Volume 9, Number 10 31 December 2021

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

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

Reevaluation of the described subspecies of

Euphydryas phaeton (Drury, 1773) with a replacement

name for Melitaea phaeton schausi (Clark, 1927).

Harry Pavulaan

606 Hunton Place NE

Leesburg, Virginia, 20176, U.S.A.

ABSTRACT. The present paper reevaluates the subspecific standing of the nominotypical and three described subspecies of

Euphydryas phaeton (Drury, 1773). The nominotypical subspecies phaeton occupies the mid-Atlantic region, with undefined

zones of contact with described subspecies borealis (F. Chermock & R. Chermock, 1940) to the north, and subspecies schausi

(Clark, 1927) to the south. Nominotypical phaeton is an intermediate phenotype between borealis and schausi, which are each

noticeably different from each other but both reasonably similar to intermediate (nominotypical) phaeton. Both borealis and

schausi were synonymized under nominotypical phaeton since about time of their descriptions, by authors and list makers who

did not justify their reasoning for essentially ignoring the original descriptions. The common belief is that there is no phenotypic

difference between the three described northeastern subspecies and all are treated as nominotypical E. p. phaeton. In the present

analysis, recently described subspecies ozarkae (Masters, 1968) bears a striking resemblance to schausi, making delineation of

the zone of contact between the two very difficult, other than by habitat and primary host preference of each. Subspecies

schausi, having been originally described within the genus Melitaea, is preoccupied by the name Melitaea schausi (Godman

& Salvin, 1901), presently considered a junior synonym of Chlosyne definita definita. Thus, a replacement name for schausi

(Clark, 1927) is necessary.

INTRODUCTION

A reevaluation of the four described subspecies of Euphydryas phaeton is presented here. Two of

the subspecies, schausi and borealis, while each described in accurate detail, were subsequently ignored

by many authors and list makers, who did not justify their reasoning for essentially ignoring the original

descriptions, thus giving one a clear picture of “armchair taxonomy” and how it affects future analyses of

lepidoptera, and no doubt other organisms. Each is evaluated here, based first on the taxon’s original

description, followed by the interpretations of authors and list makers of the time. A new analysis is

finally presented that shows convincing differences between southeastern subspecies schausi and northern

subspecies borealis. Nominotypical phaeton is an intermediate phenotype between the southern and

northern subspecies, making it very similar to each when only compared to either southern or northern

populations individually. Only when one compares schausi to borealis, are differences very evident. An

interesting issue arises when comparing ozarkae to schausi. Careful comparison of both phenotypes

shows a remarkable similarity. No new taxonomic realignments are proposed here in this regard.

Melitaea phaeton (Drury, 1773)

Melitaea phaeton was originally illustrated as a drawing in Illustrations of Exotic Entomology,

Vol. 1 (Drury, 1773: plate 21) (Fig. 2) along with a description and location (New York) on pages 42-43

(Fig. 1). Drury opted to not apply the Linnean name system, but simply referenced “phaeton” in the

Index, thus technically leaving the illustrated species in Vol. 1 unnamed. An Index to the First Volume

was published with Vol. 2 (Drury, 1773) (Calhoun, pers. corr.). That Index included the binomial names

of specimens illustrated in Vol. 1, thus making the date of description 1773 per ICZN Opinion 474 (ICZN,

1957). The name Melitaea phaeton is found along with the text in the Westwood Edition of Illustrations

of Exotic Entomology, Vol. 1 (Westwood, 1837: page 39) (Fig. 3). Westwood references Plate 21 from

the original Vol. 1 of Drury (1773); also appropriately numbered as Plate 21 in the Westwood Edition.

