in the same habitats - open

forested areas. Both feed as larvae on the same hosts - Helianthus (Sunflowers). Gorgone is specific to

species divaricatus L. Ismeria likely feeds on a wider variety of Helianthus at their mutual type locality,

but also on divaricatus. I did not find ismeria in Burke County in “swamp” habitat as was inferred by

Calhoun (2003). (He did not communicate with me when doing his paper even though I have ten years of

experience with the ecology and biology of these taxa at the type locality.) When I located ismeria in South

Carolina it was also in dry, open, upland, wooded habitat.

Fact five. Chlosyne ismeria (= nycteis ) is double brooded in coastal Georgia and South Carolina

with the first brood appearing the end of May and June, and the second brood in August and September.

First brood larvae mature in May and emerge the same year in June. It is recorded throughout the historical

literature that the species Abbot reared and painted as reproduced by Boisduval and Le Conte was,

according to Abbot’s own notes, from larvae he found in May and eclosed as an Adult May 26 of the same

year. This fits precisely with the first brood of ismeria ( = nycteis ), and is virtually impossible to be

gorgone as follows.

Fact six. C. gorgone is univoltine in Burke Co., Georgia (its type locality). Adults fly during April

and are absent by mid May and the rest of the year. I have gathered over 800 eggs and first instar larvae of

nominate gorgone from Georgia and South Carolina with about 200 of these being found in several clutches

on Helianthus divaricatus at the type locality. I sent some of these to Tom Allen in Virginia. I instructed

Richard Boscoe of Pennsylvania where to find the Colony in South Carolina so he could rear them. It is

virtually impossible to keep larvae from entering diapause. Both of these individuals and myself are very

experienced at rearing Lepidoptera. Allen had no success breaking diapause and all died (personal

communication). Boscoe only obtained adults by allowing diapause and adults emerged the next spring

(personal communication). Gatrelle was only able to induce 4 of these hundreds of larvae, from several

different females, to not diapause. Of these, only two pupated and one of the two emerging Adults was

deformed. The larvae of these two adults were obtained on 27 April and emerged 7 and 9 June, 1993. On

their ventral hind wings they were both very atypically black with restricted white. Surely a result of forced

and unnatural rearing. Both were females. Larvae were kept under intense continual light. Many were even

nudged for hours after entering the 4 th instar to try and keep them “awake”. Many died at that stage. The

rest were returned to litter in the wild where they were taken from. It would have been virtually

impossible for Abbot to rear same season gorgone even if he did know the “trick” of continual intense light.

He would have had to have had so many (and close) candles or oil lamps that the heat from that kind of light

would have dehydrated the host and the larvae. All of the larvae I gathered from the type locality and from

South Carolina were all black to the 4 th instar. The 4 mature larvae were all black except for a narrow

rusty orange line on the dorsal and a medium gray band along the base of the body. These differ

dramatically from the larvae painted by Abbot. Thus, it was not gorgone Abbot reared.

No one has seen the topotypical larvae of C. ismeria. C. ismeria nycteis, however is known to

have black and yellow striped larvae - just as Abbot figured and as noted by Calhoun page 212, “Grishin

and Boscoe observed that the mature larvae of C. nycteis are black with broad yellow or orange lateral

bands. ” (Calhoun neglected to show any larvae of C. ismeria nycteis or adults of C ismeria ismeria for

analysis. He only presented one sided data and illustrations. He also completely omitted the text of the

original description. Which, as we shall see, is the evidence that supports ismeria not being gorgone and

being a nycteis the most.)

Fact seven. Artistic license is the term we use when an artist adds something to a picture that is not

factual. The following is either a fact or artistic license. In these Abbot paintings which some have

assumed to be “gorgone” and not “ismeria”, they include the mature host plant in full bloom with the adult

male sitting on the blossom of the host. As mentioned above in fact six, gorgone is univoltine and only flies

in April. At that time of year, the host is only a few to 18 inches high. H. divaricatus does not bloom in

that region until late July to September. It is virtually impossible to find a gorgone on the flower of this

plant at the type locality! It is very likely to see ismeria (= nycteis) nectaring on it in its second brood.

Thus, we either have to attribute this “painted situation” to artistic license or the species is not gorgone.

Artistic license can not be allowed into taxonomic matters. Again, these paintings of adults on flowers has

only two options - if Abbot really found this situation in nature (which I think he did), then the only species

can be ismeria (= nycteis). But if this is to be considered a gorgone , then the painted situation is artistic

license - not real. If an artist, in this case Abbot, has a propensity to “doctor up” or “fictionalize” in one

area of a painting, they will utilize the same “license” in other elements of it. He was a dealer/artist after

all and not a scientist. My research at the type locality over 10 years produced results that are entirely

different than the situation literally portrayed by Abbot.

Fact eight. The concepts, assumptions, and identifications of Doubleday, Scudder, and others (as

cited by Calhoun, 2003) are irrelevant to what the actual concept of ismeria was to its authors - Boisduval

and Le Conte. The various non-OD paintings of Abbot are irrelevant as 1) none of them are undisputable

presentations of a single living butterfly species and 2) the OD original differs from them in several

characters. Brown pointed all this out in 1974. Brown also examined and discussed the NHM, London

painting and correctly stated it had no Code relevance to the name ismeria.

Fact nine. Only the Abbot original (in the Thomas Cooper Library) and the OD plate of ismeria

have any ICZN Code relevance to this name. All other Abbot paintings and specimens are outside the

boundaries of what the Code allows relative to the typification of the taxonomic taxon ismeria. This is

because only those specimens and data directly used by, or known to, Boisduval and Le Conte in

formulating the taxonomic taxon Melitaea ismeria are factors upon which the taxon was based (Articles:

72.4.1, 72.5.6, 73.2.1). Now that we know that the “original” painting and that which was produced in the

Boisduval and Le Conte ismeria description are virtually the same (except that the picture in the book is the

most “nycteis” like of them all), it should begin to be considered that the published plate was just as Le

Conte wanted it - conforming most to the specimens he had, or remembered, and that he knew were not

Hubner’s gorgone.

Conclusion. The only logical conclusion is that the Chlosyne Abbot reared was ismeria (=

nycteis). Abbot described these larvae as primarily yellow with black lines and painted them thusly, many

times. All of the early instar larvae of nominate gorgone I found were all black. The four mature larvae

were all black except for a narrow rusty orange line down its dorsal midline and a medium gray band next

to its black legs. No one has ever reared topotypical ismeria - except Abbot. This is the entity he reared

and Boisduval and Le Conte named ismeria. The only scientific description and name ever applied to this

entity Abbot reared and painted, and that Le Conte observed (and likely collected) in the Carolinas is the

name ismeria. Hubner painted and named gorgone. Boisduval and Le Conte textually and explicitly

described and named ismeria. Abbot had absolutely nothing to do with the naming or description of either.

Hypothesis. It is my belief that Abbot found C. gorgone in Burke County. The evidence for this are

the specimens acquired by Hubner that Hubner painted. Abbot also found a “nycteis” in Burke County,

Georgia as evidenced by 1) his painting of what he reared and 2) notes on the timing of its life cycle. It is

highly likely he attempted to rear gorgone and, given the nature of the beast, failed. He then found similar

(esp., in early instars) congeneric larvae on the same plant ( H. divaricatus) (and also on Crop Wort) and

reared it out thinking it was the same species. At this point it would be easy for Abbot to consider the two

“forms” as one species. The greatest proof of this is the Abbot paintings of the adults. Abbot is a fine artist

and all of these paintings have confused workers from the beginning - especially the ventral hind wings,

because, as should be obvious, they are a composite of two taxa he thought were one. The base of the

ventral hind wing looks like gorgone but from the middle to margin just like the “nycteis” in that region.

However, the ventral fore wing is a good depiction of that area’s “nycteis”. The dorsal fore wings look like

nothing - and differ from the NHM paintings which do look more like gorgone above. The dorsal hindwing

looks most like Burke County “nycteis” (Figures 1- 3).

6

ORIGINAL DESCRIPTION

Fact ten. The written original description. This is an exact match for coastal Georgian and

Carolinian “nycteis”. Boisduval & Le Conte could not have produced this without specimens in front of

them or one of them having seen it in nature. (Conjecture: I do not believe that these well informed and

expert scientists were unaware of Hiibner’s gorgone published 23 years earlier. It is highly probable they

had seen his work. Since they had specimens, or at least Le Conte’s personal experience, they would have

immediately recognized these are Hubner’s species (as nominate gorgone is so unlike anything else) and

would not have described it again by another name - ismeria.)

Here is the Original Description translated into English. I have never seen this presented in any of

the literature examining this situation. I have added bold and words in brackets for clarity. Figure A is the

painting accompanying the OD, B & C for are for comparison 4 . Translation by Martin Bailey, Alex

Grkovich, myself and confirmed by Don Lafontaine and Jocelyn Gill of the Canadian National Collection.

Note that the description of the ventral hindwing matches only figure B.

Ismeria Original Description (female)

The species has the size and the appearance of [the species] cinxia of Europe. The top [upperside] of all its wings

are tawny yellow [fulvous] with a lot of black splotches [stain like spots]; these are positioned randomly in the basal

area and form zigzag lines, the others forming two winding lines that traverse veins on the dorsal forewings and one

on the dorsal hindwings which is followed by a row of points of the same color [black]. The marginal borders of all

four wings are black, divided on the forewings by tawny splotches, and on the hindwings by a cord of whitish fulvous

lunules. In addition to this, the apex of the forewings are marked with four or five white dots.

The underside of the forewings differ from the upperside in that, before the edge of the margins, there is a band of

white spots preceded by three or four spots of the same [white] colour.

The underside of the hindwings are tawny with white spots near the base, then an irregular [tawny] band traversing

the median area, and then marginal lunules [semi-circles] of the same color [as the median band]. These [lunules/

semi-circles] are punctuated by a series of black points corresponding to those that appear in the same location

dorsally. The fringe of all wings are black rimmed punctuated by white.

The body and the antennas are the same as other of that species.

The caterpillar is yellow with spines and three black tinted longitudinal stripes. The head is black as well as the scaly

legs [probably the true legs] and the underside; the other legs are yellow.

The chrysalis is a cinder gray with many pale spots and small almost white dorsal tubercles.

This Melitoea is found in the Carolinas and Georgia. It is rarely in collections.

All: females. All: dorsal/ventral. A OD,

i (=nycteis). C. Topotype gorgone.

Figs. 1-3: all females, a: OD illustration of ismeria; b: Louisiana ismeria; c: topotype gorgone. Fig. 4. Hiibner’s OD of

gorgone. Fig. 5. Chevrons on ventral margin of NC paratype female of C. ismeria obsoleta. Fig. 6. Ventral, neotype male C.

gorgone, Burke County, GA. Fig. 7. Typical female C. gorgone carlota (ex. Iowa) (dorsal / ventral). Fig 8. Ventral, SC C.

ismeria ismeria. Note orange marginal line and tawny hindwing ground. All leg. R. Gatrelle except b. Photos by Joe Mueller.

The most pronounced character on the ventral hind wing of gorgone are the white arrowhead shaped

spots. Not only was this not mentioned in the ismeria OD, but the ventral hind wings are described as

tawny with a tawny marginal band of lunules with black spots - precisely as in ismeria (=nycteis ) in

coastal GA. Coastal GA males and females of ismeria (= nycteis) usually have a narrow but bright

marginal orange line on the ventral side of the wings which is never the case with gorgone gorgone. Some

ismeria individuals have orange nudums on the antennal clubs as in the OD picture.

Nominotypical gorgone have very dark ventral hind wings in most individuals (especially when

fresh) of both sexes. Dor sally, I have only two of 30 C. g. gorgone females that are broadly fulvous above

whereas it is common for C. gorgone carlota (Reakirt, 1866) females to be quite fulvous dorsally.

Gorgone males rarely and females seldom have the white pupil in the marginal spot band on the ventral

surface; I have never seen this spot on the DHW of males and it is rare on the DHW of females. This white

spot is common in males and usual and often large in females of C. g. carlota (both dorsally and ventrally).

Carlota is also usually much lighter on the ventral surface. Carlota females (Fig. 7) nearly always and

males usually have a complete row of VHW marginal white chevrons. In nominotypical gorgone, only

females frequently have this while in males it is unusual to find individuals with even most of these

marginal chevrons present. Perhaps less than 5 persons have ever seen nominotypical gorgone. Thus, most

people have only seen subspecies carlota and think of that image when the name “gorgone” is put forth.

(I consider the gorgone specimen figured by Calhoun (2003) which he purports to be from Abbot

and Georgia to actually be subspecies carlota (as labeled) and likely from Ohio. This specimen was

clearly collected by Dyson as it bears his characteristic label (Calhoun 2003). Which label, is the only

original label on the specimen as all the others were later additions. Further, because Dyson was illiterate,

the word “Georgia” on that label was added later. It is certain the inscription was not by Abbot as it is not

in his hand (see Calhoun 2003: his fig. 25). I have long series of both subspecies gorgone gorgone and

gorgone car lota males. The NHM specimen is most definitely the car lot a phenotype, and conversely, not

the nominotypical phenotype. What this specimen looks like says more, and more factually, than any

speculation on the old labels associated with it - when were they put on, by whom, did any get mixed up in

decades of handling.)

What Abbot reared was a “nycteis”. The adults he painted are a composite of nycteis and gorgone.

It is possible Le Conte had Abbot’s painting adjusted to be more like his concept, nycteis. But for almost

two centuries most workers (except F.M. Brown) assumed these were gorgone - even though the Abbot

original painting produced for Le Conte and the published copies don’t look like that taxon. In 1974 Brown

stated ismeria was not gorgone but might be nycteis. He was correct.

MODERN ASSESSMENT

The recognition that there is a subspecies of C. ismeria (described herein) inhabiting the southern

Appalachian mountains segregates subspecies ismeria ismeria from subspecies ismeria nycteis. Calhoun

(personal communication) states that the “type” of nycteis exists in the NHM, London and that the type

locality is Ohio. The taxon nycteis was described from a single specimen painting (Fig. 29). As can be

seen from the figure, the nominate subspecies has well developed and extensive dorsal spotting. Ismeria is

also a broadly spotted taxon dorsally, however, the basic patterns differ, and primarily so, on the hind wing

where in ismeria the median and submarginal spot bands are separated by a thin black line (Figs. B, 25, 27)

while on ismeria nycteis this black area is usually wider and looks more like part of the dark ground and

not a bisecting “line” (Fig. 13, but note Fig. 15). Ismeria ismeria also differs in that the black margin of the

dorsal hind wing is usually wide and thus invasive into the submarginal spot band to the extent that it

partially obliterates the distal third or even half of this band (Fig. 25).

To some, these differences would not be sufficient to consider Ohio nycteis and coastal Georgia

ismeria as distinct subspecies. But bisecting the range of these two is an Appalachian endemic subspecies

which has very restricted fulvous above. This tells us that different evolutionary processes have been at

work and that the two light subspecies (while similar) has each evolved uniquely as separated by this third

eastern US mountain taxon. This situation is evolutionarily complex and I have by no means even begun to

deal with any details of it. I am only stating the most observable manifestations of this situation and

describing the Appalachian herein as new to science. There is thus fertile opportunity for others, especially

geneticists, to investigate the evolutional path of these three subspecies in detail.

In assessing the range of this new subspecies, I have had to also assess the ranges of the other

eastern ismeria subspecies. I now consider ismeria ismeria to have a fairly wide, but narrow, range from

the entire Piedmont region of South Carolina south through southern Georgia and west through coastal

Louisiana and possibly into southeastern Texas. Ismeria is rare to uncommon throughout this range.

On the venter, ismeria is (like all ismeria subspecies) highly variable. But in general, ismeria has

three distinguishing ventral characters - all on the hind wing. 1) a complete row of marginal white or

silvery crescents, 2) fulvous submarginal lunules (especially in females) and suppressed brown marginal

patch (but in some ismeria males this area can be quite dark), and 3) a prominent post basal irregular

fulvous band. The hind wing ground is light fulvous not whitish.

Ismeria is found in varying habitat and is host associated/limited not environment associated/

limited. I have also found this to be true with ismeria nycteis in Iowa: dry prairie to wet bottomland; and

the new subspecies, C. ismeria obsoleta , in the southern Appalachian region: xeric serpentine barrens to

mesic hardwood bottomland. Human activity is both a benefit and detriment to this species. Ismeria taxa

need clearings where its host plants can flourish but not overly managed land where “weeds” are mown or

sprayed. Minimally attended rural roadsides afford good habitat.

9

Chlosyne ismeria obsoleta new subspecies

Common butterflies can be overlooked and understudied. For over 20 years I have been making

almost annual (and often multiple) research trips into western North Carolina. Chlosyne ismeria is a

common component of the mountain fauna. As such, I often encountered, but seldom collected individuals

of this species in this area. In 2003, I segregated the individuals of these mountain specimens into one

series and also relaxed and spread a few I had in papers. I ended up with a series of 15 specimens from a

wide assortment of dates and locations that were amazingly similar, and quite distinct from any other

regional populations I had ever seen or encountered.

Normally, I would not recommend describing a new subspecies from only 15 specimens of a widely

distributed entity. But this series represents a totally unbiased survey sampling over such a long time and

area that the starkly similar, and totally randomly collected individuals, can only be taken to indicate that if

nothing else, the populations in the southern Appalachian region are mostly of this distinct phenotype. My

view is that if just 60% of individuals in this region were of this unique dark phenotype that the genetic high

frequency of this phenotype warrants its subspecific recognition. It is an evolutionally significant departure

from the norm of eastern US ismeria in general. It is also important to note that all Lepidoptera in the

southern Appalachian region tend to melanic facies (J.F. Gates Clarke personal communication). In other

words, this ismeria segregate is significant as part of a larger regional evolutionary trend, and as such,

should to be noted and distinguished as such by formal subspecific description.

Chlosyne ismeria obsoleta Gatrelle, New Subspecies

Diagnosis and description. The obsoleta photos (Figs. 17-24) well illustrate this new subspecies. Dorsally: both

sexes of obsoleta adults are darker than either ismeria or nycteis with all fulvous spots and banding greatly reduced; hind

wing median and postmedian bands are distinctly colored differently in obsoleta (median: yellowish; postmedian: orange),

these bands are the same color (shades of orange) or only subtly different in ismeria and nycteis. Generally, obsoleta is

similar dorsally to subspecies drusius but not as dark, especially in females. Ventrally: males: early season obsoleta can be

boldly patterned on the hind wing, later broods can be quite lightly patterned; hind wing marginal whitish silver spots are not

well developed; marginal orange line can be well developed and similar to subspecies ismeria. Generally, very similar to

subspecies nycteis but markedly lighter patterned than subspecies drusius. Females: early season obsoleta can be boldly

patterned on the hind wing, later broods are usually very lightly patterned on both wings; dark hind wing marginal patch

indistinct to almost absent; row of six submarginal dark spots all with white pupils. Overall: obsoleta is unique in that while

the wings are dorsally melanistic they are ventrally washed out and lightly patterned, especially in females.

Types. Holotype d (Figs. 17-18): NORTH CAROLINA: Macon County, Jones Knob 4200’, 8 July 1989. Allotype

9 (Figs 19-20): NORTH CAROLINA: Macon County, Franklin Greenway, 16 May 2002. Paratypes: 10 c? d, 3 9 9 : NORTH

CAROLINA: Macon County: Jones Knob: 2dd, 1 9, 8 July 1988 (Figs. 23-24), 1 d, 8 July 1989, 2d d, 21 June 1992, 1 d,

9 June 1993. Clay County: Buck Creek: Id, 27 May 2003, 19, 25 May 2002; Chunky Gal: 19, 27 May 2003; Alleghany

County: Rock Creek Rd off Farmers Rd, 2100’: 1 d, 13 August 2003. GEORGIA: Union County: Jet. Hwy 180 & 126: 1 d,

9 May 2003 (Figs. 21-22). VIRGINIA: Grayson County: Fox Knob, FS Rd. 852, Id, 22 August 2003. All leg R. Gatrelle

The holotype and all paratypes are currently in the TILS Museum of the Hemispheres’ collection, Goose Creek, South

Carolina. Some paratypes will be deposited in other institutions.

Etymology. Obsoleta is derived from the obsolescence of the dorsal fulvous and the ventral dark markings.

Remarks. There is a tendency in females to have the ventral marginal whitish silver spots form chevrons (Fig. 5).

This is present in 2 of the 4 females. Because the sample is so small, this may or may not have taxonomic significance.

Those who encounter or collect this taxon should assess specimens for the presence of this “ gorgone like” character. The

dates of the type series encompass the entire flight season and the altitudinal range is from 2000 to almost 5000 feet. While

indeed a very small sample, it is none the less a good indicator of the presence of this subspecies in what I refer to as the

Appalachian Peninsula. This mountain range dips deep into the otherwise hot lower austral zone in North Carolina and

Georgia. It is indeed a unique faunal peninsula in the same manner as peninsular Florida in its biotic uniqueness.

Range. The range of this taxon needs to be established. It is in extreme north Georgia and apparently ranges into

extreme western Virginia (Grayson County paratype). It should be expected in the eastern mountains of Tennessee and

southern West Virginia and is likely in eastern Kentucky. However, I assume none of this. It is definite only in northeast

Georgia and southwestern North Carolina. It is a striking subspecies, especially the first brood females.

10

Figures 9-28. Chlosyne ismeria subspecies. Figs. 9-10 (D/V): d C. i. drusius : 14 July 1975, Boulder Co., Colorado (leg.

unknown). Figs. 11-12 (D/V): 9 C. i. drusius : 30 July 1948, Boulder Co., Colorado (leg Don Eff). Figs. 13-14 (D/V):

d topotype C. /. nycteis : 9 August 1984, Dayton, Ohio (leg. Andy Anderson). Figs. 15-16 (D/V): 9 C. i. nycteis : 18 July

1966, Shelby Co., Iowa. Figs. 17-18 (D/V): d holotype Chlosyne ismeria obsoleta : 8 July 1989, Jones Knob, Macon Co.,

North Carolina. Figs. 19-20 (D/V): 9 allotype C. i. obsoleta : 16 May 2002, Franklin, Macon Co., North Carolina. Figs. 21-

22 (D/V): d paratype C. i. obsoleta : 9 May 2003, Hwy. 180, Union Co., Georgia. Figs. 23-24 (D/V): 9 paratype C. i.

obsoleta : 8 July 1988, Jones Knob, Macon Co., North Carolina. Figs. 25-26 (D/V): d C. ismeria ismeria : 23 May 1963,

Marianna, Jackson Co., Florida (leg. Fuller). Figs. 27-28 (D/V): 9 C. ismeria ismeria. 11 June 2002, vise. Johnson’s

Fanding, Allendale County, South Carolina. All leg. R. Gatrelle unless otherwise noted. Photos by Joe Mueller.

11

ORIGINAL DESCRIPTIONS AND SUBSPECIES COMMENTS

Melitaea ismeria (Chlosyne ismeria ismeria)

The original description of Melitaea ismeria is presented herein on page 7 and discussion on page

8 under Modem Assessment. I here add additional comment. It is possible that subspecies nycteis does not

occur in the deep South (Mississippi to South Carolina) at all. It all depends on how narrowly or broadly a

taxonomist defines these taxa. As I stated above, I do not just consider phenotype but also the known or

likely biogeographical evolutionary path these taxa have taken to arrive at their current state of being. For

example, I now view ismeria more broadly than I did in 1989 in now considering it the subspecies of the

southern Gulf coast region, lower Georgia and all of South Carolina except the far northwest SC mountain

edge area. How this taxon transitions (if it does) east of the Appalachians up the eastern seaboard into

subspecies nycteis remains to be investigated (see discussion under obsoleta below). How it transitions (if

it does) west of Louisiana, and north in Mississippi and Alabama needs to be investigated also.

Melitaea nycteis (Chlosyne ismeria nycteis)

The original description of Melitaea nycteis Doubleday, [1847] consists of a figure and caption

(Fig. 29). John Calhoun has stated in personal communication that he is preparing a paper in which he will

formalize the type locality as Ohio (and perhaps more localized) and figure the “type” specimen in the

NHM, London and discuss its typification status (holotype or syntype). In 1848 a text was published to

accompany this painting. It was very brief and is as follows. (Don Lafontaine provided the following

additional comment. “This text was presented in the second entry under Section II. Phyciodes. Which would

effectively make it Melitaea (Phyciodes) nycteis, but since this wasn't done for the plate, which was

published the previous year, then the OD is Melitaea nycteis P)

23. MEL. NYCTEIS Doubleday & Hewitson, t. 23, f. 3 (1847).

United States (Middle States). B. M.

Subspecies nycteis , like all ismeria subspecies, should be assessed by two criteria weighed

together: phenotype and biogeographical evolution. The geographic range and phenotype of nycteis are

both broad and variable. It might be most accurate to define the range of nycteis as being where the other

subspecies are not, and its phenotype as being that which is not characteristic of the other subspecies. Thus,

nycteis ranges in the west eastward from the ranges of drusius and reversa ; in the east, north of the range of

ismeria and west, north, and northeast of the range of obsoleta. Correspondingly, its phenotype “blends” at

the range periphery into all of the other subspecies. Add to this the tendency to local and regional variation,

and nycteis becomes a rather ambiguous taxon. This broadness is what defines and delimits it.

I have consulted many books on this but will only refer to those I consider most pertinent. The two

males (page 125) figured in Butterflies of Michigan (Nielsen, 1999) reflects the above very well. The

female depicted on that page can be said to be “typical” of the nycteis phenotype. “Typical”, with nycteis

12

means: average. In Butterflies of Manitoba (Klassen et. al. 1989), nycteis is attributed to the south central

and southeastern part of that province. Subspecies reversa is limited to the western Riding Mountain region

of the province with reversa phenotype specimens being found as far east as Winnipeg. In Butterflies of

Virginia (Clark & Clark 1951), the text and illustrations can now be applied to reveal that nycteis may

only exist is that state at the northern and northeastern regions. It is also likely that in much of Virginia

populations are not technically referable to subspecies and represent a “blend zone”. In Butterflies and

Moths of Missouri (Heitzman & Heitzman 1987), the male figured is mostly the obsoleta phenotype

dorsally (too much fulvous at base of wings) and the female the nycteis phenotype dorsally. Ventrally, the

female is mostly the obsoleta phenotype (note the marginal spots are all white pupiled, but post basal tawny

band fairly bold) and the male the nycteis phenotype. This is not only not surprising but might be expected

due to the well known affinity of taxa in the Ozark plateau with the southern Appalachian fauna. The range

of subspecies nycteis in Missouri may well be limited to the northern and northeastern part of that state.

The populations in the remainder of the state (and into Arkansas) need taxonomic assessment.

Phyciodes nycteis drusius (Chlosyne ismeria drusius)

The original description of Phyciodes nycteis drusius W.H. Edwards, 1884 was in Papilio, 4: 57,

1884 and is as follows. There were no illustrations. F.M Brown (1966) restricted the TL to Turkey Creek

Junction, Jefferson Co., Colorado and designated a lectotype in the Carnegie Museum

Phyciodes Nycteis, var. Drusius. -1 give this name to the West¬

ern form of the species; distinguished by the excess of black on upper

side, and consequent restricted fulvous spaces. The fulvous is dull, and

the black pale. This is the Colorado and Arizona form of Nycteis.

This taxon ranges from New Mexico into Montana and perhaps southwestern South Dakota. James

Scott (personal communication) states this is an east slope Rocky Mountain taxon and that the west slope (in

Colorado) populations are more like subspecies nycteis. (This surely calls for taxonomic investigation.) In

Butterflies of the Rocky Mountain States (Ferris & Brown 1980), it is stated that specimens from the

Black Hills in South Dakota are intermediate between drusius and “the northeastern Minnesota phenotype”.

I take this to mean intermediate to nycteis. South Dakota is a transitional region and, like Virginia, will

have populations not technically referable to a specific subspecies.

Phyciodes nycteis reversa (Chlosyne ismeria reversa)

Phyciodes nycteis reversa F. and R. Chermock, 1940 was described in the Canadian Entomologist,

72: 83, 1940. The OD is as follows.

Phyciodes nycteis reversa new race

In nycteis, there exists a parallelism to Melitaea harrisi and its

forms; Melitaea harrisi, Phyciodes nycteis', Melitaea harrisi liggetti

(Avinoff), Phyciodes nycteis drusius (Edwards); and Melitaea harrisi

albomontana, Phyciodes nycteis reversa. This new race, although

displaying the same general characters as typical nycteis, may be easily

separated by the abbreviation and omission of black markings as in

Melitaea harrisi albomontana. The under surface has the

characteristic markings of nycteis, but less distinct.

Holotype- d, June 27, Riding Mountains, Manitoba.

Allotype- 9, June 27, Riding Mountains, Manitoba.

Paratypes- 1 to 50, Sand Ridge, Manitoba, and Riding

Mountains, Manitoba.

The holotype and allotype were taken in copulo.

13

I have never seen specimens of this outside of the literature. The evolutional parallelism mention in

the OD is noteworthy in my view. It appears from the comments of Klassen (1989), various photos in the

literature, and the OD that this is a localized subspecies endemic to the Riding Mountain region of

Manitoba, Canada. Because the tendency of both the populations surrounding reversa and other mountain

populations of ismeria is toward darker phenotypes, the light bright reversa phenotype is surely a genetic

character trait and evolutionarily significant.

Chlosyne ismeria obsoleta

Described herein, Chlosyne ismeria obsoleta’s evolutionary origin is unknown. The presence of

obsoleta like specimens in the Ozark region of Missouri and Arkansas would indicate this to be a taxon of

older rather than newer origin dating back to when these two regions were connected ecologically and

biologically (e.g. 28,000 B.P. Ice Advance, or 161,000 BP Ice Advance). It is possible that subspecies

obsoleta is present in the Ozark region as a remnant of this former continual range. An indication of this

would be the presence of obsoleta phenotypes across northern Alabama and Mississippi.

The status of obsoleta in Virginia needs assessment. The female figured in Clark & Clark (1951)

from Lunenburg County in south central VA is certainly at least transitional to obsoleta if not obsoleta.

That region of VA is apparently about as far east as species ismeria ranges in that state. If obsoleta, and or

a near obsoleta intermediate, ranges all across southern VA, then it isolates subspecies ismeria east of the

Appalachians from subspecies nycteis of the northeastern US. I have determined from specimens I have

collected in the South Carolina piedmont of Spartanburg County (north central SC), that the subspecies of

SC is ismeria (except for the narrow edge of mountains in northwest SC). Piedmont and upper coastal

North Carolina would then be the region of transition between subspecies ismeria and obsoleta.

Subspecies nycteis would then not be present in NC, SC, GA, far eastern TN, southern WV, northeast AL,

southern and southwestern Virginia.

CONCLUDING STATEMENTS

Definition. Subspecies: Any regional population of a species that has evolved into a unique reproductively

stable component of that species.

Chlosyne ismeria is a wide spread species composed of a complex set of five known subspecies.

We have the choice to simplify (lump) and basically ignore the past evolution and present reality of these

subspecies, or, to delve into this complex more specifically. The regional populations of all Lepidoptera

(species or subspecies) did not come into being in situ where we encounter them in our modem age. Thus,

just because any two populations (species or subspecies) are now in close geographic proximity, it does not

follow that evolutionally they came into being either from each other or biogeographically near each other.

I do not see a “dine” of any kind with any of these taxa at this point. I see five uniquely evolved taxa

which, at this stage in their evolution, are still the same “species”. I see that where they come into current

geographic contact there is phenotypic transition - genetic flux exists. This means nothing more than the

obvious, that these are regions of subspecifically interactive contact.

There have been objections to the use of ismeria as the binomial for this species rather than nycteis

even though the use of ismeria is Code compliant per Article 23.9. Those who favor nycteis as the

binomial may appeal to the Commission per Article 23.9.3. This is acceptable to me. However, any

attempt to sink ismeria is not acceptable as it is the available and valid name of a subspecific taxon. This

name is not a nomen dubium as is clear from the OD text, this paper and my 1998 neotypification; nor is it a

synonym of gorgone as absurdly proposed by Calhoun (2003) without any examination of the OD

description or any specimens of “nycteis” from anywhere. Calhoun simply compared gorgone with gorgone

and then with irrelevant Abbot art and declared it to all be ismeria.

14

ACKNOWLEDGMENTS

I thank Norbert Kondla for his review of and suggestions on this paper. Martin Bailey, Alex

Grkovich, Don Lafontaine and Jocelyn Gill for translation and checking of the English translation (from

French) of the Boisduval and Le Conte ismeria OD text. John Calhoun for information on the type and type

locality of nycteis. Don Lafontaine for providing original descriptions and David Wright for a copy of the

ismeria OD text. Chris Durden for glaciation time periods re Ozark Appalachian ecology.

LITERATURE CITED

BOISDUVAL, J.A., & J.E. LECONTE. 1833. Historie generale et iconographie des Lepidopteres et des chenilles de

l’Amerique Septentrionalis. Vol. 1. Paris. 228 pp.

BROWN, F.M. 1974. The Butterfly Called ismeria by Boisduval and Le Conte. Bull, of the Allyn Mus. Bull. No. 16.

12 pp.

CALHOUN, J.V. 2003. The History and True Identity of Melitaea ismeria (Nymphalidae): A Remarkable Tale of

Duplication, Misinterpretation, and Presumption. J. Lepid. Soc. Vol. 57:3, 204-219.

CHERMOCK, F.H. & R.L. CHERMOCK 1940. Some New Diurnal Lepidoptera From the Riding Mountains and the

San Ridge, Manitoba. Canadian Entomologist, 72: 83.

CLARK, A.H., and L.F. CLARK. 1951. The Butterflies of Virginia. Smithsonian Misc. Coll. 116 (7): vii + 239 pp.

COWAN, C.F. 1969. Boisduval and Le Conte; Historie generale et iconographie des Lepidopteres et des chenilles de

l’Amerique Septentrionalis. J. Soc. Bibliog. Nat. His. 5:125-134.

DOUBLEDAY, E. & W.C. HEWITSON 1846-1850. The Genera of Diurnal Lepidoptera, etc. Vol. 1, London.

FERRIS, C.D. & F.M. BROWN 1980. Butterflies of the Rocky Mountain States. Univ. of Okla. Press, Norman. 441

pp.

GATRELLE, R.R. 1998. The Rediscovery, Taxonomy, and Biology of Chlosyne gorgone and Chlosyne ismeria

(Nymphalidae) in Burke County, Georgia. TTR, Vol. 1:2, 9 pp. TILS, Goose Creek, SC.

HEITZMAN, J.R., & J.E. HEITZMAN. 1987. Butterflies and Moths of Missouri. Missouri Dept, of Conserv.,

Jefferson City, MO. 385 pp.

HUBNER, J. 1806. Sammlung exotischer Schmetterlinge. Augsburg.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International Code of

Zoological Nomenclature, Fourth Edition (1 January 2000). London, UK: The International Tmst for Zoological

Nomenclature. 306 pp.

KLASSEN, P., A.R. WESTWOOD, W.B. PRESTON, W.B. McKILLOP 1989. The Butterflies of Manitoba. Man.

Mus. of Man and Nature, Winnipeg, Man., Canada. 291pp.

NIELSEN, M.C. 1999. Michigan Butterflies & Skippers. Mich. St. Univ. Press. 248 pp.

The Taxonomic Report

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15

Volume 4

15 November 2003

Number 5

THE CASE FOR TAXONOMIC RECOGNITION OF THE TAXON

ENODIA ANTHEDON BOREALIS A.H. CLARK (SATYRIDAE)

HARRY PAVULAAN 1

HISTORICAL TREATMENT OF ANTHEDON AND BOREALIS IN THE LITERATURE

AUSTIN H. CLARK 1936: REVIEW OF THE GENUS ENODIA

line on the under side of the fore wing with a slight angle,

of the male; and in the absence of the broad furry band on

iSSK

Type: U.S.N.M. no.

Referring to E. anthedon , Clark makes the following statement:

A third form [E. portlandia and E. creola being the first two], occurring in the East from southern New Hampshire

southward to the higher altitudes of North Carolina and possibly farther (pi. 22, figs. 1, 2) is lighter, less brightly marked,

and usually smaller than true portlandia. This is the form referred to as portlandia by Skinner and Richards and by

American authors generally. It is locally frequent in the mountains of Virginia, where its quick and active movements

and its habit of keeping generally low down in the underbrush distinguish it rather sharply from the less active and

commonly high flying tme portlandia of the coastal region. Since none of the names that have been proposed for species

of this genus is applicable to it, it may be known as Enodia portlandia anthedon.

It is interesting to note that the description of “anthedon” from the “mountains of Virginia” in the

above paragraph is in fact quite suggestive of both the habitats and habits of borealis , as we discuss

below. This brings into question whether E. anthedon borealis actually ranges far southward from its

historically described range, and also whether records of “ anthedon ” from higher elevational areas of the

middle and southern Appalachian Mountain regions should actually be applied to borealis.

7 8 9

Figures 1-9. Enodia anthedon subspecies. Fig. 1 (D/V): d E. a. anthedon : 4 July 2001, Merrimack, NH (leg. Grkovich). Fig. 2

(D/V): d E. a. nr. borealis: 15 July 2001, Rt. 112, Carroll Co., NH (leg. Grkovich). Fig. 3 (D/V): d E. a. borealis: 7 July 1984,

Perthuis (Portneuf), Ouebec, Canada. Fig. 4 (D/V): d E. a. anthedon: 27 May 1994, Seneca, Montgomery Co., MD (leg.

Pavulaan). Fig. 5 (D/V): 9 E. a. anthedon: 18 May 1996, Herndon, Fairfax Co., VA (leg. Pavulaan). Fig. 6 (D/V): 9 E. a.

borealis: same data as Fig. 3. Figs. 7-8 (D/V): d holotype E. anthedon OD figures (1936). Fig. 9 (V) 9 E. a. nr. borealis: 27

July 2002, Doughton State Park, Alleghany Co., NC. Photos 1 -6 by Joseph Mueller; photo 9 by Bruce Grimes.

2

Referring to borealis , Clark then writes:

Enodia portlandia borealis is very variable, but it seems always to be distinguished by the broader dark border on

its hind wings above and by the relatively broad darker area between the row of spots and the submarginal light line

on the hind wings below.

In his Original Description of borealis, Clark notes:

...twenty two specimens are at hand from the following localities:

Manitoba: Miniota, June 17, 1926, H. Gibbon (1); July 1, 1922 (1); July 10, 1920 (12). Winnipeg,

July 1-7 (1); no date (1).

Ontario: Hymers, July 1-7 (2).

Quebec: Meech Lake, Ottawa County (1); somewhat intermediate between this and the previous

[anthedon ] form.

Maine: Sebec Lake, July 16-23 (1); July 24-31 (2); more or less intermediate between this and the

preceding form.

Clark then gives a “Key to the (...interrelationships of the...) forms included in the Genus Enodia ”:

a 1. Male with the long fore wings more pointed than those of the female, above with a broad furry band in¬

terrupted at the veins and by long triangles in the interstices; under sides of forewings with post-medial line

irregular, interrupted above vein 6, outwardly oblique between veins 6 and 4, and usually slightly indented

on vein 5; on hind wings below the fourth ocellus is smaller than the fifth. creola (pi. 22, figs. 5, 6)

a 2. Sexes practically alike; under side of fore wings with postmedial line more or less oblique from costa

to vein 4, or just above it; on hind wings below the fourth ocellus is larger than the fifth. portlandia

b 1. Wings beneath with the rows of ocelli edged with white interiorly and more or less completely

exteriorly; on the fore wing a white band runs from the costa to the region of the second ocellus, and

beyond this a narrower white band runs from the ocellus to the upper part of the first ocellus; row of

ocelli on the under side of the fore wings curved; second and third ocelli on the under side of hind wings

with elongate pupils, and fourth usually without a pupil . portlandia portlandia (pi. 22, figs. 3, 4)

b 2. No white on wings beneath; row of ocelli on outer side of fore wings below straight; all the ocelli on

hind wings below have similar circular pupils.

c 1. Dark border on hind wings above narrow and tapering anteriorly; on the hind wings below the

dark band between the light line bordering the fourth and fifth spots and the submarginal light line is

little, if at all, broader than the distance between the submarginal light line and the margins of the wing

. portlandia anthedon (pi. 22, figs. 1, 2)

c 2. Dark border on hind wings above broader and more uniform, not narrowing appreciably

anteriorly; on the hind wings the dark band between the light line bordering the fourth and fifth spots

and the submarginal light line is broader, usually much broader, than the distance between the

submarginal light line and the edge of the wing; ground color below browner and usually more

uniform

. portlandia borealis

Clark concludes his discussion of the genus Enodia by making the following observation:

Although when typically developed the four forms included in this genus Enodia are quite different, three of them

are very closely related.... It appears that typical portlandia intergrades more or less with anthedon, and the latter

intergrades more or less with borealis, the relationship between the three suggesting the relations between Cercyonis

... pegala, ...alope, and ....nephele occurring in the same general region...

3

HISTORICAL TAXONOMIC TREATMENT OF BOREALIS:

Prior to 1971

We reviewed five publications from the period prior to 1971, and found that direct reference to E.

anthedon borealis occurred in each. Macy & Shepard (1941), in their book Butterflies - A Handbook of

the Butterflies of the United States, Complete for the Region North of the Potomac and Ohio Rivers and

East of the Dakotas , included Enodia portlandia borealis , extensively citing Clark’s original description

outlined above, but added that “specimens from Minnesota belong to the race anthedon .”

In A Field Guide to the Butterflies of Eastern North America , Klots (1951) included Lethe

portlandia borealis with the descriptive comment: “Smaller, darker, dark border of HW above broader;

dark submarginal band of HW beneath broader.” The author states further: “The species as a whole

represents something of a ‘cline,’ although there is doubtless some habitat (ecotypic) segregation due to

isolation of colonies. The subspecies, not well marked, intergrade greatly, especially northward.”

In The Lepidoptera of Pennsylvania , Tietz (1952) listed Enodia portlandia borealis but it is not

clear whether he believed it occurred in Pennsylvania, since he gave no specific locations for the state.

Forbes (1960) listed Lethe portlandia borealis in Lepidoptera of New York and Neighboring States

with the following explanation: “Hardly distinct, perhaps smaller, somewhat paler and still more even in

ground color”, but neglected to specify distribution.

Cyril dos Passos (1964) compiled the first complete catalog of North American butterflies for the

Lepidopterists’ Society, published as Society Memoir #7, and listed subspecies Lethe portlandia borealis.

John H. Masters’ 1971 Paper

Masters published: A Note on Lethe anthedon borealis (Satyridae), in the Journal of the

Lepidopterists ’ Society 25(4) 1971. In this paper, he presented observations of both nominotypical

anthedon and subspecies borealis in the western Great Lakes region indicating that, despite their “slight”

phenotypic differences, there were some significant biological differences between them. Masters found

that the most useful character for separating the two subspecies was in the ground color of the ventral

hindwings. These he described as follows: “.. .in borealis it is a dull and uniform brown while in anthedon

the background seems to be composed of several shades of brown and is much brighter.” Masters also

noted: “This distinction is especially evident in looking at the butterflies in series, which avoids comparing

individual differences.” The author plotted the distribution of both phenotypes in the western Great Lakes

region and noted that the dividing line between both subspecies correlated very closely to the approximate

boundary between the Canadian and Transition Zones in this region (Roberts, 1936). “Nearly every

specimen examined”, he wrote, “could be placed reliably into one subspecies or the other. Some degree of

intergradation was noted in the character of the dark border on the dorsal hind wing, and to a lesser extent

in the other characters, but in no case was more than one character involved in the intergradation.”

From this narrative, it is evident that in using the word “slight” Masters meant only that there were

not many differences. He was not saying there were no significant differences. He clearly related that he

considered the ventral hindwing ground color a significant character to reliably and consistently distinguish

these two taxa.

Masters then stated that behavioral differences, including habitat preferences, differentiated both

subspecies, observing that “differences in habitat and habits are much more conspicuous.” Anthedon males,

Masters observed, “are very territorial and each male will occupy a favorite perch at some distance from

the perches of his nearest neighbor”. We have observed males of anthedon selecting in particular mid-level

perches on tree trunks which overlook these woodland glades or other small openings in the forest. On the

other hand, Masters stated that borealis : “...does not occupy a fully wooded environment, but prefers a

very open wooded environment with lush undergrowth”. We have found borealis especially near bogs or

4

marshes in the interface of mixed plant associations where they give way to forest. Males of borealis are

described by Masters as exhibiting “none of the territorial characteristics that are so pronounced with

anthedon ”, rather being “in fact, quite gregarious in habit”. Our observations confirm this as we have

frequently found them congregating together in bushes. They often perch on low and mid-level vegetation

but they also perch at higher levels on tree trunks and inside the leaf canopy - where they are well

concealed. Masters also states that borealis “seem to be quite ‘amiable’ together and the aerial encounters

of males, that are so common in nominate anthedon and other territorial species, never appear to occur.”

Masters concluded his paper with the following statement regarding the anthedon/borealis

subspecific relationship.

One of the major criticisms leveled at the trinomial and its usage in taxonomy is that the subspecies, as

currently defined and used, fails to distinguish between weakly and strongly differentiated geographical subspecies

and treats them all alike. Descriptions and identifications of populations are essentially based upon morphological

distinction and consequently taxonomy has been strongly oriented in this directions and populations which show

strong morphological divergence have attracted the most attention. However, we have in Lethe anthedon\ borealis, a

very good example of very marked physiological (behavioral) differences along with weakly developed morpho¬

logical differences. Differences between species, subspecies, or any taxonomic category, may be physiological,

morphological, or both, and they may be phenotypic or genotypic in each case. Fortunately physiological

differences, which are far more important in the long mn, are usually accompanied by at least minor morphological

changes which allow the taxonomist to distinguish and name the populations exhibiting them. On the other hand, as

far as we know, morphological differences are usually accompanied by at least minor physiological differences; if

they weren’t, there would be little point in pinning a name upon them.

The North American [Enodia\ are a good example of the problems in relying exclusively upon

morphological characters for species distinctions. Ehrlich (1961) cited [Enodia] as one of only fourteen North

American genera of butterflies that represented no problem to the taxonomist because speciation is quite distinct.