The illustration fairly well matches typical specimens of Euphydryas phaeton from the Mid-Atlantic

region centered around New York City. The precise origin of the original specimen illustrated by Drury

is unknown, but was likely collected by his correspondent in the New World, Thomas James, who lived

in Brooklyn, N.Y. and frequently sent specimens to Drury in England. Thus, the specimen that served for

the original illustration was most likely collected in the rural western end of Long Island. See discussion

in Calhoun (2010) and also Pavulaan (2020) for details and circumstances surrounding Drury’s collection

and personal contacts. Interestingly, the illustrated type in Drury (1773) is more aligned with the schausi

phenotype. Assuming it was collected in New York, it represents a variant. [This paper will not attempt

to refine the original TL. ]

Fig. III. and 1V. Expands three inches and al- different fhapes and fizes, sin all abouttwenty-fix

moft-three quarters.

Upper-jide.—The Antenne, Head, Byes, Back and

Abdsmen, are black.—'T he Superior (rmgs, alfo are

black ; with nine red {pots placed near the exter-

nal edges ; above which are two rows of {pots, of

a lemon colour, with fome other {mall ones placed

near the upper edges. —The fuferier FPings, like-

wife, are black, having eight red {pots placed clofe

to the exterrial edges; the two'next the abdomi-

nal corner, being almoft united. ‘Thife, as alfo

thofe on t.c fuperior wings, being nearly of a tri-

angular form. Above thefe is a row of lemon

coloured fpots, fhaped like crefcents ; and above

that is another row of lemon coloured fpots, that

are nearly round, all placed circularly, anfwering

to the fhape of the wing.—Lhe £dees of all the

Wings, are eve.

Under: fide —The Palpi and Legs, ave red.—The

Superior fF ings, are black, with a red indented

margin. Along the external edge, 1s "a row of

Jemon-coloured crefeents above it. Near the an-

terior edge, about the middle of each wing, are

two larger red (pots, and a lefier ome at the cor-

ner, next the fhoulders. ‘The remainder of the

wings, is {prinkled with lemon culoured [pots, of

diferent

on each.—The Jnferisr ings, are black, havin

‘their-external-edzes margined with triangular re

dpots; above which, is arow of lemon coloured

¢refcents; and-over thefe, is arow of round Je-

mon coloured fpots. “The remaining part of the

wings, is covered with lemon coloured and red

fpots ; the former very:{mall, and the latter:pretty

large, angularly fhaped.

I received it from New York, there my cor-

refpondent allures ine, he has caught them in June

and September; fromwhence he ts indueed to be-

lieve there are two broods in a year.

T have not feen it defcribed in any author.

Fig. 1: Original description of Melitaea phaeton [unnamed] in Illustrations of Exotic Entomology, Vol. 1 (Drury,

1773: pages 42 and 43)

Fig. 2: Original illustration of Melitaea phaeton in IMlustrations of Exotic

Entomology, Vol. 1 (Drury, 1773)

MELITVZEA PHAETON.

Plate AAL. fe. 3, 4,

Unik: Lepideptern. SmeTiox: Dorn, Fasitv: Ayimghoalida, 40,

Gaixte. Melita, Pobre. Argynnis, Jaf. ef Grow, Papilio (Thin. Peetiv.), rary.

MBLITAA Poantes. All subretundatia intecerrimie, niertis, simngularim extine Spr, pagind omg) mbites

fulyo flavoque macolotis, (Expans. Alor. 3 une. 0 lin, tere.)

Syx. Papilio (Dau. Festiv.) Phaeton, Drery, App. c. 2. Pode. Bat. Set. DD. 1. pp. a Vo. 140, Cromer,

ew, WEE C.D. Angynots Pheetontea, Latr. ef God. Bae. Meth, ix. p. S68. Hoielur, HMiet. mat, Lep,

mw. 7. Hf 3

Hagitat: Sew Vork: token in Jone onl joterau lg r, Whence there ar protealsly two bremle in tha WF

Liner Suade, Body al Winigts black. “The anterior peur of the latter with nine red spots placed near

the external edges; above which are two rows of spots, of a lemon colour, with some other small ones

placed near the upper edges. Posterior wings with eight red triangular spots, placed close to the

external elges, Above these is a row of lemon-coloured crescents: sul above that is another row ra |

lemon-coloured round spots. “he wings are entire.