Since that time field work and behavioral studies have forced us to increase the number of recognized species from

three to five and we have become aware of the classification problems in the populations of borealis and

[Satyrodes] fumosus Leussler, both of which are still treated as infraspecific, but with reservations.

Masters thus concluded that, along with the nearly allopatric nature of their morphological

[phenotypic] differences, the physiological distinctions discussed above justify the retention of Enodia

anthedon borealis as a valid, separate and distinct subspecies of Enodia anthedon.

We state that these two may in fact be sibling species. The sinking of borealis into anthedon,

based on assessment via museum specimens devoid of observations of living behavior, has effectually

closed the mental door of investigating the possibility of possible speciation.

After 1971

For the post-1971 period, we reviewed a rather extensive list of publications in order to evaluate

the status of Enodia anthedon borealis in those publications.

Hooper (1973), in Butterflies of Saskatchewan lists Lethe portlandia borealis and states that

“[borealis] is found in poplar woods where it often lands on leaves about five feet from the ground and on

tree trunks”.

In Butterflies and Skippers of New York State (Shapiro, 1974), the author lists only species entry

anthedon , but states: “Specimens from eastern New York (Clinton Co. to lower Hudson Valley and S.I.

[Staten Island]) lighter and brighter than those from farther west.”

Brower (1974) makes an interesting comment in A List of the Lepidoptera of Maine, in which he

lists Enodia portlandia anthedon, but states: “Part of the type series of E. p. borealis Clark were from

Maine. An attempt has been made to restrict borealis to Canada.” The author appears to have considered

his examined material as referable to nominotypical anthedon.

Thomas C. Emmel, in The Butterflies of North America (In Howe, 1975), listed subspecies Lethe

portlandia borealis and states: “This pale northern subspecies grades southward into [subspecies]

anthedon .” Plate 13 in Howe illustrates a female specimen (#10) of anthedon (Streator, Ill. 29 Aug. 54)

appearing somewhat darker and displaying more wing contrast than the female specimen of borealis (#13)

(Menominee Co, Mich., 1 Aug. 1950), which appears considerably more pale by comparison, while

displaying less contrast in wing markings.

Pyle (1981), in the National Audubon Society Field Guide to Butterflies , makes no direct reference

to borealis , but notes that “...in the much cooler northwestern portion of its range, the Northern Pearly

Eye [anthedon\ shifts it habitat to more open woodlands with sedge marshes; the butterfly changes its

behavior as well, gathering in groups and perching on bushes...”, a statement absolutely in agreement

with Masters’ description of borealis.

Miller & Brown (1981) compiled the second complete catalog of North American butterflies for

the Lepidopterists’ Society, published as Memoir #2 . In this catalog, the authors listed borealis as a

junior synonym of Enodia anthedon (entry #711) without explanation. It was after this treatment that

the authors of most subsequent guides of a regional scope decided to follow suite.

In a journal paper entitled Butterflies of the Ottawa District (Layberry, et. al., 1982), the authors

list subspecific entry Lethe anthedon borealis.

Despite the arrangement in Lep. Soc. Memoir #2, Hodges (1983) listed subspecies Enodia

anthedon borealis (as entry #4568.1(a)).

Tilden & Smith (1986) followed Miller & Brown, making no reference to borealis under the

species entry for Enodia anthedon , in A Field Guide to Western Butterflies.

Scott (1986), in The Butterflies of North America, similarly provided no mention of borealis under

the entry of Lethe anthedon (Scott dropped most North American butterfly subspecies from usage and

apparently listed only those which he personally considered sufficiently different from other subspecies

in morphology).

The authors of The Butterflies of Manitoba (Klassen, et. al., 1989) state: “Two subspecies are

recognized. Enodia anthedon borealis occurs in southern Manitoba.” Extensive life history is included.

The habitat is described as “damp deciduous forest edges, clearings, glades, roads and trails, especially

where the underbrush is thick.”

Ferris (1989) compiled a supplement to the Miller & Brown ‘Catalogue/Checklist’

(Lepidopterists’ Society Memoir #3) and retained the designation of junior synonym by making no

changes to the status of borealis as listed in Lep. Soc. Memoir #2,

Opler & Malikul (1992), in the revised Peterson Field Guide to Eastern Butterflies , did not include

borealis in the guide’s checklist, though the inclusion of “Damp deciduous woods, usually...near

marshes” hints of borealis habitat.

The authors of Alberta Butterflies (Bird, et. al., 1995) following authors of other Provincial guides,

listed Enodia anthedon borealis , and describe the habitat as “poplar woods near streams and lakes”.

In The Butterflies of Canada (Layberry, et. al., 1998), borealis is, rather interestingly, not listed in

the ‘Checklist of Canadian Butterflies’, nor did the authors mention it in the text under species entry

Enodia anthedon. The omission of borealis from both the checklist and the text was made despite the

authors’ describing almost in detail the preferred habitat of borealis , stating that “.. .it is found only in rich

deciduous or mixed wooded areas, usually where the undergrowth is thick” (this is again suggestive of the

habitat for borealis as described by Masters), and also despite figuring a male specimen of borealis (pi. 18,

fig. 1) from the Riding Mountains National Park, Manitoba, located 60 miles north of the town of Miniota,

Manitoba (the origin of fourteen of the twenty two specimens used by Clark in his Original Description

of borealis). The omission of borealis in The Butterflies of Canada also suggests the application of junior

synonym status of borealis to anthedon (despite the same authors’ 1982 publication Butterflies of the

Ottawa District in which, as we state above, borealis was listed as a valid subspecies of anthedon).

6

Opler & Wright (1999), in the revised Peterson ‘Field Guide to Western Butterflies’ likewise

make no reference to borealis , while appearing at least in part to describe the habitat of borealis as

“Damp deciduous woods, usually...near marshes”.

In Michigan Butterflies & Skippers (Nielsen, 1999) Ists only species Enodia anthedon without

any reference to subspecies.

And finally, Le Guide des Papillons du Quebec (Handfield, 1999) describes only Enodia

anthedon, while referencing Masters’ 1971 paper on borealis in the Bibliography, and offering the

following notes, first on habitat: “Partly shaded clearings, undergrowths and edges of humid deciduous

and mixed forests, boggy or sedgy”; and then on behavior: “...it is encountered often along the edges of

humid woodlands, perching high enough on vegetation or on tree trunks with its head down...with its

wings closed, it is difficult to observe...” (translations from the French). It should be noted, in conclusion,

that a number of these publications make indirect references to borealis in terms of habitat and/or

behavior, while otherwise making no direct reference to borealis.

Many of the newer major regional field guides mentioned above (Opler & Malikul, 1992; Opler &

Wright, 1999; also Brock & Kaufman, 2003) make minimal reference to subspecies.

FIELD OBSERVATIONS IN NEW ENGLAND AND ELSEWHERE IN THE EAST

We embarked on this study after corresponding over our personal observations of Enodia

“anthedon ” in New Hampshire. During July 2001, one of us (Grkovich) encountered two colonies of

“anthedon ” in widely different parts of the state.

The first colony, of anthedon , was observed on July 4, 2001, near the town of Merrimack, in the

south-central part of the state, 10 miles north of the Massachusetts State line at approximate elevation of

300 feet. Here, along the Rookery Trail, behind (east of) the Fidelity Investments Inc. buildings on

Constitution Road near Exit 10 of Interstate 93, specimens were observed along a gravel access road

through dense Transition Zone pine-oak-birch forest, with individuals perching on birch and oak leaves in

territorial fashion along the edges of the road/trail, invariably 4 to 6 feet above the surface. Perched

individuals maintained distinct territorial behavior, as the average distance between two specimens was

not less than approximately 30 feet. Others were observed inside the forest cover away from the trail,

perching on and flying short distances between leaves and small tree trunks. No individuals were

observed perching anywhere at a distance other than the previously stated 4 or 6 feet above either the

surface of the trail or the ground inside the forest. Perched specimens were quite wary and were rather

difficult to approach, and tended to take flight in response to any sudden human movements, sometimes

relocating to another perch along the trail, but also flying into cover. Individuals (Fig. 1) were in good

condition but demonstrated some wear and were no longer fresh; it was thus estimated that the flight had

begun approximately one week to 10 days earlier during the fourth week of June. A second visit to the

site was made on August 25, 2001 with only a very worn anthedon female being found.

On July 15, 2001 a second colony, this time of borealis , was encountered in an entirely different

type of habitat in the lower Canadian Zone, along the Kancamagus Highway (Route 112) in Carroll

County, near the boundary of Grafton County at elevation of approximately 1500 feet. The habitat is

located 2.6 miles west along the Kancamagus Highway from the intersection of Bear Notch Road, which

extends eight miles north to Highway 302 at the town of Bartlett. The colony was found approximately

1600 feet north of the highway, in a partly shaded but open woodland with very thick moist undergrowth

of grasses, dense sedges and bushes, and other plants such as blackberry, black cohosh etc. The open

forest lies in a low, wide flood plain of the nearby Swift River, and is dominated by firs, spruce, sugar

maples, poplars, birches, willows etc.

As one hikes the distance from the highway into the habitat, one first encounters a small boggy

area (however, no typical bog butterfly species have been found there), then a large open, moist field with

a deep and thick grassy, sedgy area, after which the open field gives way to the open forest. The first

specimens were encountered approximately 150 feet into the woodland, where numerous specimens

(about two dozen) were seen occupying a rather restricted glade area which measured no more than 50

feet in width by 100 feet or so in length. The differences in behavior between these specimens and those

of the colony at Merrimack were recognizable at once. These displayed a decidedly gregarious behavior

quite suggestive of a colony of Asterocampa celtis or clyton , occurring in groups as numerous as a half

dozen or more. They were perching on the bushes and grasses from approximately 6 feet to as low as a

foot above the ground, but also on tree trunks inside the leafy canopy as high as 8 to 12 feet high where

they would become well concealed.

No territorial behavior whatsoever was observed in this colony, as opposed to the individual

behavior noted at Merrimack; sometimes as many as three or more specimens were observed perching on

the tree trunk or plant - or even on the same large leaf. The flight of these, as observed, was quicker and

more direct than the Merrimack anthedon. Perched specimens, particularly when sitting upon low

vegetation, appeared also to be not nearly as wary and were more easily netted. Also in contrast to those

at Merrimack, specimens encountered here were in fresh condition, indicating a very recent emergence of

the brood. A follow-up visit to the habitat was made on August 8, 2001, with no specimens being found at

that time; apparently, the flight had ended.

8

No specimens were seen during a visit on July 15, 2002. This likely indicated only that the flight

had not yet begun due to an abnormally cold, wet and late spring in New England in 2002. On a visit to

the site on August 11, 2002, a few specimens were observed. These were obviously quite past peak, and

were testimonial that the flight period was ending. The site was visited again on July 15, 2003 which

yielded similar observations of numerous specimens and similar behavior as made July 15, 2001.

These behavior, habitat, and flight period observations of the borealis colony at the Kancamagus

Highway are quite in contrast with those of the more southern anthedon colony at Merrimack. Similar

observations of colonies of borealis have since been made during early July 2003 at Pinkham Notch,

Coos Co., NH at about 2,000 ft. elevation and also in a lush Upper Transition Zone / lower Canadian

Zone habitat near Green River, Guilford Center, Windham Co., VT in mid to late July 2003.

Our conversations revealed that Pavulaan had encountered and collected a small series of

anthedon along the same highway (the Kancamagus) at the Jigger Johnson Campground of the White

Mountain National Forest near a rocky area adjacent the Swift River. The specimens taken at this location

matched specimens of nomi notypical anthedon from Rhode Island, Maryland and Virginia. This location

is only 2 miles east of the borealis colony described above. The habitat (dense, mixed deciduous

streamside forest dominated by firs and sugar maples, but generally with a rather sparse undergrowth

consisting primarily of ferns) and behavior (widely-dispersed individuals perching chiefly on tree trunks

in territorial fashion rather than on ground vegetation) is rather similar to those of the colony at

Merrimack, and is also typical for anthedon in areas farther to the south. These observations would

suggest that the area along the Kancamagus Highway lies within a tension or contact zone between two

populations of anthedon having differing origins. It is also remarkable that two such differing colonies of

E. anthedon , one displaying characteristics and habits of nominotypical anthedon , and another displaying

characteristics of borealis , have been found in such close proximity to one another.

It is interesting to note that Grkovich also visited the site of the anthedon colony at Jigger Johnson

Campground on August 11, 2002 and failed to observe any adult specimens of anthedon , despite

favorable weather and the presence of specimens of borealis at the nearby habitat for that subspecies

earlier that same afternoon.

We have closely observed and documented the behavior of various populations of anthedon at

several widely distributed, more southerly locations. These locations of anthedon include: In Rhode

Island, along the Blackstone River near Quinnville, and the Arcadia Management Area near Escoheag; in

Maryland, Cedarville State Forest near Cedarville, McKee-Beshers Wildlife Management Area near

Seneca, and Great Falls National Park near Potomac; in Virginia, Great Falls National Park near McLean,

Runnymede Town Park in Herndon, Hemlock Overlook Regional Park near Clifton, private lands near

Waterford, and in Shenandoah National Park near Front Royal; and finally, one location in Missouri,

August A. Busch Memorial Wildlife Area near Weldon Spring.

At each of these locations, habitat and adult behavior was remarkably similar, with adults

displaying a preference for dense deciduous woodland with a sparse understory consisting primarily of

non-woody plants and grasses (similar to the habitat of the colony of anthedon observed at Merrimack,

NH). Some of these locations have an almost savannah-like appearance, were it not for the dense tree

canopy which provides total shade, and are generally surrounded by denser woodland. This habitat type

provides a relatively unobstructed view and flight area for the adults. The adults (generally the males)

have been observed perching on bare earth or on tree trunks and were extremely wary of approaching

humans. Their habit of frequently flying from one tree to another and keeping several meters' distance

from nearby adults gives the impression of territorial behavior. Colonies were generally very small

(fewer than five individuals being seen during each visit) and were confined to a small area (generally less

than 30 meters in diameter).

Some colonies seemed to persist in a particular location for only one or two years, subsequently

relocating a year later by a distance of several hundred meters, or being replaced by other colonies in the

general vicinity. But the colony at Quinnville, Rhode Island persisted at the same location throughout the

9

study period. Despite these observations, questions remain over divergent behavioral patterns and habitat

selection in different regions and between some colonies in relatively close proximity to one another, such

as those in northern New Hampshire.

In consideration of the statement of Clark (1936) referred to above, regarding the “ anthedorT from

the “mountains of Virginia”, one wonders how far to the south (or also to the north) the two taxa may,

unbeknownst and undocumented, occur in such proximity to each other while at the same time remaining

distinct. The question of how far south borealis might range in the Appalachians has been responded to

in part by the discovery of an unusual colony of anthedon in the southern Appalachian Mountains on July

27, 2002 by Bruce Grimes and Clyde Kessler. They observed and photographed several “ anthedon ’ (Fig.

9) at Doughton State Park along the Blue Ridge in Alleghany Co., North Carolina. These displayed

considerably more brownish ground coloration on the undersides than is normally seen in anthedon from

that region. This noticeable difference prompted them to reported this finding to us.

Grimes also informed us that individuals were observed displaying what could only be described

as “communal” behavior on a wooded hillside above a boggy area. This behavioral pattern is similar to

that described by Masters (1971) for borealis and to the behavior we have observed for borealis in

northernNew Hampshire. (Interestingly, Satyroides appalachia had previously been observed in this exact

same area. There is no waterway present in the area, and the forest can be described as having some

understory growth, but not dense.) These observations were made at approximately 3500 feet elevation

within generally Transition Zone forest of the southern Appalachians. Our conclusion is that this represents

a mid-high altitudinal colony of borealis far to the south into the range of nominotypical anthedon and

thus has significant implications relative to the taxonomic relationship between borealis and anthedon.

Of similarly great interest to us was the documentation by Grimes and Kessler of nominotypical

anthedon approximately 125 miles to the northeast along the Blue Ridge, at approximately 4000 feet

elevation on Apple Orchard Mountain in Botetourt Co., Virginia. Grimes and Kessler provided

photographic evidence showing the grayish brown ventral surface with the presence of a faint violet tinge

characteristic of nominotypical anthedon.

DISCUSSIONS OF COMPARATIVE PHENOTYPIC CHARACTERISTICS

According to the Original Descriptions (Clark, 1936) for both subspecies as cited above, borealis

differs phenotypically from anthedon as follows:

1. Upper surface darker, with the dark margin of the hind wings broader and more uniform.

2. Lower surface darker and more brownish, usually with the ground color less varied and sometimes quite

uniform, with only faint indications of a narrow lighter line enclosing the rows of spots on the fore and hind

3. On the hind wings beneath, the dark band between the light line is, beyond the fourth and fifth sports,

broader - usually much broader - than the distance between submarginal line and edges of the wing.

An examination of the series of E. anthedon housed in the U.S. National Museum of Natural

History (Smithsonian) collection in Washington D.C. intended to confirm whether the phenotypic

differences cited by Clark (1936) were indicative of true geographic variation (as opposed to mere

individual variation). Careful examination revealed that regional variation does exist (Map: Fig. 10), but is,

by most accounts slight and is initially suggestive of a cline (by evidence of phenotypic characters only).

However, the cline is not continuous, and has some inconsistencies suggestive of post-glacial movements

from several directions. It might be theorized that populations of nominotypical anthedon and borealis

came into secondary post-glacial contact and reintegrated phenotypically at different points in their range.

Yet, in some regions reintegration appears to be incomplete, as evidenced by behavioral and slight

phenotypic differences between nearby colonies, which may also be indicative of early stages of speciation.

10

To supplement Clark's descriptions, we note the following additional characters that distinguish

borealis from anthedon:

1. Upper surface not necessarily darker but with a paler brown cast than anthedorv, anthedon having a more

grayish brown cast. Wing markings more washed in appearance, having less contrast than in anthedon.

Specimens of borealis from Manitoba are generally lighter than anthedon , while specimens of borealis

from eastern Canada and northern New England are at least as dark as anthedon, but with a more brownish

cast. [One factor in the appearance of northern borealis being lighter than anthedon may be in the age of

specimens. Fresh specimens from central Canada were not available to us.]

2. Lower surface of the fore wings displaying considerably more whitish wash exterior to the jagged

postmedian brown band and between the row of eyespots and the wing margin. This character is subject to

great individual variation in both subspecies, but the degree of whitish wash is generally greater in borealis

and lesser in anthedon when comparison is made between series.

3. Lower surface of the hindwings lacks the peculiar violet tinge frequently found in anthedon populations,

especially from those in the southern portion of the species’ range. This violet tinge is most pronounced in

the interspace between the straight postbasal band and the jagged median brown band. In anthedon, the

violet tinge is frequently expressed in the white outer "halo" which encases the row of hindwing eyespots.

4. Borealis appears to be slightly smaller (by 2 mm ) than nominotypical anthedon, on average. A moderate

sample (10 specimens of Canadian borealis and 20 specimens of “true” anthedon) showed the forewing of

anthedon to be roughly 28 mm. in length, while only 26 mm. in borealis.

Based on all of the distinguishing features, examination of the limited series available at the

U.S.N.M. revealed that the nominotypical anthedon phenotype mainly inhabits the southern portion of the

species’ range, primarily the central and southern U.S., southern Appalachian Mountains (with the

exception of the North Carolina colony) and adjacent Piedmont and a narrow area along the coast into

southern New England. A single specimen from Digby, Nova Scotia was of the nominotypical type.

Borealis, in the "true" sense (displaying little tendency toward intermediate characters), ranges throughout

the northern portion of the species' range, primarily in the Great Lakes region, westward to Alberta, and

north to the limits of the species’ range in south-central Canada. Norbert Kondla provided a photograph

of a typical borealis from Elk Island, Alberta (leg. Bob Carrol, July 6, 2001). This specimen appears to

be a darker (more “typical”?) variant of borealis. Specimens assignable to borealis range to the south end

of a broad “tension” zone stretching from northern Minnesota, through Wisconsin, Michigan, eastern

Ohio, through central Pennsylvania, eastern New York and northern New England. Within this tension

zone, borealis, anthedon, and intermediate forms occur.

Masters (1971) concluded that there was a dividing line between both subspecies in the region

immediately west of the Great Lakes that correlated very closely to the approximate boundary between

the Canadian and Transition Zones in that region. This could not be verified from the U.S.N.M. series.

Ironically, the distribution of specimens in the U.S.N.M. series indicated a broad band across Wisconsin

where both borealis and anthedon types occurred, including intermediates. However a reasonably distinct

line of demarcation appears to exist to the east of that region, mainly through Michigan. Interestingly,

this line corresponds to the division between sibling species pairs Pterourus glaucus and P. canadensis,

Phyciodes tharos and P. cocyta “selenis ” [sensu Nielsen, 1999] and also between Celastrina ladon and C.

lucia in Michigan (Nielsen, 1999). While the term "blend zone" might seem tempting, we prefer to refer

to this as a “tension” zone or zone of contact. Interestingly, populations in the southern Great Lakes

region (Michigan and Ohio) and southern Ontario contain a high degree of intermediates which retain the

basic pale brown coloration of borealis but the sharpness and contrast of wing markings that characterize

nominotypical anthedon.

On the other hand, specimens from northern New England and eastern Ontario appear to be slightly

darker and grayer brown like anthedon but with wing markings more washed-out in appearance, as in

“true” borealis. What is more interesting is that no intermediates were examined that occurred south of

11

the tension zone. In other words, intermediates are evident only from areas where borealis occurs. South

of the tension zone, only “true” anthedon types are evident with the exception of the North Carolina

colony. However, north of the tension zone (that is, north of areas where “true” anthedon has been

recorded), intermediates range broadly into the southern and eastern portions of the range of borealis. We

suspect that phenotypic review of such limited series such as those of the U.S.N.M. masks what is truly

happening in nature. Documentation of the field biology of the intermediate specimens is critical to

further study, and is currently unknown to us other than our New Hampshire observations and those of

Grimes and Kessler in North Carolina. Additional and expanded field observations would likely reveal

much more to us about their nature.

Of special interest to us was the zone of contact in New Hampshire. We examined short series of

specimens of E. anthedon anthedon and E. anthedon borealis from the two New Hampshire colonies

described above, in order develop and compare the relative phenotypical characteristics of both.

Reference is made to Figure 1 (Merrimack, N.H., 4 July 2001) and Figure 2 (Swift River, Rt. 112W,

Carroll Co, N.H., 15 July 2001). While slight, we note the following phenotypic differences between

specimens from the two colonies:

1. Overall ground color of both the upper and undersides of the wings are lighter in anthedon than in borealis.

2. Position of the second, small ocellus in Cell M2/M3 of the forewing is positioned more inwardly toward the

discal cell in anthedon than in borealis.

3. While the general lighter area between the postmedian dark line and the row of ocelli of the fore wing is

more extensive and is (perhaps) somewhat more whitish-brown in borealis than anthedon, the narrow

lighter line enclosing the rows of ocelli on both forewing and hindwing is noticeably narrower in borealis

than in anthedon. This is especially noticeable on the hindwing; and there the line is particularly narrow

and faint on the marginal side of the ocelli.

4. The lower ocelli of the hindwing in cells M1/M2, M2/M3, M3/CU1, and CU1/CU2, average slightly smaller

and somewhat less elongate in borealis than in anthedon.

5. The dark lines on the underside are somewhat more extensive and are thicker in borealis than in anthedon.

This is particularly true in the innermost, submedian line extending toward the anal angle of the hindwing;

in some anthedon the line is incomplete and disappears only a short distance beyond the cell, while in

borealis the line is more complete and often is fully connected to the postmedian dark line above the sixth

ocellus.

6. The dark band enclosing the ocelli of the hindwing beneath is broader in borealis, as noted in the Original

Description. However, in our New Hampshire specimens of borealis, the dark band is also broader at each

ocellus and thus across the entire submarginal area of the hindwing, not only at the fourth and fifth ocelli.

The corresponding dark band enclosing the four ocelli of the forewing appears to us to be somewhat wider

in borealis as well.

7. Finally, the ground coloration on the underside of borealis has only a tendency toward a tinge of violet, in

both the lighter and darker areas of the wings. This violet tinge is stronger in anthedon and becomes more

pronounced and “pearly” the farther south it ranges into the southern Appalachian region.

While we anticipate that future examinations of longer series of specimens from these two

colonies will reveal as expected that specimens intermediate in appearance to anthedon and borealis occur

within both, the phenotypic differences between specimens examined from the two colonies described

above, while slight, appear to be rather consistent and are also consistent with the characters described for

anthedon and borealis in their Original Descriptions. These differences are initially less conspicuous than

the well-documented differences between the two in terms behavior and preferences of habitat. We can

perhaps also conclude that had the latter two differences not been noted between individuals of the two

colonies, then the slight differences in phenotypic characters along with the agreement of these color and

pattern characters with the Original Descriptions, might have remained uncovered and thus may have as a

result continued unnoticed.

CONCLUSIONS

Based upon our research and observations documented in part herein, we theorize that populations

of an ancestral Enodia were separated during the final glacial maxima (or perhaps during a previous one),

and as a result these now separated populations embarked onto and progressed along the path toward

speciation. These populations eventually developed on the one hand slight phenotypic differences, but on

the other hand more importantly they also developed significant behavioral differences as well. Full

speciation between the two, however, did not become fully evolved during this period of separation. After

the passing of the most recent Ice Age and the subsequent warming of the climate, the ranges of these

separated populations were extended once again and as a result the two came once again into contact

along a broad front within the Transition Zone (which is in itself a zone of contact or tension, between the

northern Canadian Zone and the southern Austral Zones). This broad front stretches from the Dakotas to

Ohio, and thence eastward into Maine. It is entirely possible that an undocumented second and narrower

tension zone or front between the two extends southward along the higher and cooler Transition Zone

regions of the southern Appalachians as well. The conclusion of the Ice Age would have also interrupted

the process toward full speciation, with the two taxa thus overlapping in what we today observe as the

current tension zone between them.

Under the scenario theorized above, reintegration of the taxa appears to be occurring at this time.

However, secondary contact appears to not have occurred uniformly throughout the broad tension zone, as

evidenced by the differing degrees of intermediacy observed in specimens from the Great Lakes region on

the one hand, and through eastern Ontario and Quebec and into northern New England on the other. As

described above, intermediates occur in Ohio and Michigan, which retain the basic pale brown ground

color or hue of borealis but display the greater contrast in wing markings of anthedon. In eastern Canada

and in northern New England, intermediates occur which possess the darker gray-brown ground color or

hue of anthedon; yet the wing features in these are less contrasty and are more washed-out, as in borealis.

These intermediate forms in general appear to occur well to the north of the northern limit of “true”

nominotypical anthedon phenotypes, where they gradually blend into “true” borealis populations, which

occur in the northern and western portions of the species’ overall range. While these have been

documented as occurring north of the northern range of "true" anthedon populations, we have however

found no borealis or intermediate types south of the tension zone, with the exception of the single high-

altitude colony in western North Carolina discussed above.

While full speciation between the two is not complete, full reintegration is also apparently either

incomplete or inconsistent, as evidenced by known colonies of the two occurring in close proximity to

one another. These colonies, as documented from New Hampshire and North Carolina, occur in differing

habitat and display differing behavioral traits. The fact that a colony referable to borealis persists in

North Carolina appears to us to be evidence of the likelihood that speciation did in fact occur there in its

earliest stages. One might well be left to wonder about the fact that such a unique colony would persist so

far south and yet remain distinct following the most recent glaciation. It is also possible that such isolated

colonies may be eventually doomed to extinction due either to a hypothetical further climate change or by

ultimate reintegration into nominotypical anthedon at some point in the future.

COMMENTS ON TAXONOMIC CONCLUSIONS

A review of subsequent literature following the description of borealis by Clark in 1936 clearly

demonstrates individual author preferences over whether to accept the taxon borealis as either a

subspecies or as a junior synonym of anthedon. This likely reflects either: (1) an individual author’s

personal interpretation of what delineates or defines a subspecies; (2) an author’s personal bias for or

against the naming of phenotypically differentiated populations as subspecies; (3) an author’s personal

13

experience (or lack of) with the taxon in question; or (4) a blind following of, or a reluctance to deviate

from, prevailing or popular taxonomic usage or the prescribed arrangements of popular authors.

We acknowledge that, while the taxon borealis differs only slightly in phenotypic characters from

its sister subspecies, nominotypical anthedon , and that many authors may consider this too minimal to

differentiate subspecies, Masters pointed out that there were physiological differences as well. We

believe that our observations confirm Masters’ conclusions. Some authors might have either overlooked

these physiological differences or simply considered them too minimal also. In the absence of a more

thorough study that examines this issue in much greater depth, we encourage future authors and

listmakers to thoroughly research outstanding issues such as the current one, prior to revising taxonomy

without providing detailed justification. Thus, we retain usage of the original description of borealis by

Clark (1936) as valid, and consider the paper by Masters (1971) as reinforcing the status of borealis at

subspecific rank. The present paper in addition poses the question as to whether primitive speciation

between anthedon and borealis should be further considered. Such issues will require DNA analysis to

fully resolve.

ACKNOWLEDGEMENTS

We thank the following persons for their assistance: Bruce Grimes and Clyde Kessler for their

observations of E. anthedon and photograph of the North Carolina borealis ; John Burns for providing

access to the Smithsonian NMNH collection, Washington D.C.; Norbert Kondla for information on

borealis in Alberta; Richard Green, Senior Reference Librarian (Entomology and Invertebrate Zoology),

Smithsonian NMNH, Washington D.C., for scans of the holotype plates from Clark, 1936.

LITERATURE CITED

Bird, C.D., G.J. Hilchie, N.G. Kondla, E.M. Pike & F.A.H. Sperling. 1995. Alberta Butterflies. Provincial Museum

of Alberta, Edmonton, Alberta, viii + 349 pp.

Brock, Jim P. & Kenn Kaufman. 2003. Butterflies of North America. Kaufmann Focus Guides. Houghton, Mifflin

Company, Boston, Massachusetts. 384 pp.

Brower, A.E. 1974. A List of the Lepidoptera of Maine - Part 1 the Macrolepidoptera. Life Sciences and

Agricultural Experiment Station, University of Maine, Orono, Maine. 136 pp.

Clark, A.H. 1936. Notes on the butterflies of the genus Enodia and description of a new fritillary from Peru. Proc.

U.S. Nat. Mus. 83:251-259.

dos Passos, C.F. 1964. A synonymic list of the Nearctic Rhopalocera. Lepid. Soc. Mem. No. 1.

Ferris, C.D. 1989. Supplement to: A Catalogue/Checklist of the Butterflies of America North of Mexico. The

Lepidopterists’ Society Memoir No. 3. vii + 103 pp.

Forbes, W.T.M. 1960. Lepidoptera of New York and Neighboring States - Agaristidae through Nymphalidae

including Butterflies - Part IV. Memoir No. 371. Cornell University Agricultural Experiment Station, New

York State College of Agriculture, Ithaca, New York. 188 pp.

Handheld, L. 1999. Le Guide des Papillons du Quebec. Volume 1. Broquet, Boucherville, Quebec. 982 pp.

Hodges, R. W. (editor), 1983. Check List of the Lepidoptera of America North of Mexico. E.W. Classey Ltd. And

The Wedge Entomological Research Foundation. London. 284 pp.

Hooper, R.W. 1973. Butterflies of Saskatchewan. Saskatchewan Dept, of Natural Resources and Museum of

Natural History, Regina, Saskatchewan. 216 pp.

Howe, W.H. 1975. The Butterflies of North America. Doubleday & Co., Inc. Garden City, N.Y. xiii + 633 pp.

Klassen, P., A.R. Westwood, W.B. Preston & W.B. McKillop. 1989. The Butterflies of Manitoba. Manitoba

Museum of Man and Nature, Winnipeg, Manitoba, vi + 290 pp.

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

Houghton, Mifflin Company, Boston, Massachusetts, xvi + 349 pp.

Layberry, R.A., J.D. Lafontaine & P.W. Hall. 1982. Butterflies of the Ottawa District. Trail & Landscape 16(l):3-59.

14

Layberry, R.A., P.W. Hall & J.D. Lafontaine. 1998. The Butterflies of Canada. Univ. of Toronto Press, Toronto,

Ontario. 280 pp.

Macy, R.W. & H.H. Shepard, 1941. Butterflies - A Handbook of the Butterflies of the United States, Complete for

the Region North of the Potomac and Ohio Rivers and East of the Dakotas. University of Minnesota Press,

Minneapolis, MN. vii + 247 pp.

Masters, J.H. 1971. A Note on Lethe anthedon borealis (Satyridae ). J. Lep. Soc. 25(4):256-261.

Miller, L.D. & F.M. Brown. 1981. A Catalogue/Checklist of the Butterflies of America North of Mexico. The

Lepidopterists’ Society Memoir No. 2. vii + 280 pp.

Nielsen, M.C. 1999. Michigan Butterflies and Skippers - A Field Guide and Reference. Michigan State University

Extension, East Lansing, Michigan. 248 pp.

Opler, P.A. & V. Malikul. 1992. A Field Guide to Eastern Butterflies. Peterson Field Guide Series. Houghton

Mifflin Company, Boston, Massachusetts, xvii + 396 pp.

Pyle, R. M. 1981. The Audubon Field Guide to North American Butterflies. Alfred A. Knopf, New York. 924 pp.

Roberts, T.S. 1936. The birds of Minnesota. University of Minnesota Press, Minneapolis, MN.

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

Press, Stanford, CT. xii + 583 pp.

Shapiro, A.M. 1974. Butterflies and Skippers of New York State. Search 4(3): 1-60.

Tietz, H.M. 1952. The Lepidoptera of Pennsylvania. Pennsylvania State College, School of Agriculture,

Agricultural Experimental Station. State College, PA. xii +194 pp.

Tilden, J.W. & A.C. Smith. 1986. A Field Guide to Western Butterflies. Peterson Field Guide Series. Houghton,

Mifflin Company, Boston, Massachusetts, xii + 370 pp.

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15

Volume 4 30 April 2004 Number 6

A TAXONOMIC EXAMINATION OF HARKENCLENUS TITUS

(LYCAENIDAE: THECLINAE) IN THE EASTERN UNITED STATES

RECOGNITION AND RESOLUTION OF TAXONOMIC PROBLEMS BY THE

DELINEATION OF OLD NAMES, NAMING OF A LONG RECOGNIZED (BUT HERETOFORE

UNDESCRIBED) SUBSPECIES, AND DESCRIPTION OF A WIDE RANGING NEW SUBSPECIES.

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445-3413

ABSTRACT. Harkenclenus is retained per dos Passos 1970 as a genus distinct from Satyrium. The names titus and

mopsus have long been considered to apply to two subspecies. These are demonstrated to be synonymous as both names are

based on southeastern US populations. The holotype of titus (NHM London) is examined and its type locality established as

Screven County, Georgia. A neotype is established for the synonymic name mopsus, type locality, Screven County, GA. This

leaves the long recognized northern subspecies without a name; it is described as new subspecies Harkenclenus titus winteri,

type locality Sherbom, Massachusetts. Harkenclenus titus campus, type locality Shelby Co., Iowa, is described as a new

subspecies. Subspecies campus has long been recognized as distinct from northern titus winteri, but wrongly assumed to be

subspecies mopsus (=titus). Harkenclenus titus watsoni is assessed and considered valid. Four subspecies of titus are thus

known from eastern North America: H. titus titus, H. titus winteri, H. titus campus, H. titus watsoni. Each subspecies is

phenotypically defined and their ranges delineated. H. t. titus ranges from north central Florida up the coastal plain possibly

to Maryland. H. t. watsoni ranges west of a line from south central Texas north to at least western Oklahoma; its western

boundary in New Mexico is uncertain. H. t. winteri ranges from the Atlantic across the northern US and southern Canada to

subspecies immaculosus', not in southern Appalachians. The range of H. t. campus extends from the Black Belt prairies of

Alabama to eastern Texas and north and northeast to the ranges of H. t. immaculosus & H. t. winteri.

Additional key words: Evolutionary relationships H. titus occidentalis, Floridian relict.

HISTORY AND PROGRESSION OF THIS PROJECT

The North American butterfly commonly known as the Coral Hairstreak was described in 1793 by

Fabricius as Hesperia titus. It is a distinct part of the North American fauna and easy to identify. The only

modem taxonomic debates have been over what genus it is best placed in - Satyrium Scudder, 1876, or

Harkenclenus dosPassos, 1970, and the periodic questioning of “in Anglia” =Newfoundland as the type

locality (Miller & Brown 1981). Because Newfoundland has been given in the literature as the titus type

locality, the nominate subspecies has long been accepted as the ventrally moderately marked and rather un¬

contrasting populations from southeastern Canada into Manitoba, and the northeastern US south to Virginia

and west to the east slope of the Colorado Rockies (Ferris & Brown 1980, Layberry et al. 1998).

Hubner described Chrysophanus mopsus in 1818 with “Georgia in Florida” as the type locality.

The name mopsus has long been applied to all titus from Georgia t> Texas, and north to Virginia and

Colorado (e.g. Klots 1951, Ferris & Brown 1980). There are some exceptions, as Scott (1986), pg. 360,

who attributes all eastern population to H. t. titus and all western populations to H. t. immaculosus (W.P.

1 Research Associate, FSCA, Gainesville, Florida. Staff Researcher, TILS, Goose Creek, South Carloina.

Fig. 1. Jones’ 1785 leones 9 titus. Fig. 2. Hiibner’s 1818 d Chrysophanus mopsus. Fig. 3. Fabricius’ 1793 9

Hesperia titus holotype (NHM, London). Fig. 4. H. titus holotype labels. Fig. 5. 9 C. mopsus neotype (data in text).

Fig. 6. 9 H. titus, 6 June 1992, nr. Jet. 3 & 394, Orangeburg Co., SC. Fig. 7. d H. titus, 4 May 1975, nr. Torreya,

Liberty Co., FL. Fig. 8. d H. titus, 13 May 2002, Axon Rd., Orangeburg Co., SC. Figs. 9-10. d holotype. H. titus

campus (data in text). Fig. 11. d paratype. H. t. campus, 5-10 June, vise. Clinton Lk., Douglas Co., KS (leg. Adams).

Fig. 12. d Paratype. H. t. campus. 2 June 1975, Athens, TX. (leg. Bordelon). Fig. 13. d H. t. campus. 24 June 1991,

8 mi. n. Walker, Ellis Co. KS (leg. Kuhn). Fig. 14. d H. t. watsoni. 11 May 2003, nr. Mendard, Menard Co., TX. (FW

17 mm. leg. Kuhn). Fig. 15. 9 H. titus. ex. pupa 8 April 1993, Wakulla Springs, Wakulla Co., FL (leg. Slotten). Fig. 16.

9 Allotype. H. t. campus (same data as 11). Fig. 17. 9 Paratype. H. t. campus, 1 June 1945, Faulkner Co., AK (no.

leg). Fig. 18. 9 H. t. campus, 8 July 1980, 10 mi. east Nobel, Cleveland, Co., OK. (leg. Davenport). Fig. 19. 9 H. t. nr.

watsoni (same data as 18). Fig. 20. d Paratype. H. t. watsoni, Kerrville, TX (AMNH). Fig. 21 d holotype. H. t.

immaculosus, July, Provo, UT (AMNH). Fig 22 allotype. H. t. immaculosus (same data as 21). Fig. 23 D/V

d holotype. H. t. winteri (data in text). Fig. 24 D/V 9 Allotype, H. t. winteri (data in text). Fig. 25 d paratype. H. t.

winteri (data in text). Fig. 26 9 Paratype. H. t. winteri (data in text). Fig. 27 d H. t. immaculosus, 2 August 1952,

6,400’, Warm Springs Rd., nr. Sun Valley, UT (no leg). Fig. 28 9 H. t. immaculosus (same data as 27). All leg.

Gatrelle except as noted. 13-14 same photo/lighting: by Kuhn. AMHN watsoni and immaculosus photos, D. Wright.

Photo of titus holotype, NHM staff. 18-19 photos, D. Walker. All other photos by Joseph Mueller. Not to scale.

Comstock, 1913). The only other described subspecies in the eastern area is watsoni Barnes & Benjamin,

1926. Throughout the literature, subspecies watsoni (when recognized as valid) has been attributed to the

area of south central & west Texas, north to southwest Oklahoma, possibly extreme southeast Colorado and

west into New Mexico. All of tiis appears to render all the subspecific eastern taxa of titus as well

understood entities with stable nomenclature. However, the examination of this taxon has shown that it is

both a nomenclatural and taxonomic quagmire.

My field experience with the Coral Hair streak is in Iowa, Florida, Georgia and South Carolina.

The significance of this, is that these populations are in taxonomically unique and rarely sampled regions.

In Iowa, two phenotypes were found; one was what has been conventionally known as titus titus in

northeastern Iowa (Howard County, 1975); the other was in central Iowa (Johnson County, 1975), west

Iowa (Shelby County, 1967), and southwest Iowa (Guthrie County, 1975) which fell within the literature

concept of subspecies mopsus in the western prairie region. True mopsus was collected in Liberty County,

FL in 1975, in south coastal South Carolina (Aiken & Orangeburg counties: 1976, 77, 88, 90, 91, 92, 98,

2002), and most importantly, in 1994 at its type locality in both Screven and Burke counties, Georgia.

These specimens from the Midwest and southeast indicated that the conventional application in the

literature of the name mopsus to western and Midwestern populations was incorrect. Thus, about 1977, a

taxonomic assessment of titus east of the Rocky Mountains was undertaken and by 1980 series from the

above locations demonstrated that these prairie “mopsus” populations were either: 1) an undescribed

subspecies or 2) perhaps an expression of H. titus watsoni as described from south central Texas - because

they were not mopsus.

A search began for specimens of watsoni for comparison. This took many years as I was not able to

travel to those museums which housed type material, was unable to obtain the loan of these specimens, and

was not able to find any in private collections until 2002. However, this potential watsoni specimen was

from Oklahoma, and Texas topotypes were needed for unequivocal determination. This resulted in periods

of years of little activity re titus. With the inability to determine just what watsoni was, the effort to reach a

taxonomic conclusion on the western and Midwestern prairie populations became a dead end.

However, from 1980 to 2001 some significant information was obtained. 1) A visit in 1994 to

Mississippi State University, and an examination of its collection, revealed that Mississippi titus were of

the same phenotype collected in southwestern Iowa, and thus, what the literature incorrectly called

“mopsus” in the prairie states. During that visit, knowledge of the Black Belt Prairies of Mississippi and

Alabama was acquired and why the Mississippi titus are of western prairie affinity and evolutionally

disassociated with the titus of the southeastern coastal plain. 2) Specimens of titus in the FSCA collection

Gainesville were examined and 3) specimens borrowed from private collectors. The examination of these

specimens showed 1) that mopsus is restricted to the Southeast from north Florida up the eastern seaboard

into Virginia and 2) that all the prairie populations from Mississippi to Texas, Colorado, and central Iowa

were the same taxon and lacked a name - unless they were broadly referable to watsoni.

In 2001, this project changed dramatically. To that point, the research was only seeking to determine

the taxonomic placement of prairie titus - was it an undescribed taxon or did it belong within watsoni.

While acquiring copies of original descriptions in general, the original descriptions of titus and mopsus

were obtained. It was immediately evident that a big problem existed. The figure on which the name titus

was based (fig. 1) was clearly the southeastern phenotype - traditional mopsus. Further, the OD

illustration of “mopsus” (fig. 2) was intermediate to the northern phenotype associated with the name titus.

This problem was confirmed in 2002 through correspondence with Harish Gaonkar (page 4) who stated that

the actual type locality of titus was not “Newfoundland” but Georgia! Hlibner had restricted the type

locality of the name mopsus to “Georgia in Florida” in his original description of that taxon. All of this

renders the name mopsus a junior synonym of titus with both names applying to the subspecies of the

southeast and eastern seaboard with ventrally dark grayish brown wings, small black spots, reduced black

spotting on the VFW (esp. of males), and with bright white halos.

3

TITUS TAXONOMY

No type or syntypes exist for mopsus as all American Hubner types are lost or destroyed 2 . The

holotype of titus exists in the British Museum of Natural History (fig. 3). Because the names titus and

mopsus are based on the same southeastern subspecies, this leaves the northern and northeastern subspecies

(traditionally assumed to be titus titus) without a name - having never been described.

In 2003, photographs of a fresh nearly topotypical male watsoni were obtained (fig. 14). This

confirmed that the wide ranging prairie populations are not related to watsoni and that watsoni is indeed a

unique and rare southwestern taxon.

The background research has unveiled two matters that need to be addressed and resolved. One is

the matter of the wide ranging prairie populations that have long incorrectly passed as subspecies mopsus.

This situation (the original area of investigation) is easily taken care of by the description and delineation of

this prairie entity as a new subspecies, which is accomplished herein. The second, and paramount matter,

is having found that titus and mopsus were actually proposed for the same zoological taxon, which negates

the long standing nomenclatural application of these names by synonymizing the name mopsus and leaving

the northern subspecies without a name. Regarding the mopsus!titus synonymy, there is no easy solution.

The two courses of action relative to the names titus and mopsus are these:

1. Ignore the type specimen of titus, both ODs, the historical facts about the type localities and

leave the names titus and mopsus in application (usage) as they have been. Not simple. To do this, an

appeal would have to be made to the International Commission of Zoological Nomenclature to set aside a

valid holotype and ignore all historical data and erect a neotype and type locality for titus that would be

historically and scientifically fiction. Not only is titus not known from Newfoundland, it does not range

anywhere near there (Butterflies of Canada, 1998, page 132 map). What would be the advantage of this?

Maintain popular usage? North American butterfly hobbyists seldom utilize scientific names and

overwhelmingly choose to just call all titus subspecies the “Coral Hairstreak”. Further, most hobbyists

Guides use only binomials. Thus, maintaining current usage would sacrifice science for a trinomial

nomenclature seldom even utilized by recreational lepidopterists.

2. Maintain the historical and ICZN Code compliant synonymy and erect a new name for the

heretofore undescribed northern subspecies. What would the disadvantage of this be? Professionals and

taxonomists would need to adapt to using a different set of trinomials for this group of subspecies.

The advise of the taxonomists who reviewed this paper was to implement solution two.

Harish Gaonkar is generally accepted as the expert on Fabricius and his work. Here is his email of

November 12, 2002. (I have put some key words in bold.)

“The following notes may be useful on Fabricius's sources in Britain and in Denmark. I am afraid the

problems with Fabrician types are not as neat as they some times appear to be...

1) Hesperia titus Fabricius, 1793.

F described the species from the Collection of Dm Drury and stated that it was, “in Anglia Dom.

Drury...” Fabricius would have studied this material either in 1787 or in 1790. The locality given by Fab

was obviously wrong, because Dmry often did not know from where many insects came from, although

Dr. Gerhard Tarmann of Innsbruck, Austria, relayed the following information in 1998: “There is some Hubner material

there although most of Htibner’s material was destroyed by a fire. There was a man called Mazzola who bought some of

Hubner’s original material. As this man has taken away all of Hubner’s labels and replaced them with his own printed labels,

it took years to find out that some of the material in Mazzola’s collection is in fact original Hubner material. However,

although the Mazzola collection is in Vienna, there is only European material involved. There are no possible Hubner types of

American butterflies existing. I got this inf ormation from Dr. Sabina Gaal, Naturhis-torissches Museum Wien (NHMW).”

, although I can not look it up in Copenhagen. Fab also

i. When the Drury Collection was sold in P 1804 (or 1805) many type material described by

Taken ah in all - with the external evidence etc. that I know -1 tt

imitted by

il to J. E.

ly do as you think is fitting.”

November 13, 2002 as

it 650 butterflies, out of v

1) Hesperia titus 1793: p. 297, no. 130. Accordin

us, 130”, which may be the one written by Drury hi

I confirm that there are NO specimens of this [titi

volumes of the leones for the first tir

the LECTOTYPE. As I said, adding two and two toget

supposed to curate. So the actual specimen of “thus” m

Vienna, were destroyed by fire in 1848! So forget about

e 1787 by Jones. That is

d on the TYPE then in the Drury

Hubner’s name was synonymized by Butler (Catalogue of the Diurnal Lepidoptera described by

Fabricius in the Collection of the British Museum, [1870]: 191)

3) John Abbot.

Most of John Abbot's drawings and paintings of North American Insects (also butterflies) are held by

the Entomology Library in the BM.

Although I have not done careful research, I have a "feeling" (not subjective though) that his specimens

are in the Linnean Society Collection, London. And why do I think that? Some years back, I helped two of

my colleagues Martin Honey and Malcolm Scoble on Linnean Butterflies. It appeared to me then (and now)

that some of the specimens in LS Collections were added later by Sir James E. Smith to the Linnaean

Collection, particularly from North America. We know that Smith received a lot of material from your area

(first from Virginia and later from Georgia) sent by Abbot. William Jones also got some specimens, and

so did Dru Drury. So, I think (in many cases know) that specimens sent by Abbot to these three are to be

searched in the Linnaean Collection (LS) AND Hope Dept, of Entomology, University of Oxford.

The LS now houses many specimens that were used by Jones for his leones and described by

Fabricius directly from the illustrations themselves. Oxford contains many specimens of William Jones

himself which came there through many others, among others through the John Francillon Collection.

Harish”

The pertinent facts from the above are these. 1) The published type locality of “in Anglia Dom.

Drury” is “obviously wrong” in Harish’s expert opinion. 2) It is his view that the specimen the name titus

was based on came from the “southern parts of the USA” and likely from John Abbot ex “Virginia and later

from Georgia". (Abbot moved from Virginia to Georgia in 1775). 3) The type was illustrated by Jones in

his leones and exists in the main collection of the British Museum.

Contacting the NHM, London confirmed that the type is there and in the Lycaenidae drawer as

Harish stated (fig. 3). This specimen is here determined to be the holotype by monotypy. A comparison of

this female specimen with the Jones leones’ female illustration shows they have the same markings and spot

positions. The titus holotype is typical of southeastern females traditionally known as subspecies mopsus

of Hubner (TL Georgia in Florida). No other specimens were figured or mentioned, thus there are no

syntypes. A holotype label has been sent to Kim Goodger, NHM, for placement on this specimen.

Butler (1870) synonymized the names titus and mopsus as below. While correctly recognizing

that the taxa represented by these two names were synonyms, he was incorrect in assuming New England as

the habitat. Specifically, because Hubner had stated that Georgia was the habitat of mopsus - and because

the titus holotype is clearly the southeastern phenotype. Thus, this paper is not the first to state that the

names are synonymous. It is the first to determine and that the region inhabited by both these taxa, based on

the proceeding data, is coastal Georgia.

3, Strymon Titus.

Hesperia (E.) Titus, Fabricius f “ alia integemmis, fuscis imma¬

culate ; postids subtus ocellatis strigaque postiea maculari fulva:

habitat in Anglia." S. Mopsus, Buhner, Rvot, BchmtL Outrage, figs,

135,133 (1S0G),

Fabricius, EnL SysL ill. p. 297. n. 130 (1793).

North America, (Probably from Mr. Milne’s collection) B.M,

The specimen of P. Titus in the National Collection bears an old

label, on which is written “ Titus, 130,” evidently a reference to the

description by Fabricius. It is quite possible that this is the typo

from Drury’s collection, received through Mr. Milne, as it answers

in ovpj-v parriculai' io t!>*■ «]'>Lri])1i<Mi. and has jliogotlier the up-

pcarance of a very old specimen; its true habitat appears to be New

England.

6

TYPE LOCALITIES

(TITUS)

The phenotype of the titus holotype is Southeastern. Harish Gaonkar states that he considers it likely this

type specimen originated from John Abbot from either Virginia or Georgia. It is considered unlikely that the

specimen originated from Virginia because 1) Abbot only resided in Virginia from 1773 to 1775 when he relocated

to Georgia due to the impending American Revolution (Harris, 1972), 2) Fabricius likely saw the specimen

“...either in 1787 or in 1790” and published the name in 1793 - 18 years after Abbot left Virginia, and 3) Abbot

did not figure titus for the 1797 book on the Lepidoptera of Georgia by J.E. Smith which makes it possible he did

not even discover titus in Georgia until after 1797 - after Fabricius’ description of titus (which would mean this is

not an Abbot specimen). However, we know Abbot found titus in coastal Georgia as his painting of it was

published in Boisduval and LeConte, 1833 (http://www.sc.edu/libraiy/spcoll/abbot/part_ l.html Image 11, and

historical notes. University of South Carolina (USC) Thomas Cooper Library web site). This painting depicts a

typical male and female of the Georgian phenotype. The issue of Virginia as the potential IL is important as some

populations there are likely not no min ate titus, but intermediates.

Fabricius’ OD citation of “in Anglia” (Newfoundland in the literature) is false. This is confirmed by the

fact that H. titus has never been recorded anywhere near Newfoundland (Butterflies of Canada, 1998). Butler’s

rendering of “in Anglia” as “New England” is literal but, as Harish noted, an error. Recommendation 76A.2 of the

ICZN Code says: “A statement of a type locality that is found to be erroneous should be corrected.” The type

locality of Hesperia titus Fabricius, 1793 is here corrected to: Millhaven Plantation, Screven County, Georgia.

This is because Abbot collected on this Plantation (Harris, 1972) and I have found it still there today. Abbot lived

in adjacent Burke County (1776-1806) until he was 55 and moved to Savannah (see above web site), found many

of his taxa in these counties, and Harkenclenus titus still occurs in both counties today.

(MOPSUS)

Hiibner established “Georgia in Florida” as the TL of mopsus. At that time, “Georgia in Florida” meant

Spanish (coastal) Georgia. At one time this region extended from Beaufort, SC south into Florida. In the early

1800’s, this was a commonly used phrase to denote coastal Georgia. The specimen from which Hiibner described

mopsus was obtained from a Dr. Andersch (see mopsus OD below). Martin Spies provided the following from a

German web resource (http://www.zalf.de/deid/index.htm) and suspects this may be Hiibner’s Andersch: “Johann

David (or Daniel) Andersch (1768-1847), died in Tilsit (now Sovetsk in the Russian enclave between Poland and

Lithuania that also contains Kaliningrad”. Spies states, “There's no indication in the above that he lived in the

U.S.” He considers it likely that Andersch was a wealthy Dr. and “.. .private collector who acquired material from

all over the world.” Andersch is frequently mentioned by Hiibner, 1818, and it is reasonable to conclude

Andersch was a customer of the European agents who sold Abbot’s specimens. It is also possible Andersch also

bought Abbot art - perhaps the veiy painting for Hubner’s mopsus plate because the Hiibner figures of mopsus

are identical to an Abbot painting at the USC Thomas Cooper Library (image 13). This Abbot figure even has the

name “mopsus” (1818) penciled beneath it. This male painting has heavier spots than the # 11 image that was the

Boisduval and Le Conte plate original. The Hiibner painting is quite atypical.

It is wise to delimit the type locality cf mopsus by designating a neotype to eliminate all ambiguity and

possible future instability relative to the names titus and mopsus in accordance with the ICZN provisions laid out

in Article 75.3 and subsections. All provisions of 75.3.1, 3.2, 3.3, 3.4, 3.5, 3.6 and 3.7 are here considered quoted

by this reference and specifically addressed and met within the applicable sections of this paper. The type locality

of Chrysophanus mopsus Hiibner, 1818 is here clarified by neotypification (fig. 5) to Millhaven Plantation,

Screven County, Georgia. Neotype deposited in the NHM (London) and bears the following labels. A red label,

hand lettered “NEOTYPE: Chrysophanus mopsus, Hiibner 1818”; small white label, hand lettered “9 H. t.

mopsus ” and type set print “Ronald R. Gatrelle, COLLECTOR”; a medium size white label hand lettered, “June 9,

1994, Screven Co., GA, M illh aven Plantation, Brier Creek”. All words in black ink.

GENERIC PLACEMENT

The taxon titus was originally described in the genus Hesperia (now restricted to skipper butterflies). The

next oldest name is mopsus described in Chrysophanus, which was suppressed by the ICZN in 1959. C.F.

dosPassos (1970) proposed Harkenclenus as the replacement name for Chrysophanus with mopsus as type

species = subspecies titus. DosPassos treatment was very brief and is as follows.

On page 28.

HARKENCLENUS nom. nov.

pro Chrysophanus Hiibner, 1818 (opinion 541, name 1235)

Type: Chrysophanus mopsus Hiibner, 1818 (opinion 541, name 1235)

(= Papilio titus Fabricius, 1793) (opinion 541, name 1605)

On page 36.

The International Commission on Zoological Nomenclature by opinion 541 suppressed among other names

Chrysophanus Hiibner, 1818, and placed it on the Official Index of Rejected and Invalid Names in Zoology for the

purpose of the Law of Priority but not for those of the Law of Homonymy. Consequently, a replacement name is in

order. For that purpose Harkenclenus has been chosen, being an arbitrary combination of the first syllables of the

name of my friend and colleague, Harry Kendon Clench. The new name is masculine.

Clench in Ehrlich & Ehrlich (1961) was the first to begin combining other genera into Satyrium Scudder,

1876. In his 1961 assessment, Clench not only determined Chrysophanus titus was not a Satyr ium, but that it

was monotypic (page 191). In 1978, Clench again assessed various world genera and synonymized many genera

(and thus species) into Satyr ium - but not genus Harkenclenus (1970) nor species titus. His analysis continued

his 1961 position that titus was not a Satryium (delimited species on page 281). Lafontaine (Butterflies of Canada,

1998, page 25) proposes the inclusion of titus in Satyrium and presents data in support of this. In the last 7 years,

most lists and books have placed the taxon titus in the genus Satyrium. The assertion of this paper is that there is

insufficient criteria to place titus into Satyrium. Until more genus level in-depth research is presented specific to

titus , it is deemed best to leave the species titus in Harkenclenus per dos Passos 1970 - and several other

publications between 1970 and today. The perspective here is that this also serves to keep synonymization to a

minimum. Retaining titus in Harkenclenus is the most conservative position.

(In review, this section was discussed and various versions considered. Because the primary objectives

of this paper are to 1) properly define the names titus and mopsus according to the type specimens and historical

record and 2) assess the other regional populations from that base, the consensus was that the paper is best

served by limiting this section to basic information and author’s opinion because the generic placement issues are

outside the paper’s primary purposes.)

TAXONOMIC DESCRIPTIONS

Harkenclenus titus titus (Fabricius, 1793)

Original Description

Latin (Transcribed as in original.)

130. H. R. alis integerrimus fuscis immaculatis: Titus. 130.

posticis subtus ocellatis strigaque postica ma-

culari fulva.

Papilio Titus. Jon. fig. pict. 6 tab. 44. fig. 2

Habitat in Anglia Dom. Drury.

Statura omnino praecedentium. Alae omnes

supra fuscae, immaculatae. Subtus itidem

fuscae, anticae striga postica e lineolis albis

nigrisque, posticae lineola media strigaque e

punctis nigris, albo cinctis. Versus margi-

nem maculae rufae, puncto nigro notatae.

English Translation (By: Dr. Rienk de Jong, NMHN, The Netherlands.)

H[esperia] RJurales] entire wings dark and without spots: Titus

underside of hindwings with eye spots and a discal series

of tawny spots.

Papilio Titus. Jon[Jones] fig. pict. 6 tab. 44. fig. 2

Lives in Anglia Dom. [N. American English territory] Drury.

With general build of foregoing species [artaxerxes]. All wings

dark on upperside, without spots. Underside equally dark,

forewings with a distal series of white and black short lines,

hindwings with a central short line and a series of black,

white-ringed specks. Toward the margin reddish-brown

spots, marked with a black speck

10

County: no location, 1?, 1 June 1945 (leg. unknown). TES

Bordelon). The holotype is deposited in the Museum of the

10): IOWA: Shelby County, vise, of Defiance, 8 July 1967 (leg. R. Gatrelle). Allotype

)f Defiance, Id 1 , 8 July 1967; Guthrie County: vise. Sheader Prairie, 4dd, 1 9,28 July

airie, Id, 28 June, 2d d, 1 9, 2 July, 19,3 July 1975 (all leg. R. Gatrelle). KANSAS:

rson County: Athens, Id, 2 June 1975 (leg. C.

se Creek, SC. C

s: 3 d &

(Ferris & Brown, ^ 981), Butterflies and Moths of Missouri (Heitzman &

id east of Ohio. One A

, It is not known if, or

‘d in Butterflies of Georgia (Harris, 1972)

ia is unknown to me. (S

i. If this is so, it in

at the range of campus are qi

s. 1) The range of winteri is al

le & Oak forests ofth

.rida anefup the Sandhills region of Georgia, South Ca

;welve 9 cotypes. 12 fir¬

es of the Eastern United States” al

m of S[trymon]. titus is to be found in *

is. (Figs. 21-22 & 27-28.) This ta

by the extensive amount of red (fulvous) scaling that is fr

5) on plate 52 figures 27 & 28 depicts beautiful examples

a. It flies in one brood from early Jr

Map A. USGS range map. Historical site records by county and state geopolitical boundaries. Map B. Geological relief

map. Subspecies’ ranges overlaid to biogeographical regions. Stars indicate type localities; lines, range limits. Black: H.

titus titus. Encircled Appalachian areas are where titus is rare or restricted by high elevations. Southwest and northeast

range/blend zones undetermined and indicated by open line. Red: H. titus winteri. Southern boundary corresponds

closely to glacial limits. Western transition to campus and immaculosus needs determination. Lavender: H. titus

campus. Northwestern, southwestern and northeastern limits/transition areas need verification. Green: H. titus

immaculosus. Eastern area of transition with campus and winteri needs to be worked out. Absent in many areas within

the range of subspecies. Light yellow: H. titus occidentalis. Northeast California and northwest Nevada. Only in

mountain ranges in this region. Yellow: H. titus watsoni. Northern limit/blend with campus undetermined.

14

CONCLUDING REMARKS

During the review of this paper, David Wright informed the author that he had photos of type material

of watsoni (fig. 20 paratype male) and immaculosus (figs 21-22 holotype and allotype) in the American

Museum of Natural History (AMNH). The addition of these adds greatly to the substance of this paper.

Also, just prior to going to press, Ken Davenport emailed photos of three female titus he had collected

east of Nobel, Cleveland County, Oklahoma, 4-8 July 1980 (figs. 18-19). These are significant specimens

as they are from the area where watsoni and campus meet. Fig. 18 is a typical campus while fig. 19 is near

watsoni with reduced and yellow VHW marginal spots. The Canadian River is a geological feature which

seems to be the basic boundary between campus and watsoni in Oklahoma.

The ranges of the titus subspecies differ greatly (Maps pg. 14). H. t. occidentalis has the smallest

range and is the rarest of the subspecies. The next smallest range is that of H. t. watsoni , after that H. t.

titus , H. t. winteri , H. t. immaculosus and H. t. campus . None of these subspecies are considered

“common”, although they may be locally not uncommon at the right time and place.

John Hyatt (recent pers. comm.) stated he has “... found none from localities above about 1400-1500’,

or below 1200” in eastern TN. Together with the USGS range maps and the author’s extensive collecting in

western NC above 1500’, titus appears to be absent from the southern Appalachian Mountains. If this is so,

an extensive north south barrier exists between those population east and west of these mountains - and

apparently as far north as Pennsylvania.

It is possible that nominate titus ranges through north GA and around the Appalachian Mountains in a

narrow band up into extreme eastern TN. This is because Hyatt has determined the titus in east TN as

“mopsus” and James Adams (pers. comm.) has stated the titus in northwest GA are also the “mopsus”

phenotype. However, I have not seen specimens to confirm either of these determinations. Thus, they may

be intermediates or subspecies campus.

Having lived in Pensacola, Florida, It is probable that titus titus may inhabit the sub-Appalachian hill

country of Escambia County in the Cantonment area and into southeast Alabama. The taxon in the Black Belt

of Alabama should be campus.

In the southeastern US, the author’s experience is that unless Asclepias tuberosa is found, especially

subspecies rolfsii (Vail) Woodson (Butterfly-weed), the Coral Hairstreak will seldom be located. Thus,

not finding this taxon does not mean it is not at a location, just that it is very difficult to find unless the

proper nectar source is present. Then again, this taxon may be so selective about nectar sources, that even

where the larval host is abundant, titus will be absent without these resources. If this is so,

conservationists need to support the availability of these “weedy” Milkweeds in areas where titus taxa are

known to occur.

Unfortunately, it was not noted, nor now remembered, on what flowers campus was found at Sheeder

Prairie in Guthrie County, Iowa. The following uncommon species were found with campus at this location:

Satyrium acadica (W.H. Edwards, 1862), Euphyes bimacula illinois (Dodge, 1872) and E. dion ssp.

(W.H. Edwards, 1879). In the one visit to this site in 1975, it was a small hillside prairie and rather

invaded with weeds and surrounded by agriculture. It is unknown if the site is still extent, and if so, if any

of these taxa are still found there. When the campus type was taken, the author resided in Defiance, Iowa.

A detailed notation was not placed on that specimen as to where it was caught. The recollection is that it

was (in 1967) not far north of Defiance along Hwy. 59, east side of road, at the edge of a wooded area

going up a hill. S. calanus falacer (Godart, 1824) was also found at the type locality.

Norbert Kondla brought two 2003 publications to the author’s attention during review. One on the

Butterflies of North Dakota (Royer) and one on South Dakota butterflies (Marrone). The photos in these are

in stark contrast. The North Dakota individuals are very near immaculosus , if not that taxon, while the

South Dakota individuals are boldly marked campus. The Dakotas need a subspecific range analysis for

titus. Kondla considers the titus in south west Canada also in need of more taxonomic work.

15

ACKNOWLEDGEMENTS

I thank the people who provided advise on and/or reviewed this paper: David Wright, Norbert Kondla, Don

Lafontaine. I also thank: the Photo Unit at The Natural History Museum (NHM, London) for the photo of the

holotype of titus; Kim Goodger, Curator of Butterflies, NHM (London) for locating the type and facilitation; Harish

Gaonkar, Denmark, and the NHM (London) for the essential historical information on Fabricius and the titus type

specimen without which this paper could not have been done; Dr. Rienk de Jong, Department of Entomology,

National Museum of Natural History, Leiden, The Netherlands for translation of the titus OD; Martin Spies,

Zoologische Staatssammlung, Munich, Germany for the translation of the mopsus OD and general linguistic

consultation; David Wright, Norbert Kondla, Don Lafontaine for providing copies of ODs and other documents;

Ricky Patterson for loan of Mississippi specimens; Charles Bordelon for the loan of Oklahoma, Texas and Arkansas

specimens; Jeremy Kuhn for photos of topotypical watsoni’, James Adams for north Georgia phenotype information,

and John Hyatt for east Tennessee range data; Joseph Mueller for photography; and Ken Davenport and David

Wright for additional photos of Oklahoma and type specimens respectively.

LITERATURE CITED

ALLEN, T.J. 1997. The Butterflies of West Virginia and Their Caterpillars. Pittsburgh, PA: University of Pittsburgh

Press, xii + 388 pp.

AUSTIN, G.T., and J.F. EMMEL. 1998. New Subspecies of Butterflies (Lepidoptera) from Nevada and California.

Pages 501-522 in: Emmel, T.C., editor. Systematics of Western North American Butterflies. Mariposa Press,

Gainesville, Florida. 878 pp.

BARNES and BENJAMIN. 1926. Xerox copy of Original Description of Strymon titus race watsoni in: Bulletin of the

S. California Academy of Sciences, 25: 94 (1926).

BROWN, F.M., D. Eff & B. Rotger. 1957. Colorado Butterflies. Proceedings, Denver Museum of Natural History.

Denver Colo. 369 pp.

BUTLER, 1870. Catalogue of the Diurnal Lepidoptera described by Fabricius in the Collection of the British

Museum. Digital Scan of 191.

CLARK, A.H., and L.F. CLARK. 1951. The Butterflies of Virginia. Smithsonian Misc. Coll. 116 (7): vii + 239 pp.

CLENCH, H.K. 1961. Tribe Theclini. pp. 177-220, in: P. R. Ehrlich & A. H. Ehrlich, How to Know the Butterflies.

Dubuque, Iowa; Wm. C. Brown Company: [vii] + 262 pp, 525 figs.

_. 1979. The Names of Certain Holarctic Hairstreak Genera (Lycaenidae). Journal of the Lepidopterists'

Society 32(4): 277-281

COMSTOCK, W.P. 1913. A New North American Butterfly in the Family Lycaenidae. Bulletin of the Brooklyn

Entomological Society 8:3. pp 33-36.

DOS PASSOS, C.F. 1970. A Revised Synonymic Catalogue with Taxonomic Notes on Some Nearctic Lycaenidae.

Journal of the Lepid. Soc. 24:1, pages 26-38.

EBNER, A.E. 1970. Butterflies of Wisconsin. Milwaukee Public Museum. 205 pp.

FABRICIUS, J.C. 1793. Entomologica Systematica Emendata. III. Hafniae. Xerox copy of 3:297.

FERRIS, C.D. & F.M. BROWN 1980. Butterflies of the Rocky Mountain States. University of Oklahoma Press,

Norman. 441 pp.

HARRIS, L., Jr. 1972. Butterflies of Georgia. University of Oklahoma press, Norman OK. 326 pp.

HEITZMAN, J.R., & J.E. HEITZMAN. 1987. Butterflies and Moths of Missouri. Missouri Dept, of Conserv.,

Jefferson City, MO. 385 pp.

HOWE, W.H. 1975. The Butterflies of North America. Doubleday & Co., Inc. New York, NY. 633 pp.

HUBNER, J. 1818. Zutrage Sammlung exotischer Schmetterlinge. Augsburg. Xerox copy of 1:24 and figs. 135-136.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International Code of

Zoological Nomenclature, Fourth Edition (1 January 2000). London, UK: The International Trust for Zoological

Nomenclature. 306 pp.

IFTNER, D.C., J.H. SHUEY & J.V. CALHOUN. 1992. Butterflies and Skippers of Ohio. Ohio St. Univ. (in co-op

with) Ohio Dept, of Nat. Res. Div. of Wildlife and Ohio Lepid. Columbus, Oh. 214 pp.

JONES, W. 1785. leones: Papiliones Nymphales, etc. Digital Scan of titus plate.

KLASSEN, P., A.R. WESTWOOD, W.B. PRESTON, W.B. McKILLOP 1989. The Butterflies of Manitoba. Man.

Mus. of Man and Nature, Winnipeg, Man., Canada. 291pp.

16

KLOTS, A.B. 1951. A Field Guide to the Butterflies of Eastern North America. Peterson Field Guide Series.

Houghton, Mifflin Company, Boston, Massachusetts, xvi + 349 pp

LAYBERRY, R.A., P.W. HALL, AND J.D. LAFONTAINE. 1998. The Butterflies of Canada. Toronto, ON:

University of Toronto Press. 280 pp.

MARRONE, G.M. 2003. Field Guide to Butterflies of South Dakota. South Dakota Dept, of Game, Fish and Parks.

478 pp. Scan of titus section.

MILLER, L.D., AND F.M. BROWN. 1981. A catalogue/checklist of the butterflies of America north of Mexico.

Lepidopterists’ Society Mem. 2:i-vii, 1-280.

NIELSEN, M.C. 1999. Michigan Butterflies & Skippers. Mich. St. Univ. Press. 248 pp.

ROYER, R.A. 1988. Butterflies of North Dakota, an Atlas and Guide. Minot State University. 192 pp.

_ . 2003. Royer, R.A. 2003. Butterflies of North Dakota. Minot State University Science Monograph

Number Two. 192 pp. Scan of titus section.

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

Stanford, CA. 583 pp.

SHULL, E.M. 1987. The Butterflies of Indiana. Indiana Academy of Science, U. of Indiana Press, Indianapolis, IN.

262 pp.

THOMAS COOPER LIBRARY. University of South Carolina web site. John Abbot art and historical information

http://www.sc.edu/library/spcoll/abbot/default.html

UNITED STATES GEOLOGICAL SURVEY. Northern Prairie Wildlife Research Center. Butterflies of North

America Web Site, http://www.npwrc.usgs.gov/resource/distr/lepid/bflyusa/bflyusa.htm

WATSON, C.N. & J.A. HYATT. 1988. Butterflies of Northeast Tennessee. Journal of the Lepidopterists’ Society.

42(1): 19-31

* For Code purposes, new names published 14 May 2004.

17

Volume 4

10 May 2004

Number 7

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

A Concise Update of the Information Provided in

THE BUTTERFLIES OF SOUTHERN CALIFORNIA (1973)

by Thomas C. Emmeland John F. Emmel

Ken Davenport 1

6601 Eucalyptus Dr. #325

Bakersfield, California 93306

This paper’s purpose is to bring together in one resource a concise, but thorough, report on the current status of

butterflies discussed in the Butterflies of Southern California by Thomas C. Emmel and John F. Emmel, published

by the Natural History Museum of Los Angeles County, 1973. This update deals primarily with new species or

subspecies, taxonomic matters, and new distributional information. The family order fellows A Catalogue /

Checklist of the Butterflies of America North of Mexico by Clifford D. Ferris, editor, published by the

Lepidopterists’ Society as Memoir No. 3, 1989. Species order follows the Emmel, Emmel & Mattoon checklist for

California in Systematics of Western North American Butterflies, Thomas C. Emmel, editor, Mariposa Press

Gainesville, Florida (pgs. 825-836), 1998. The area delimited as southern California here is the same as in The

Butterflies of Southern California, except that all of San Luis Obispo County is included. Readers will need to

research several publications to read these changes since 1973.

The primary resources from which this updated information is drawn are: annual Season Summary reports

published in the News of the Lepidopterists’ Society, various scientific journals (including Journal of the

Lepidopterists ’ Society, Bulletin of the Allyn Museum and Journal of Research on the Lepidoptera) and articles and

information reported by various researchers and observers. Much such information is found in the Systematics

publication. In Systematics, there are 73 chapters written by a number of authors, with many of the chapters

including information on the southern California fauna. Information for Kern County is found in Butterflies of

North America 3. Butterflies of Kern and Tulare Counties, California by Ken Davenport, published as a

contribution of the C. P. Gillette Museum of Arthopod Diversity, Colorado State University.

FAMILY HESPERIIDAE (SKIPPERS)

Subfamily Megathymidae:

YUCCA GIANT SKIPPER Megathymusyuccae harbisoni J. Emmel & T. E mm el.

Update: M. coloradensis is now considered conspecific with M. yuccae. The new subspecies harbisoni was described in the

SYSTEMATICS publication. Subspecies martini is now known to occur in the southern Sierra Nevada of Kern County.

MOJAVE GIANT SKIPPER Agathymus alliae paiute Roever.

Update: Also known as ALLIE'S GIANT SKIPPER. The name paiute was applied in 1998 by Kilian Roever (SYSTEMATICS

book). E mm el & Emmel had stated California populations would be named by Roever back in 1973.

GENTRY’S GIANT SKIPPER Agathymus gentryi Roever.

Update: Southern California populations were previously known as BAUER'S GIANT SKIPPER {Agathymus baueri ) in Emmel

and Emmel. BAUER'S QANT SKIPPER is now considered a species ranging from the Phoenix area into northern Arizona

(some western Arizona populations are gentryi ). Others including NABA view all of these as ARIZONA or ARYXNA GIANT

SKIPPER {Agathymus aryxna ). See SYSTEMATICS book and Roever’s paper.

1 Staff Researcher, The International Lepidoptera Survey. Field Associate of Entomology, Natural History Museum of Los Angeles County.

Museum Associate, Colorado State Museum at Colorado State University. E-mail address: kdavenport@tils-ttr.org

ATLAS of WESTERN USA B

•d is a dot in extreme eastern

,ly as a rare stray.

sduvaliana (C. & R. Felder).

10 Co., 18 April 1992 (John F. Emmel and Bruce Gri

Update: Besides Roever’s Imperial County record in Emmel & Emmel (1973), Larry Orsak

lists an old record for Orange County: Doheny Palisades 15 IX 35 by Charles Rudkin. A n

FAMILY LYCAENIDAE (COPPERS, HAIRSTREAKS and BLUES)

of Orange County)

as P the canyon on the south side of Butterbredt Peak in Kern County. However it should be noted that populations on the east

j: One of the states most rare bi

10

^ (1987/1988 Vol. 26:1-4) sh

FAMILY NYMPHALIDAE (BRUSH-FOOTED BUTTERFLIES)

13

patdu'iu'heFWaf

Update: Long considered a fall species, most workere now view zephyrus to be the western representative of P. gracilis.

Update: There are many older records (1930's) from San Luis Obispo County (1 male Oceano 9 IX 32 and 1 female Oceano 8

X 32 ; Huasna 16 IX 32; Arroyo Grande 16 IX 32 and 17 IX 32 (all Homer Edgecomb) and the LACM has a specimen from a

unsuccessful. The Ribes divencatum host plant grows along Morro Creek and the butterfly may still occur there. It still occurs

' in a 1994 paper. 1

tharos are in the LA

FAMILY DANAIDAE (MILKWEED BUTTERFLIES)

rsaassrj

CHECKLIST OF THE BUTTERFLIES AND S

S OF SOUTHERN CALIFORNIA

23 more to be rare

S AND REGULAR STRAYS LIST

Family Papilionidae

17

Family Danaidae Duponchel

Family Satyridae Boisduval

71.

Family Libytheidae Boisduval

Family Riodininae Grote

h. v.prattii J Emmel & T. Emmel

derma cythera (W. H. Edwards)

a. c. cythera (W. H. Edwards)

Family Lycaenidae Leach

ehru behrii (W.

19

20

. channelensis J. Emmel & T. Emmel

PART II: RARELY R

21

Nymphalidae:

San Bernardino County, 5 April 1934 by G. H. and J. L. Sperry.

nty in the 1950 Season Summary. Extensive collecting in the

22

23

Volume 4

25 September 2004

Number 7A

i# The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

A Concise Update of the Information Provided in

THE BUTTERFLIES OF SOUTHERN CALIFORNIA ( 1973 )

by Thomas C. Emmel and John F. Emmel

- ADDITIONAL SOUTHERN CALIFORNIA UPDATES -

FAMILY LYCAENIDAE (COPPERS, HAIRSTREAKS and BLUES)

PART II: RARELY RECORDED SPECIES LIST

Satyridae:

LIKELY FUTURE CHANGES

Lycaenidae:

2

Volume 4

25 September 2004

Number 8

DESCRIPTION OF A MULTILEVEL CRYPTIC NEW SPECIES OF

PHYCIODES (NYMPHALIDAE: MELITAEINAE) FROM THE

SOUTHERN APPALACHIAN MOUNTAINS.

RONALD R. GATRELLE 1

126 Wells Road, Goose Creek, South Carolina 29445-3413

ABSTRACT. Phyciodes incognitus is described as a new species from the medium to high elevation hardwood forests of the

southern Appalachian Mountains, type locality: Duncan Ridge Road, 3700 ft, Union County, Georgia. P. incognitus is verified

from four colonies in Union County, Georgia and Clay and Macon counties, North Carolina. It is projected to range, in

suitable habitat, as far north as montane Pennsylvania (indicated by photos). P. incognitus possesses a unique set of character

traits that are virtually identical to both Phyciodes tharos and Phyciodes cocyta but at differing character positions - biology,

morphology, genitalia, mtDNA, phenology. P. incognitus is multi-brooded and sympatric with P. tharos and P. batesii

maconensis. P. incognitus can usually be easily distinguished in the field from sympatric tharos - especially males. Where

incognitus ranges north into the range of P. cocyta, it will be difficult to distinguish their females, and many males, except by

mtDNA or phenology. A lectotype of marcia Edwards, 1868 is designated from Kanawha River, Kanawha County, WV. P.

marcia is a synonym of P. cocyta. The currently known mtDNA data for the tharos- group of taxa is determined to have

limited, but specific, usefulness as a taxonomic tool in this group. It is concluded that sibling biological species are present

with similar mtDNA. Thus, mtDNA is only useful in this group to determine speciation where taxa have significantly different

mtDNA. Similar mtDNA does not indicate multiple species are not present. Specimens collected during this study suggest that

a second undescribed tharos- like species is present in the study area.

Additional key words: rearing, diminutor, orantain, marcia

TAXONOMIC OVERVIEW OF EASTERN PHYCIODES

Scott (1994) examined all the original descriptions and status of type specimens associated with

the tharos-growp of North American Phyciodes Hubner, 1819. At that time, Phyciodes tharos (Drury,

1773), Phyciodes cocyta (Cramer, 1777), and Phyciodes batesii (Reakirt, 1865) were considered the only

taxa (species or subspecies) in this group to occur in the eastern region of the continent. Of these, tharos

and cocyta can be very similar, especially in females. In examining the descriptive literature, Scott found

that even though there were only two (then known) species in the eastern region that could be confused,

the literature was not only inconclusive but potentially disruptive to the long held stability of what the

names tharos and cocyta delimited. Scott correctly designated neotypes for tharos Drury, cocyta Cramer,

and euclea Bergstrasser, 1780 to preserve stability. Scott noted the name marcia W. H Edwards, 1868

and while he did not recognize it as a subspecies of tharos himself, he pointed out that the name was

validly proposed at the species level and was thus available to any who wanted to “split” tharos into black

clubbed {tharos tharos) and red clubbed {tharos marcia) subspecies.

1 Staff Researcher, The International Lepidoptera Survey, Goose Creek, South Carolina.

Research Associate, Florida State Collection of Arthropods, Gainesville, Florida.

Scott’s 1994 paper resolved any possible historical ambiguity of the names tharos and cocyta.

Some workers (personal communication) have questioned some of his neotypifications. I do not. As

Scott points out, Drury described tharos as a taxon with brown antennal clubs. It does not matter that

some of the OD and associated paintings present red clubbed specimens. This is because female tharos

frequently have either brown or red nudums on their clubs and some males have red tipped clubs. Plus,

paintings are subject to artistic license, even failure. There are also various arguments about cocyta ,

selenis W. Kirby, 1837, and morpheus Fabricius, 1775. However, these arguments are precisely why

neotypes needed to be designated to settle the nomenclature and preserve stability. These names are left

as Scott dealt with them (Scott 1994, 1998). The stabilizing approach is to deal only with present

taxonomic issues in this genus - which are still many and complex.

In 1998, Scott published again on Ihe North American Phyciodes and presented significant new

discoveries. This included the description of an easterly western subspecies (< orantain ) and an eastern

subspecies ( diminutor ) which were tentatively, and ambiguously, placed in tharos and cocyta

respectively. The ambiguity exists because after describing these as subspecies of tharos and cocyta ,

Scott then presents alternate taxonomic placement of both as subspecies of one another and as individual

species. While unorthodox, this was nonetheless reasonable because it correctly reflected two things: 1)

that these taxa exist in nature as distinct organisms and 2) that their correct specific status was then

uncertain due to various factors indicating different possible relationships. Technically, it is important for

taxonomists to note that Scott acted as first reviser within his own paper and thereby presented both

orantain and diminutor as species. It is this researcher’s position that while these two taxa may be

subspecies of something, they are not subspecies of tharos or cocyta respectively (discussed later in this

paper).

In his 1998 paper, Scott reexamined the name marcia and determined that the lectotype designated

by F. M. Brown (1966) was invalid. However, only one of his reasons for invalidation is Code compliant

- it is not a syntype (as confirmed later herein). Scott then left the name marcia without a singular type

and without species level circumscription. Scott focused on the common usage of “ marcia ” as a form

name and left it at that. However, it was proposed at species rank and remains available, and more

importantly, applicable to a specific organism. Therefore, this name remains as a “loose cannon” among

the available names within the eastern Phyciodes and is thus specifically addressed and resolved herein.

Wahlberg, Oliveira and Scott (2003) studied the mitochondrial DNA variation in Phyciodes and

concluded the following.

“The results indicate that mitochondrial DNA sequences must be used with great caution in delimiting

species, especially when infraspecific samples are few, or introgression seems to be rampant”; and, "... the utility

of mtDNA on its own in assessing the boundaries of traditionally recognized species (e.g. Weins & Penkrot, 2002)

is suspect. One must combine all possible knowledge, including morphological, ecological and molecular, to

understand the species boundaries of groups of very closely related species. Our study has raised more questions

than it has answered and will certainly help focus future research on the process of speciation in the tharos- group of

species of Phyciodes ”.

This researcher was in periodic contact with Wahlberg and Scott contributing some specimens and

comments toward their study. They have also been consulted at various times on ma tters relating to the

research presented herein. Chiefly, several NC specimens of both tharos and new species incognitus

(described herein) were sent to Wahlberg for mtDNA sequencing, and photos of immatures and adults

were sent to Scott for taxonomic assessment. This current paper, by documenting the existence of two

genetically cryptic and two phenotypically cryptic species within Phyciodes , confirms the Wahlberg et al.

2003 conclusions quoted above re determining taxonomic relationships in Phyciodes.

In 1998, this author published the discovery and description of Phyciodes hatesii maconensis from

the southern Appalachian mountains, type locality: Jones Knob, Macon County, NC. In May 2002, it was

discovered that a cocyta- like taxon also existed at the Jones Knob TL. It has now been determined that

three sympatric Phyciodes species occur in the Jones Knob general area: batesii maconensis , tharos , and

2

new species incognitus. This new species was not noted during the years of maconensis study for two

reasons. First, was a preoccupation with studying maconensis , and second, it occurs in a micro habitat

within the Jones Knob general area not previously searched until 2002. While tharos occurs ubiquitously

throughout the Jones Knob area, this new species is habitat restricted.

Today, there are six known and described Phyciodes taxa in eastern North America: tharos Drury,

1773, cocyta Cramer, 1777, batesii Reakirt, 1865, maconensis Gatrelle, 1998, diminutor Scott, 1998 and

incognitus Gatrelle, 2004 (herein), with marcia W. H. Edwards, 1868 determined a synonym of cocyta.

Figures 1-9. Mature larval heads of P. incognitus and P. tharos at varied angles. Figs. 1, 2, 5 & 6 left. P. incognitus. Note

more rectangular shape and darker face. Figs. 3, 4, 6 right, 7 & 9. P. tharos. Note round face and heavy markings, especially

around mouth and eyes (handlebar mustache). Fig. 8. Face of Phyciodes tharos from Edwards’ Butterflies of North America.

Figures 10-15. Larvae of P. tharos and P. incognitus. Figs. 10 & 14. Mature larvae of P. incognitus. Figs. 11-13. Mature

larvae of P. tharos ;12 in extremely bright light to artificially enhance colors. Fig. 15. Clutch of mature first instar, and newly

molted second instar, larvae of P. incognitus. First instar larvae appear hairless to naked eye and in low magnification. Figure

16. Eggs of P. tharos. Figures A-E. Antennal shafts and clubs of P. tharos and P. incognitus. Fig. A. Round club with black

nudum: male P. tharos. Fig. B. Oval club with black nudum: female P. tharos. Fig. C. Round club with partial orange nudum,

female P. tharos. Fig. D. Elongate club with orange nudum: male P. incognitus. Fig. E. Oval-elongate club with orange

nudum: female P. incognitus. Photos: Joseph Mueller.

3

Iktf &*lSr

29l_J^ 30 '

tefc# Sjfei, **8 ^-ggr

It^y W 1

sssiism

L Ma Mb

Figures 17-43, Phyciodes species (exact size). P. tharos : figs. *176', *18/29 6\ *19c?, *22 9, *23/34 9, *24d\ 39b, 40c?, 41 9: fi

Sept. 88, Barr Lk^Adams Co., COJeg Scott. P. diminutor : figs. 25 1,261 paratypes: /11 June 95, ^ex pupa 31 July 96, 3 mi. NE

" ' * len, Freeborn Co., MN, leg Scott. P. cocyta: fig. 37. * topotype, P cocyta, 10 July 92, Sydney, Cape Breton Is., Nova f '

figs. *27<?’ *28 9, 30/35o', 31/369, *32o', *33 9,38* figs. 30/35 holotype (datain text); figs. 31/36 allotype (data ) in text); fig. 27

paratype, 9 May 03, Union Co., GA, Duncan Ridge Rd; fig. 28 paratype, 16 May 02, Jones Knob, Macon Co., NC; figs,

paratypes, 5 Sept. 03, Clay Co., NC, Sally Gap Rd, fig. 38 paratype, ex pupa 19 June 03, (same site as 27). Figures F-K, ge,

(key in text); P. tharos: F & I; P incognitus : G-H, J-K. Figures UP, pupae: figs. L-M, P. tharos (M, dominant phenotype); fi

EDWARDS’ MARCIA

The name marcia W. H. Edwards, 1868 was not dealt with fully by Scott (1994 & 1998) and thus

remains as potentially disruptive to the nomenclature. This is because its typification remains unsettled.

This is corrected herein by lectotypification. The following is an enumeration of the problems that

presently exist that make this lectotypification necessary.

1) Scott (1994) addressed W. H. Edwards’ 1868 name marcia and noted it is an available species

level name. Scott also noted that in the OD Edwards stated that species marcia was large and had red

antennal clubs in both sexes. Scott then followed the tradition based association of the name marcia,

1868 with tharos - an association began by Edwards in his 1874-1884 Butterflies of North America 2 .

Scott stated that there existed in nature a small sized northern Phyciodes with orange-red nudums on its

antennal clubs, which he, at that time, assumed to be a tharos. Scott gave its range as, “ S Maine to C

N.Y. (and some Pnn. colonies) W to Sask.-Alta.-Dakotas-W Neb.-Wyo.-Colo.-extreme W Okla.” Scott

then applied the 1868 large species name marcia to this small, orange clubbed entity concluding it was a

weak northern subspecies of tharos differing only in having orange-red nudums. Scott gave no indication

in 1994 that “ marcia ” was only a form name.

In his 1998 paper, Scott had decided that his 1994 tharos marcia concept was an undescribed

taxon which he then described as, Pharos orantain or cocyta orantain or orantain ”. However, Scott

noted that the name marcia was problematic to his describing the new taxon orantain as 1) both his

orantain and marcia 1868 had orange clubs and 2) in 1994, Scott had applied the name marcia to this

same organism. Scott (1998, page 7) presents a largely subjective analysis and reasons why he concludes

F. M. Brown’s 1966 lectotype of marcia is invalid and why marcia (of 1868) is a tharos “form” only

(even though described as a species) limited in range to the eastern US. Scott did not examine the

lectotype nor any of the many Edwards marcia specimens in the Carnegie Museum. He simply based his

assessment on the black and white photo of the lectotype in Brown’s 1966 paper. Scott then described

orantain as a western taxon (by above multi-combinations) and states that orantain “... does not occur in

eastern North America”. Scott then redescribes the remainder of his 1994 tharos marcia concept

organism as taxon “cocyta diminutor or orantain diminutor". This position is untenable. The name

marcia is not a form per its 1868 OD. The OD range of marcia included montane New York which is

well within the range of diminutor. Thus, the name diminutor has to be considered as possibly what

Edwards meant by species marcia 1868. If northern and eastern diminutor is the same species as

orantain , the combinations, by priority, would then be marcia marcia [=diminutor\ and marcia orantain

and not orantain orantain and orantain diminutor as Scott proposed as first reviser.

2) The discovery of a multi-brooded, large-winged, orange-red nudum cocyta- like non -cocyta

taxon in the southern Appalachian mountains that is sympatric with black clubbed tharos necessitated that

this researcher also examine Edwards’ marcia to determine if this “new” taxon was what Edwards meant

by species marcia. Scott’s relevant papers were in hand. Copies of Edwards’ 1868 species marcia

description, Edwards’ 1874-84 section on Phyciodes tharos (introducing form marcia ), and Brown’s 1966

lectotypification paper were obtained; also, Brown’s marcia lectotype and 66 syntypes by loan from the

Carnegie Museum. Together, these resources provide the evidence of what Edwards found, where he

erred, and what is the best way to move forward with the least damage to current usage of names.

2 The problem is that Edwards’ 1868 marcia and Edwards’ 1874-84 marcia apply to two different species. In the former,

Edwards described a new species - thus, marcia, 1868 is both available and delimited per the 1868 description and those

syntype specimens. Edwards 1874-84 used the name marcia for a form of species tharos - thus, marcia 1874-84, as an

infrasubspecific homonym, is unavailable from that date and application. Edwards did this because he erred in confusing his

first entity (= species cocyta ) with the spring form of species tharos (and other taxa) he reared years later. See points 3 & 4.

3) Edwards’ 1868 description of Melitaea marcia is textually thorough but lacks illustrations.

Two features stand out in this description. First, the large wing expanse of the new species: 1.3 to 1.5

inches in the male and 1.7 inches in the female. Second, the orange nudum of the antennal club of both

males and females, “...antennae black above, whitish below; annulated with white; club black above,

fulvous below.” Because the dorsal and ventral wing markings of the eastern tharos-group Phyciodes are

so similar and variable, only batesii can be descriptively eliminated by Edwards’ wing description. The

large size and club color specifically indicates something other than tharos, as tharos is relatively small in

the Appalachian region and the clubs of eastern male tharos are either all black or only slightly red tipped

(Allen 1997). There are two large sized Phyciodes species with orange-red nudums in the Appalachian

region - cocyta and incognitus (described herein). Edwards’ 1868 marcia is likely one of these two with

high probability.

Edwards stated that he considered marcia to range from New York to Louisiana, but he mentioned

only one actual site, stating, “I have found Marcia common on the Kanawha River, W. Va., in June.

Batesii is of earlier flight and rather rare in that region.” This site is thus where Edwards collected the

syntypes of his new species and establishes the Kanawha River, WV as the marcia type locality.

There are a large number of specimens labeled “ marcia ” in the Edwards collection at the Carnegie

Museum (Brown 1966). These are apparently assumed to all be syntypes because they have Edwards’

hand written labels on them. Sixty six of these “syntypes” and the marcia lectotype (invalidated by Scott

(1998)) were sent on loan for this research. One box of 30 specimens was almost totally destroyed in the

mail, but 22 of those still possessed enough wing fragments and antennae to be determinable to species or

species group. Thus, a total of 58 “syntype” specimens, all labeled “ marcia ” by Edwards, were assessed.

Thirteen of the 58 specimens are reared individuals with 6 of those being aberrations due to being

placed on ice as pupae. One aberration lacks antennae and is thus not determinable to species. The 12

with antennae are comprised of 3 tharos and 9 with orange nudums and of larger size which are thus not

tharos. None of the reared specimens have locality data.

There are 7 collected tharos for a total of 10 tharos out of 58 total specimens. These 7 are from

NC (3), TX (2), Hunter, NY (2). The 8 destroyed specimens have only the thorax remaining on the pin.

From their labels and small thorax size, it is probable that 5 were tharos - GA (3) and NC (2). This is

because the 3 other NC specimens among those determinable are clearly tharos. This would make a

maximum potential of 15 tharos out of 66 specimens sent (22%). The remaining 38 specimens all have

strongly orange nudums on their clubs. Of these 38, 20 males and 7 females (27) are strongly to likely

determinable as species cocyta (6 being Canadian - Labrador, Anticosta and Montreal; and 1 from New

Mexico). Three are possible incognitus. The remaining 8 specimens are either small cocyta or diminutor.

The 66 specimens sent may or may not be a typical sample of Edwards’ specimens - both in

phenotype and location. The high percentage of non -tharos was unexpected (58 minus 10 tharos for 48,

mostly larger, orange nudum specimens = 82% non -tharos). It was also surprising to find only 1 probable

and two possible new species incognitus among them. It was expected that many P. incognitus would be

among the “syntypes” and that it would thus be demonstrable that the “new” entity found at Jones Knob,

North Carolina would be a rediscovery of Edwards’ 1868 species marcia. But such is not the case. As

stated earlier, there are two large winged, orange nudum Phyciodes species in the Appalachians and one

of them is what Edwards called species marcia. From the material exami ned, that species is cocyta, to

which marcia Edwards, 1868 is thus a synonym. This is further confirmed by the following.

Alex Grkovich (pers. comm.) had a very large female specimen he collected from Greenbrier

County in southeast West Virginia determined by Wahlberg as species cocyta by mtDNA analysis. This

is the furthest south in the Appalachians that cocyta has been confirmed by mtDNA examination. This

confirms that cocyta’s range extends fully into the area from which Edwards collected his specime ns of

marcia on the “Kanawha River, West Virginia”. Edwards’ 1868 Kanawha River site is the same location

as Coalburg, where he lived. In 1894, the town of Coalburg had its name changed to Cabin Creek. This

area is only 55 miles west northwest of Greenbrier County.

6

In examining the Edwards series, 18 cocyta specimens are labeled (by Edwards) from “Ka”,

“Kan” or “Kan a ”. At first, this was thought to mean Kansas. However, this cocyta phenotype does not

occur anywhere near Kansas (Wahlberg et al. 2003). It was noticed that two forms of location notations

are on his labels. They either have a location of US state or CA province (Texas, Ga., NC., W. Va., N.

Mex., Colo., Labrador, Anticosti) or specific local sites (Montreal, Coal, Coal b, Ka, Kan, Kan a , Hunter,

W mtns.) In reading Edwards 1868 and 1874-84 all the these locations are mentioned, but not Kansas.

This worker’s position is that Ka, Kan and especially Kan a is the Kanawha River site. If so, then

specimens so labeled are most likely the valid syntypes of Edwards’ 1868 species marcia. This locality

comprises the largest percent of specimens (of those sent). Additional proof of this is a unique inscription

on one specimen. It is a small cocyta or normal sized diminutor female specimen with one intact antennal

club with an extensive orange nudum. The label says “Marcia female, C Kan a Sept, caught”. The word

caught surely refers to Edwards having caught this himself versus having reared it. “Caught” is on the

bottom line by itself, the same area Edwards made his rearing notations (e.g. “on ice” or “reared”.)

4) Type localities, types, and Edwards point of error re marcia as a form. According to the

International Code of Zoological Nomenclature (ICZN), where syntypes are involved, they collectively

constitute the type and the sum of their locations comprises the type locality. A type locality is the place

the type specimen(s) is/are from. Edwards’ 1868 type locality is NOT “New York to Louisiana” - that

was his range statement. The type locality is the place the syntypes he had came from, and the only site

he stated having specimens from is “Kanawha River”. Only specimens seen or referred to by Edwards for

his 1868 description of Melitaea marcia are syntypes. Any of Edwards’ specimens not utilized in his

1868 description of marcia can not be syntypes. The fact that Edwards has labeled all of these “ marcia ”

does not make them syntypes. That is simply an ID he wrote on them some time during his life.

Several of the specimens on loan have dates on them around 1877 - years after the species marcia

description. All specimens with post 1868 dates are excluded from being syntypes by the Code. Of those

without dates, a great many are disqualified as syntypes simply by their location (e.g. Canada). In the

OD, the only specific location is the Kanawha River. However, in Edwards’ Butterflies of North America

(1874-1884), two primary areas are specified relative to his research into tharos seasonal polymorphism.

These primary areas are Coalburg 3 , WV and Hunter, in the Catskills of NY. Specimens from these, and

any other locations mentioned in his BNA in conjunction with his polymorphism studies, are excluded

because those studies did not begin until after 1875. In the Butterflies of North America Phyciodes tharos

account, Edwards states that he did not rear any specimens before 1875 (emphasis mine).

“In 1868,1 described, as a distinct species, ... calling it Marcia. ... But there were such resemblances also

to Tharos that it was not possible to determine its specific value, unless the butterflies could be bred from eggs, and

as yet [1868] the food-plant of the larvae, and the larvae themselves, of both Marcia and Tharos were unknown.

But in 1875, the food-plant was discovered by Mr. Mead. He states, in Can. Ent. VII., p. 161, that he planted ... all

common composites... introduced a number of females of this species. A few days later, on examining the leaves,

he found eggs deposited on the Aster Nova-Angliae, and on no other plant... This happened in the month of July,

near the last of the month, at Hunter, N.Y., among the Catskill Mountains.”

There are 5 Hunter, NY specimens among those on loan for this study. Three have dates of 1876

or 1877, two lack dates. There is no evidence Edwards had any specimens from Hunter, NY in 1868.

There is sufficient evidence, including specimen dates, to show that all Hunter, NY specimens are not

1868 syntypes - including the “lectotype” designated by F.M. Brown in 1966. It is further reasonable to

conclude that most specimens labeled “ marcia ” by Edwards’ (some dated as late as 1886), from many

locations, are from his extensive post 1868 collecting and rearing of Pearl Crescents and not 1868

syntypes.

3 Coalburg and Kanawha River are the same site. In 1868, Edwards referred to (and labeled) specimens from that site as “Kan,

Ka or Kan a ”. By 1875, he was referring to (and labeling) specimens from that site as “Coal, Coal b”.

It is a fact that Edwards described marcia as a species distinct from tharos as he states, . .hitherto

considered a variety of Tharos , from which it is readily distinguished by the purple or pearly hue of the

underside of secondaries.” It is factual that Edwards did not consider marcia a mere seasonal form until

after 1875. It is a fact that a great many (perhaps vast majority) of what Edwards considered tharos form

marcia are species P. cocyta as evidenced by his extant collection. It is a fact that Edwards also had

specimens of tharos he considered form marcia. Because specimens of both cocyta and tharos (and likely

diminutor ) are among the specimens reared by Edwards, it is unarguable that Edwards was confused

about what species he was dealing with because he didn’t detect these species in hand. How did this

happen?

The above quote in Edwards’ marcia OD states that he considered the coloration of the ventral

surface the character of differentiation between marcia and tharos rather than size and club color. While

Edwards noted wing size and club color in the 1868 OD, it is apparent from his specimen IDs, that he had

become oblivious to the major differences in size, dorsal markings, and club color between tharos and

cocyta just seven years later as being indicative of speciation. In reading his presentation of means and

methods, it is stated that he only gathered egg masses from wild females and never tried to cross any of

the phenotypes. He simply focused on rearing one seasonal form from the previous seasonal form.

It is now well known, from various rearings of Eastern tharos and cocyta in this region, that

cocyta always has bold orange clubs while tharos always has all black clubs (with occasional red tips) in

males and mixed clubs in females (Allen 1997, Scott 1994) (Figs. A-C). It is thus knowable that all the

egg clusters that Edwards reared would have each produced the above distinctively different, and species

specific, adult antennal morphology. Perhaps some egg clusters were combined on feeding plants as he

was only after seasonal variation (season to season, not family to family comparisons) and may have

thought he was getting variable nudums and phenotypes from the same clusters. However, we know from

his accounts that he did segregate some egg clusters. In fact, it is his account of such segregation that

reveals his error. Here is the BNA quotation (emphasis mine).

“This second generation was just one month behind the second at Coalburgh. So far only could I trace the

Catskill [Hunter, NY] generation this year; but as, in 1875, Mr. Mead obtained eggs on the 27 th July and following

days, the larvae from which all hibernated, that would be the second laying of eggs of the season, and the resulting

butterflies the first generation of the following year.

“The foregoing Coalburgh [=Kanawha River] observations were supplemented by others in August, 1877.

Between 14 th and 20 th , I obtained three lots of eggs, from which the larvae in due time emerged. Those of the first

all went on to maturity, giving butterflies after middle of September, the last emerging 26 th . But of the other two

lots all became lethargic [diapause]. The reason for this difference I could not conjecture. It certainly was not

owing to any change in the weather [evidencing outdoor rearing]. In the field the species was abundant from 15 th to

25 th August (this being the third brood of the year). But one month later, when the fourth brood should be flying,

examples were remarkably scarce.’’

The “reason” that eluded him is quite evident today, they were Phyciodes cocyta; whose larvae all

go into diapause at that time of year (end of July through late August). This is also what happened to Mr.

Mead’s larvae. It is also why the supposed second generation was on the wing only one month after the

previous emergence. The previous entity would have been tharos and/or diminutor and/or incognitus. It

takes a minimum of 35 days to rear these species indoors under perfect warm conditions. In the

mountains, with long and humid cool nights, mornings, and evenings it takes much longer. Thus, one

brood being virtually on the heals of another is an indication of multiple species being present. Larvae

from multiple females obtained at the normal flight season of cocyta and which all enter diapause strongly

indicates a non -tharos species being present.

These accounts and the Edwards specimens evidence that Edwards was so focused on seasonal

variation that he missed the fact he was dealing with two to four species - tharos, cocyta, diminutor and

incognitus. All of which likely occur in West Virginia and at least two of which {tharos and cocyta) have

been unequivocally identified among the specimens he both collected and reared, and labeled as

“marcia”.

8

5) Melitaea marcia typification. Edwards never provided any evidence showing marcia of 1868

to not be a species. He merely reared and collected parallel seasonal forms from multiple species and

arbitrarily called them all “marcia”. Edwards applied an 1868 valid species name to a seasonal form after

1875. There are thus two types of taxa going by the name marcia - one a valid species (; marcia = cocyta)

and one a seasonal form occurring in many Phyciodes species (inc. tharos f. marcia , cocyta f. marcia ,

diminutor f. marcia, incognitus f. marcia.). Though the same, the names have no taxonomic connection.

A species can not be changed to a form solely because someone gives it a different concept.

Brown (1966) approached Edwards’ marcia specimens as tharos forms because of Edwards’

erroneous presentation of marcia as only a seasonal variant in his Butterflies of North America. Brown

designated a lectotype for the species marcia 1868, but did so per an infrasubspecific concept authored in

1874-1884. The Brown specimen has a notation on its label by Edwards that it is the “type” of Edwards’

form concept (Fig. 39b). The specimen Brown utilized is not a syntype of marcia 1868 and is thus invalid

as a lectotype (per Scott 1998). This is because the specimen is from Edwards’ post 1875 research as

evidenced by 1) appearing bred, 2) matching the BNA form A 1-2 figures and, 3) being labeled from

Hunter. The “A” is part of a form code (A, B, C & D) created and employed by Edwards in his post 1875

research. The primary disqualifying factor is its place of origin - Hunter, NY. (This situation was

presented to the official ICZN internet list serve. The only thing that can invalidate this lectotype is if it is

not an 1868 syntype. Which it is not.)

The above research into species marcia W. H. Edwards, 1868 supports and confirms Scott’s 1998

invalidation of Brown’s 1966 lectotype of marcia 1868. However, this specimen is the actual “type” of

Edwards’ infrasubspecific BNA early season form marcia of species tharos. This is because it is labeled

as “type” by Edwards and is clearly the BNA model for figure A 1-2. While no longer having antennae

(Figs. 39b), the painting made while it did, has black clubs (Fig. 39a). The wing shape, all markings, and

size confirm Scott’s 1998 diagnosis of this specimen as female - not male. Thus, being female and

having black clubs confirms that this specimen is a partially aberrant reared tharos. The name marcia

may therefore continue to be used for the heavily marked spring forms of Phyciodes in general and tharos

specifically.

The preceding means that a proper lectotype for Melitaea marcia Edwards, 1868 is needed to

provide what the Code refers to as the “objective standard of reference” by which taxonomists can apply

this name to a taxon that bears this name. Among the Edwards specimens sent, a male and a female stand

out as likely true syntypes and typical examples of his 1868 original description. The male has orange

nudum and is 1.3 inches in expanse. The female lacks antennae but is 1.7 inches and with contrasting

forewing colors typical of cocyta. Both are from the Kan a location. The male has a “D” on its label

which would have been added later. Also, many of these specimens were surely relabeled because the

labels are all so similar (same paper and pen markings) even among those that have very different dates

(e.g. 1876 & 1886). Because 1) both his 1868 species and post 1875 form have the same name and 2) all

specimens have the name marcia on them, one can not use the presence of the name marcia on a label to

date specimens or confirm syntypic status.

Trying to do the best one can with this complex situation, I here designate this male (Fig. 43) as

the lectotype of Melitaea marcia Edwards, 1868. This retains the type locality as Kanawha River

(Coalbrug), West Virginia. This specimen and the paralectotype female (Fig. 42) are, with very high

probability, species Phyciodes cocyta. Melitaea marcia Edwards, 1868 is thus either a synonym of Phy¬

ciodes cocyta cocyta or a valid name for a “southern” brighter subspecies (compared to typical Canadian

cocyta (Fig. 37)) of cocyta in the mid Appalachian Mountains - Phyciodes cocyta marcia. Its labels are

figured with the lectotype and the specimen is returned to the Carnegie Museum, Pittsburgh, PA.

This maintains stability of all eastern North American Phyciodes names in their current usage. It

insulates the new species/subspecies ( orantain, diminutor, incognitus) from any potential confusion with

the name marcia. These recently described non -tharos taxa need to be free from any associations with the

long ^ras-associated and form-associated name marcia.

9

SPECIES STATUS OF PHYCIODES DIMINUTOR SCOTT, 1998

Alan Wormington of Ontario, Canada has relayed the results of his Ontario Phyciodes studies and

details of his discovery of what he identified as P. cocyta in Union County, Georgia on 18 August 2002.

The following are selected comments from his personal communications on his Ontario research.

“Here where I live, Point Pelee National Park in southern Ontario, I have been studying butterflies for

decades and have written several annotated lists... In 1995 I put in a special effort to understand the brood

relationships of crescents here, when I realized the whole thing was a complete mess. Finally, I came to the

conclusion that the "mess" was actually the result of a third entity involved that obviously had never been described

in the literature. The two species that "normally" are mapped for this area, and across much of southern Ontario,

etc., are cocyta and tharos.

“In addition, I worked on determining which "new" species was actually new. Cocyta is obviously distinct,

but of the other two I couldn't determine which was true tharos and which one was undescribed. Shortly thereafter

I was able to visit the National Museum in Ottawa, where Don Lafontaine showed me the original drawings of

Dmry (1773) that apparently act as the type for tharos.

"The drawing matched what I had been calling "Summer" Crescent (since the first brood does not emerge

until very late in June). The drawing showed orange tips to the antennae clubs, which is what I had isolated as

"Summer" Crescent. Thus the other entity, by default, is the undescribed taxon — or is it? This other entity is the

one with very rounded and jet black antennae clubs, and tends to be small in size (also, there is less size differences

between the sexes compared to other crescent species). I have given it the name "Early" Crescent since the first

brood appears very early, starting most years around May 10 or shortly thereafter. In southern Ontario, it tends to

be slightly less common than the other crescents, and tends to be found on dunes, alvars, very dry fields, etc. Some

of what I discuss above is briefly mentioned in The Butterflies of Canada text”.

Scott’s 1994 neotypification of tharos stabilized that name to its traditional concept as the small

sized, round club with black nudum, early and multiple brooded Eastern species = Wormington’s Early

Crescent. In 1998, Scott described diminutor = Wormington’s Summer Crescent. In 1995, Wormington

had correctly determined three species to be present in Ontario - tharos, cocyta and diminutor. He also

relayed this significant range observation about species diminutor.

“I don't like the name "Northern" Crescent, since Summer Crescent [< diminutor ] occurs farther north in

Ontario than cocyta. I prefer the common name "Orange" Crescent, which I think was used first by some authors.”

Wormington’s range observations, in conjunction with Scott’s 1998 diminutor and cocyta range

data, confirms that the ranges of diminutor and cocyta broadly overlay one another over thousands of

square miles and that in many areas the two are sympatric. This makes it impossible that these two are

“subspecies” of one another. Scott (1998), as first reviser, proposed diminutor as a species distinct from

cocyta specifically because of their overlaying ranges, sympatry, and different phenology. That status is

followed herein. The taxonomic assignment of diminutor as a subspecies of cocyta in Wahlberg et al.

(2003) is incorrect. It is a falsifiable taxonomic conclusion. The similar mtDNA of cocyta and diminutor

must be assessed taxonomically against their biology, phenology and morphology which renders the

conclusion that they are distinct, often sympatric, species with very similar (to identical) COI mtDNA.

While common names have no scientific standing, their greatly expanded use today calls for a

reexamination of the common names for these species. Wormington’s common names of Early Crescent

(tharos ), Orange Crescent ( cocyta ) and Summer Crescent (< diminutor ) are good common names for these

species (especially in the northern US and Canada) and this usage is supported here. The term “pearl

crescent” was coined only in reference to form marcia - which occurs in many early season Phyciodes

species. In other words, there is no single Phyciodes species that is a “pearl” or “pearly”. With the

modern knowledge that there are multiple species in eastern North America which all may have pearly

undersides in spring and at times late fall, the name Pearl Crescent should be dropped as a common name

because it can lead observers (especially beginners) into confusion. This is because they may ID all short

photo period pearly HW marked individuals incorrectly as species tharos , and in summer be confused

because no species have pearly markings at that season.

10

DOCUMENTING A NEW SPECIES

DETERMINATION OF SPECIES

The most recent taxonomic placement of tharos -group Phyciodes to species and subspecies is in

Wahlberg et al. 2003. That paper is here considered to have two foundational taxonomic flaws. One, is the

acceptance of the taxonomic determinations of specimens sent per the determinations by their senders.

Second, is a presupposition that the overall taxonomy (strictly following that of Scott) was correct. The

mtDNA sequences and resulting tree diagrams were molded together with that taxonomy and those

identifications. The product is various taxa (names) being found in multiple (polyphylectic) locations in

clades B, C, D, and E (pg. 262). Primary among them is the name selenis which is found in all four

clades. It is suggested here that 1) various named individuals may be misidentifications and 2) various

taxonomic associations may be incorrect. Incorrect either to subspecific associations or undescribed taxa

being named as something they are not. See Funk and Omland (2003) relative to taxonomic assumptions

and gene trees.

This assessment seems to be indicated by comments in the paper itself.

“However, species defined using non-DNA characters were not well defined using mtDNA characters, and

indeed there were a large number of poly- and paraphyletic mtDNA lineages in different taxa of Phyciodes.” (pg.

263) and:

“...we prefer to interpret our results in the framework of the traditionally held concepts of species in the

thar os- group. Our results can be interpreted to be in strong conflict with the traditionally defined species in the

thar os-group, especially if one would redefine species based on mtDNA. Since mtDNA disagrees so strongly with

what any good field biologist can observe in nature regarding Phyciodes, we will question the traditional concepts,

but will defer any actual changes to them until further investigations have taken place.” (pg. 264).

The phrases “prefer to interpret” and “can be interpreted” are significant because this renders all

such derived at conclusions as unauthoritative due to their subjectivity and reasonable alternatives. The

flaw in this paper was a predetermined, but partially erroneous, taxonomy and/or occasional incorrect

individual specimen identifications. It is the view here that not nearly enough “field biology” has been

done to definitively place various Phyciodes taxa in correct taxonomic associations. Too many specific

and subspecific associations are still being based simply on the general “look” of various entities.

The mtDNA sequences presented in the Wahlberg study should be applied in two different ways

in determining taxonomic relationships in the tharos- group of taxa. One, is that significantly dissimilar

haplotypes between populations should be taken as evidence of speciation regardless of phenotypes where

no other data (e.g. biological) is available. Second, is that similar and even identical haplotypes should be

rendered null as taxonomic indicators when biological data shows two or more taxa to have species

integrity when sympatric in nature.

Example. In clade C, both the batesii batesii male and cocyta selenis female are from the same

date and location. (Males of cocyta selenis were also sequenced from this site and date.) It is very

possible that this “ selenis ” female is actually a misidentification of an atypical batesii female (compare

this female with batesii females 73-10 and 95-9). It has also long been this author’s position that batesii

lakota and batesii batesii are the same subspecies with variable phenotypes (this is supported by the

similar haplotypes in C of MN “ lakota ” and Ont. “batesii”. One can see from the phenotype why the C

cocyta selenis identification was made, but the irtDNA would demonstrate that that taxonomic “visual”

determination is in error and it is actually a variant female of batesii - with which it was collected. Thus,

everything in C would actually be P. batesii batesii and C would be monophyletic rather than para¬

phyletic.

Example. The taxa orantain and diminutor are therein considered subspecies (per Scott) of tharos

and cocyta respectively. However, this is not possible with diminutor and cocyta (as detailed above) due

to their wide sympatry. They are simply a pair of cryptic species with similar phenotypes and mtDNA.

Scott was correct in 1998 to suggest these two might prove to be two species.

11

Example. It is also unlikely that tharos and orantain are conspecific because Scott (1998) easily

crossed orantain with a “ selenis”. (Some of those crosses are housed here in the TILS collection.) The

taxonomic association of the orange nudumed orantain with true eastern black nudumed tharos is simply

an assumption. The biological compatibility of orantain and this “ selenis” supports their being

conspecific and works against the hypothesis that orantain is a tharos merely because they have some

superficial wing similarity. But what is Scott’s selenis used in this cross? (And what are the many

“selenis” spread ihroughout Wahlberg’s clades B, C, D and E?. It is possible that those “selenis” are

actually a menagerie of taxa perhaps described or undescribed.) Scott was correct to postulate that

orantain might be a species - so likely so in fact, that he actually described it as such therein as first

reviser! It is just as logical to consider it a subspecies of “ selenis” or a full species than a subspecies of

tharos with which no biological connection has been demonstrated. The fact that orantain has mtDNA

similar to species tharos only proves it is not species cocyta. This is because cocyta and tharos were

demonstrated by their mtDNA to not be closely related (Wahlberg et al., pg. 263).

Example. The nesting of batesii anasazi Scott, 1994 in clade E should be taken as evidence that

anasazi is actually a genetically cryptic full species or a subspecies with the “ selenis” with which it is

nested rather than a subspecies of batesii.

Example. The location of various individuals, as those of batesii lakota in clades B, D, and H,

can be taken as evidence that peripheral populations (e.g. Alberta “lakota”) are not subspecifically

equivalent. Or, that a specific individual may be a hybrid.

The larger issue however, is that it is now evident that there are an undetermined number of

Phyciodes species within both the tharos and cocyta mtDNA general haplotypes. This helps to explain

the why of this statement in Wahlberg et al.: “However, species defined using non-DNA characters were

not well defined using mtDNA characters, and indeed there were a large number of poly- and paraphyletic

mtDNA lineages in different taxa of Phyciodes.” One point here, is that this is due (to an unknown

degree) to the flaw of assuming the taxonomic delineations were correct to begin with, and then

maintaining and incorporating them into the taxonomic placement of names within the cladistic

conclusions.

The paper, On the use of genetic divergence for identifying species, by J. Willem H. Ferguson

(2002), offers perspectives applicable to the taxa in the tharos-group of Phyciodes. Here are some

excerpts (emphasis mine). The last quote is the final concluding remark.

“Coyne & Orr (1989) investigated pre-mating and post-mating isolation in several Drosophila species and

concluded that, among sympatric species pairs, pre-mating isolation arise more rapidly then post-mating isolation.

...pre-mating isolation may arise before post-mating isolation in many cases. This would appear to indicate that the

degree of genetic divergence required for pre-mating isolation is often less than that required for post-mating

isolation.

“Firstly, species with strong pre-mating isolation and weak post-mating isolation are likely to have little

genetic divergence because pre-mating isolation could be brought about by fewer than 10 loci.

“The aim of this paper is to contribute towards operational molecular yardsticks for identifying separate

species. It does not aim to disqualify genetic divergence as a useful tool in systematics. It is useful in many ways,

e.g. h population-level analysis and phylogeography, but on its own it is not useful for identifying separate

species. Systematists need tools that are parsimonious, have well understood foundations, and that can be used

consistently across a wide range of taxa. In terms of the identifications of new species, genetic divergence fails on

all three of these criteria.”

The pattern emerging out of the tharos- group is that several to many species are present that are

cryptic to human detection in one character but not in others (i.e. biological vs. molecular). Thus, to

detect and define species, multi-level data must be examined. With respect to mtDNA analysis, a species

will be demonstrated to be distinct from one species thereby, but not distinct from another thereby. For

example, P. diminutor is proven to not be P. tharos thereby, but can not be proven to not be P. cocyta

thereby. Likewise, P. incognitus is confirmed to not be P. cocyta thereby, but can not be proven to not be

a P. tharos thereby. In both instances, sympatry and biology reveal the false positive of the mtDNA data.

A NEW CRYPTIC APPALACHIAN SPECIES

This author has been collecting and researching butterflies in the southern Appalachian Mountains

since the mid 1970s. This has involved scores of days in the field over nearly 30 years at many and

diverse locations in northern Georgia and western North Carolina. The areas of heaviest concentration

have been in Macon and Clay counties, NC and Union and Rabun counties, GA.

Phyciodes tharos is common throughout the southern Appalachian region - except in dense forest.

Many specimens of tharos were sporadically collected over those years from many sites. Due to the

discovery of P. batesii maconensis in this region in June, 1992, all regionally encountered Phyciodes were

given at least some visual attention during the maconensis May - June flight period and 1992 to 1998

research period. It was thus surprising on 16 May 2002 to discover a cocyta like Phyciodes phenotype

never before personally encountered in the southern Appalachian region, and to do so within the P. batesii

maconensis Jones Knob type locality area of Macon County, NC.

The area was a small narrow ridge top meadow surrounded by hardwood forest. This area had not

been previously investigated. It is a man made meadow which exists as a remnant of an old apple orchard

(determined by size/condition of apple trees). Both tharos and this large, red clubbed, cocyta looking

entity were equally common on 16 May 2002. They were fairly easy to determine even in flight by size

and flight pattern: tharos having a lower, faster and more zigzag flight. P. tharos tended more to the

central meadow area while the large orange clubbed entity flew more to the margins, esp. females. P.

batesii maconensis was also present but uncommon. In addition, a single Chlosyne gorgone (Hubner,

1810) was found that day for a new county record and only the second record for the state of North

Carolina. This all indicated this small site to be rich in regionally unusual and rare butterflies. These

butterflies would have simply been identified as P. cocyta had they been found later in the year - late June

through August. But mid May is at least a month before the beginning of the cocyta flight period.

After this initial find, intensive searching was conducted in 2002, 2003 and spring of 2004 to

locate new colonies from north Georgia through western North Carolina into extreme southwestern

Virginia. Dozens of areas were searched but only two other colonies located, both in Clay County, NC.

A third colony was discovered 18 August 2002 in north Georgia in Union County when Alan

Wormington visited the area. Wormington’s Georgian discovery and observations are significant in light

of his research of Ontario Phyciodes. Wormington provided a copy, as follows, of his full report to the

Southern Lepidopterists News (SLN) editor detailing his discovery and determination of this population.

“On August 18 I drove along Duncan Ridge Road (about 15 miles?) which is a gravel hne which starts at

Hwy. 180 in Union County, Georgia. There were hundreds of crescents here and of dozens closely examined, all

were definitely Orange (=Northern) Crescent (cocyta). According to various books, the species should not be

present within 200-300 (?) miles of this area. Elsewhere on my trip (at lower elevations) I saw other crescents that

were obviously tharos.

“Here in southern Ontario, I closely scrutinize all crescents as I have isolated a third "species" that is not

recognized in the scientific literature. So I am quite familiar with crescents in a broad sense. The cocyta that I saw

in Georgia looked no different than the cocyta in Ontario — same for the tharos .”

By the time this record was published in the SLN, the cocyta- like Jones Knob entity had also been

found at two sites in Clay County, NC and observed to be multiple brooded by rearing and wild caught

specimens. Thus, when Wormington’s record appeared in the SLN, the immediate suspicion here was that

this was actually another colony of this cocyta- like taxon and not P. cocyta. This was confirmed in May

of 2003 when I found it to also occur commonly at the Duncan Ridge Rd. site in a spring brood. This

brood began to emerge May 2 nd and was in full flight by May 10. P. tharos was already present in this

area on April 15. These four cocyta- like populations have now been demonstrated to not be species

cocyta, diminutor or tharos by collecting of adults, rearing families from all sites of sympatric taxa, and

by mtDNA sequencing of sympatric specimens of both species.

13

Sites and Occurrence

The four sites new species Phyciodes incognitus has been found to occur at are: Jones Knob, 4000

ft., Macon County, NC; Buck Creek, 3200 ft., Clay County, NC; Sally Gap Rd., 2200, Clay County, NC;

Duncan Ridge Rd., 3700 ft., Union County, GA. At Jones Knob, Sally Gap, and Duncan Ridge the

species is common and found in narrow openings in rather day hardwood forest. At Buck Creek, it is

uncommon and found either as a stray into more open areas or (more commonly) at the edges of

hardwood forest or specific micro areas (i.e Jeep trails through forest).

At Buck Creek, P. tharos is very dominant. This is an extensive generally open Serpentine area.

At Sally Gap and Jones Knob, both species are found in equal numbers. Both areas are long

narrow “cuts” through mature forest. Jones Knob is a ridge meadow, Sally Gap a gravel road. However,

at the Sally Gap road location, P. incognitus is only found in one 300 ft. section going up a hill.

At Duncan Ridge Road, new species P. incognitus is very dominant. P. tharos is quite uncommon

in this area and usually found in the few more open areas the road passes through. This is a narrow,

shaded, one lane jeep trail often going along the side of the mountain with steep slopes on either side of

the roadway. P. tharos is common on Hwy 180 leading to Duncan Ridge Road, especially at the open

base areas at Vogel State Park. P. incognitus is absent from these lower open areas along the highway.

From May of 2002 to July of 2004 the following counties were searched extensively for additional

colonies of P. incognitus. Rabun and Union counties, GA; Oconee County, SC; Grayson and Carroll

counties, VA; Clay, Macon, Avery, Yancy, Haywood, Watauga, and Alleghany counties, NC. A few

other counties were spot checked (stop and scan immediate area) while driving to / from main search

areas. Field research dates were as follows: 2002: 18 April, 7 May, 16 May, 24-25 May, 2 Aug.; 2003:

14-15 April, 26 April, 2 May, 9-10 May, 27 May, 2 June, 5 Aug., 12-15 Aug., 21-22 Aug., 5 Sept.; 2004:

19-20 April, 20-21 May, 8-9 June, 28 June, 8 July. In the mountain region, P. tharos begins its flight

between the end of March and mid April depending on elevation. P. tharos is more or less continual until

frost. P. incognitus begins its flight between first and mid May depending on elevation. P. incognitus is

mostly absent by the first week in July and does not appear again in numbers until late July and becoming

common again by mid August with stragglers through September.

This phenology helps explain why P. incognitus has been missed by collectors in the historically

well known, long, and heavily collected region of Duncan Ridge Road. (The overall region is known as

Cooper’s Creek.) Most collectors do not venture into this area until the first of July when they are seeking

Speyeria diana (Cramer, 1775), Erora laeta (W.H. Edwards, 1862) and other choice species found there.

At that time, P. incognitus is virtually absent except for rare out of season solitary emerging individuals.

Further, it is likely that its similarity to the common P. tharos would contribute to its being ignored by

visiting lepidopterists interested in uncommon to rare taxa at that time.

Three other sites have been identified that appear to harbor populations of P. incognitus.

1) The historical site of the junction of the Kanawha River and Cabin Creek at Coalburg, West

Virginia. At least one and possibly three of Edwards’ specimens sent for study are of this taxon.

2) The figures in Butterflies of Ohio (1992) of P. pascoensis (=cocyta) on Plate 29 from Elk

County, PA are, with high likelihood, P. incognitus. These specimens are of the incognitus phenotype

and lack the broad “smeared” dorsal patches of cocyta. Thus, P. incognitus ranges north well into the

range of both P. diminutor and P. cocyta. Also, the 5 th row figures on Plate 28 of this book are species P.

diminutor not tharos f. marcia as indicated by smeared pattern, dates, small size, and red nudums.

3) Ted Wilcox of North Carolina has confirmed (pers. comm.) that a colony of P. incognitus

occurs in Watauga County, NC. Ted stated: “The site is located in southwest Watauga County, NC near

Avery County, NC and the Tennessee state line. Mostly wooded area with a narrow driveway with grassy

area on each side. Elevation 3500ft.” Ted sent several diagnostic photos of specimens from this

population (header figs., page 1). Photos taken 9 August 04.

14

Rearing

At least one family of both P. tharos and P. incognitus were each reared from Macon, Clay, and

Union counties. Females of both species were collected the same day and same spot for each comparative

rearing. 100% (n=300 +/-) of reared P. incognitus had fully orange to red nudums in both sexes (Figs. D

& E). Five males had dark red nudum. 100% (n=140 +/-) of all reared P. tharos males had black nudums

(Fig. A) with about 15% of those having orange-red tipped clubs. Approximately 70% of reared P. tharos

females (n=200 +/-) had partially orange nudums (Fig. C), 25% all black nudums (Figs. B), and 5% all

orange nudums. Both all black and all orange nudum female tharos were utilized as parent stock. Parent

female club color had no relativity to female nudum color or male % of red tips in reared offspring.

All specimens of both species were reared under identical conditions and on the same species of

Aster (not determined). The first rearing of each species was conducted under continual light to prevent

diapause. All subsequent rearing was under natural photo period (reared in north facing windows). No

larvae of either species entered diapause from May through August rearings. All rearings were in an air-

conditioned environment with a consistent temperature of 75 to 78 degrees. The glass rearing containers

(small baby food jars for early instars and gallon jars for final instars and pupation) were kept

unventilated to insure high humidity (+90%). In the wild, this temp would not be much higher during the

day but would be much lower at night. This would make the larvae develop slower in the wild. Humidity

in their natural mountain habitat would be about the same. Food was changed daily.

In captivity, the minimum time from egg to adult was 35 days for each species. P. tharos tended

to have a more extended developmental period - maximum of 59 days with about 20% of immatures

taking prolonged developmental time. The P. incognitus egg to adult maximum was 50 days with only

sporadic individuals taking prolonged development. This reflected the occurrence in nature of P.

incognitus having two more or less well defined broods while P. tharos tends to occur in long overlapping

ill-defined brood waves. The lowest numbers of P. tharos tend to occur in July (at locations over 3000’).

The larvae of both species are very similar in all instars except for the shape and markings on the

mature head capsules (Figs. 1-9). In P. incognitus , the head is frontally flatter and from the frontal view

looks rectangular (height greater than width) or egg shaped; with less extensive white markings about the

eye and toward the lower area of the frontoclypeus (Figs. 1-2, 5-6 left). In P. tharos , the head is frontally

more bulbous and rounded in outline; with extensive markings about eyes and up aside the frontoclypeus

(Figs. 3-4, 6 right, 7-9). The sum effect is that the heads are diagnostic for species determination. Note

the similarity of Edwards’ BNA drawing of a tharos larval head (Fig. 8) and the specimen in Fig. 9.

The pupae vary greatly in color in both P. tharos and P. incognitus (Figs. L-P) in the region, and

to the same degree. ( P. tharos also has a blackish gray morph (rare) and a pale morph (uncommon) as in

Figs. N & O.) However, in P. tharos the vast majority of all pupae from 4 families were form M, while in

5 families of P. incognitus , the majority of pupae were form O. The pupae of P. tharos seemed “rougher”

on the thorax than in P. incognitus (Figs. Mb & Pb). No definitive pupal color or morphological character

differentiating the two species has been determined - only generalities.

All reared tharos (Figs. 40-41) were smaller than reared incognitus (Fig. 38) (as they are in

nature). All reared specimens of both species had the VHW marginal dark patch correspondingly darker

than in their wild counterparts. All reared specimens were very uniform in their respective phenotypes for

each species. This was surprising for tharos as it has a wide variety of co-occurring dorsal pattern

phenotypes in the wild. Note that figures 17-19, 22-24 were all caught the same day and place. All of

these specimens were also sent to Wahlberg for mtDNA sequencing who reported that their sequences all

nested within tharos. The variation contrast between wild and reared tharos poses the possibility that

another undescribed tharos like species is present in this region. This is especially true at the Buck Creek

location where a “ tharos ” (Fig. 24) is found throughout the season with heavily patterned dorsal wings as

in P. diminutor (Fig. 25) but with all black clubs and tharos mtDNA.

15

Uniformity of captive rearing conditions alone would not account for the lack of this phenotypic

variation in tharos. This is because in the wild all simultaneous broods are also subject to identical

(though different from captive) conditions. It is not logical that large numbers of similarly fresh, or worn,

wild individuals of the same species would be found at the same place and time with notably different

markings, yet have this variation non existent in reared families. What has been found in many rearings

(by this worker) of the Phyciodes species in western North Carolina, is that all three taxa ( maconensis,

tharos, and incognitus) produce very stable phenotypes rearing after rearing (family after family) for each

species.

Further, we now know that the many phenotypes reared and figured by Edwards are partially due

to his having reared multiple species and not “merely” seasonal forms of one species. Thus, it has never

been demonstrated that rearing single eggs clusters will yield very different dorsally marked tharos

individuals. (Edwards’ obtained dorsal variants by placing some pupae “on ice”, which yields aberrations,

not natural forms.)

With the recognition of species tharos, diminutor, cocyta and incognitus in the East, it is now

evident that all that was considered u tharos variation” by past generations of lepidopterists was a severely

flawed assessment of the Eastern Phyciodes. Hopefully, this old mindset will not be hard to discard.

Today, the occurrence of phenotypically different sympatric tharos-group populations should be viewed

with an inclination that sibling species may be present. This is complicated by the fact that we now know

that some species in this group have similar mtDNA (i.e. diminutor and cocyta , and tharos and

incognitus). All of this emphasizes Scott’s 1998 position that rearing is essential to correctly determining

the taxonomy of the tharos-group.

Independent Analysis

(Single vs. Multi-tool Analysis)

Twelve specimens of P. incognitus were sent to Wahlberg for mtDNA sequencing. They were

from various sites and dates. All of these individuals had COI sequences similar or identical to P. tharos.

As a molecular biologist, Wahlberg determined these specimens as P. tharos regardless of their larger

size, different phenotype, different morphology, and sympatry. However, when later presented with the

rearing data, he also acknowledged the potential of a genetically cryptic species. Photos of larvae, pupae

and adults of P. incognitus were sent to Scott for his assessment. As a taxonomist, Scott determined these

same specimens as P. cocyta. In this we see how specialists tend to render conclusions out of their own

field of expertise. Without any other data, their conclusions (while opposite) would both be considered

correct from the limited perspective of single tool analysis.

But when assessed with multi tool analysis, this taxon was evidenced to be neither species cocyta

or tharos - thus exposing both single-tool based conclusions as incorrect. One element of the multi-tool

analysis was Wahlberg’s mtDNA sequencing which confirmed that this Appalachian taxon is not species

cocyta. Another element was Scott’s determination (which included knowledge of rearing results) that

this taxon is not species tharos. In these conclusions, they were both correct. The third element was the

consistent rearing data (including larval head morphology). The fourth was sympatry, but restricted to

select locations - tharos being found almost everywhere but incognitus restricted to forest ecotones and,

so far, at only a few sites from north GA to PA (suspected).

One specimen each of P. tharos and P. incognitus was sent to David Wright for dissection and

photography of genitalia (Figs. F-K), and his opinion. His observations were:

“On the balance, after studying the male genitalic mounts under the microscope and as digital images, I can

not find any significant differences between your Phyciodes tharos and new species. That may be an important

finding, indicating that the n. sp. is closely related to tharos , at least closer to tharos than to batesii or any stray

western taxon. In summary, these findings place these two species (tharos and n. sp.) in the tharos-group,

according to Scott (1994), and closest (or identical) to the tharos-cocyta subgroup.”

16

Figures F pharos) and G ( incognitus ) are dorsal views of the tegumen and the two gnathos hooks.

Figures H and K are the aedeagus of incognitus , with K being a magnification of its strongly toothed

terminal tip. Figures I ( tharos ) and J ( incognitus ) are of the medial view of the valve. Personally, I find

the differences in overall shape of the structure, number and positions of hairs worthy of further

investigation. In particular, the tuft of hairs on the lower valve stem of tharos which is absent in

incognitus. (These printed images can be greatly magnified (zoomed) in the CD version of this paper.)

At high magnification, several minute differences can be noted in Figs. I & J. The question of whether

17

I wish to express special thanks to the following persons without whom this research and paper

could not have been accomplished: Niklas Wahlberg, James Scott, David Wright, and Norbert Kondla. I

also thank Alan Wormington and Ted Wilcox for their contributions of field data which greatly added to

this paper. I thank my son-in-law, Joseph V. Mueller, for his continued assistance in many areas

including photography and all things “computer”. Last but not least, I thank my wife of 38 years,

Marilyn, for allowing me to spend so much money and time on my “hobby”.

LITERATURE CITED

nae. Trans. Amer. Ento. Soc. 92, pgs. 357-468.

, W.H. 1868. Notes on a Remarkable Variety of Papilio turnus, ai

P 1884. The Butterflies of North America. Vol. 2, 357 pgs. Houghtc

1, J.W.H. 2002. On the use of Genetic Divergent

Society. 75, pgs. 509-516.

, K.E. OMLAND. 2003. Species-Level Paraphyly and Polyphyly: Frequency, Causes.

idrial DNA. Annual Reviews Evol. Syst. 34. pgs. 3«

GATRELLE, R.R. 1998. Two new Nym

3, pgs 1-8.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International Code of

Zoological Nomenclature, Lourth Edition (1 January 2000). London, UK: The International Trust for

Zoological Nomenclature. 306 pp.

ILTNER, D.C., J.H. SHUEY & J.V. CALHOUN. 1992. Butterflies and Skippers of Ohio. Ohio St. Univ. (in co-op

with) Ohio Dept, of Nat. Res. Div. of Wildlife and Ohio Lepid. Columbus, Oh. 214 pp.

LAYBERRY, R.A., P.W. HALL, AND J.D. LALONTAINE. 1998. The Butterflies of Canada. Toronto, ON:

University of Toronto Press. 280 pp.

SCOTT, J.A. 1994. Biology and Systematics of Phyciodes (Phyciodes) Papilio (New Series). 7, pgs 1-120.

Published by Author.

_ 1998. Phyciodes (Phyciodes)\ New Discoveries, New SUbspecies, and Convergence. Papilio (New

Series). 10, pgs 1-42. Published by Author.

WAHLBERG, N., R. OLIVEIRA & J.A. SCOTT. 2003. Phylogenetic Relationships of Phyciodes Butterfly

Species (Lepidoptera: Nymphalidae): Complex mtDNA Variation and Species Delimitations. Syst. Ento.

Royal Ento. Soc. 28, pgs. 257-273.

_ 2003. Phylogenetic Relationships of Phyciodes Butterfly Species (Lepidoptera:

Nymphalidae): http://www.zoologi.su.se/research/wahlberg/Phyciodes/phyciodes.htm World Wide Web site

for study’s specimens..

Financial support to TILS research can be given through our web site at http://www.tils-ttr.org

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