Under Side. Anterior wings black, with a red indented margin; within the external edge is a row

of lemon-coloured crescents, Near the costa, about the middle of each wing, are two larger red spots,

and a smaller one at the corner, next the shoulders, ‘he remainder of the wings is sprinkled with

lemon-olourec spats, of ciflerent shapers and sizes, in all about tw enty=s1X on each, Posterior wings

black, having their external edges marciiedd with triangular rec SpHOLS | above which is a row nil

lemon-coloured crescents; and over these is a row of round lemon-coloured spots. The remaining part

of the WHILE is cowered with lemon-coloured and rec Spats ; the former very small, and the latter rather

large, and angularly shaped.

Fig. 3: Revised description of Melitaea phaeton in II|lustrations of Exotic Entomology, Vol. 1 (Westwood Edition

of Illustrations of Exotic Entomology, Vol. 1 (1837).

For purposes of synonymy, the aberrant forms “superba” (Strecker, 1878) and “phaethusa”

(Hulst, 1881) were each described from specimens taken on Long Island, New York, thus remaining

synonyms of E. p. phaeton. Pelham (2008) misspelled “phaethusa” as “‘phaetusa’’. Aberrant form

“streckeri” (Ellsworth, 1902) was described from a specimen taken in Broome County, N.Y., thus

associated as a synonym of E. p. phaeton. Hiibner (1816) described Melitaea phaétaena, by description

apparently an aberrant form of phaeton [translated to read: “The wings brick red colored, cheerful yellow

bands and with black lines alternately drawn’’] - which was subsequently misspelled by Barnes &

McDunnough (1917) as Euphydryas phaetana. Godart (1819) misspelled the species as Melitaea

phaetontea, a synonym, which was subsequently misspelled by Barnes & McDunnough (1917) as

Euphydryas phaetoneta. Herrich-Schaffer, G. A. W. (1865) misspelled the species as Melitaea phaedon,

a synonym. Holland (1889) misspelled the species as Melitaea phaétona, a synonym.

3

Euphydryas phaeton schausi (Clark, 1927)

In 1927, Austin Clark described new subspecies E. phaéton schausi. He compared nominotypical

specimens of what he referred to as the “southern form” from Maryland and Virginia against specimens

of what he referred to as the “northern form” from New York, Massachusetts and New Jersey. Clark

makes comparisons between specimens from Cabin John, MD and Alexandria, VA (schausi) against

samples from: Stoneham, Lincoln, Weston, and Newtonville, MA; Kendall, NY; and New Jersey (no

locality). He states: “We find no difficulty in distinguishing specimens from New Jersey and northward

from those from the vicinity of Washington. Typical examples of each are very distinct...’’ Clark’s

original description follows:

“Characters.- Closely resembling E. phaéton phaéton (pl. 1, figs. 1-4) from eastern Massachusetts, but with the ground

color of the upper surface of the wings deep velvety black, usually, but not always, duller and more grayish in the

females, instead of blackish brown, and the light markings white instead of light straw yellow; on the fore wings the

orange spots in the middle and at the tip of the cell are usually much reduced and commonly (occasionally in the

northern form) entirely absent; the eight orange spots along the margin of the wing are smaller, due to the broadening

of the band of black scales along the veins between them and a rounding off of their outer angles by an invasion of

black scales; they are frequently very much reduced in size, especially in the females, and may be almost wholly

obliterated by black scales; in the northern form the three apical spots are usually noticeably larger than the others,

extending inward between the veins for a greater distance, but in the southern form these spots may be all of the same

size, as is usual in the females, or they may decrease regularly from the apex posteriorly, as is usual in the males; on

the hind wings there is very seldom any trace of orange except for the submarginal row of spots, which are restricted

by a broadening of the narrow black border of the wings and a heavier development of black scales along the veins,

especially in the females; beneath, the marginal band of orange spots is narrower than in the northern form with a

more deeply crenate inner margin, and the orange markings in the basal half of the hind wings are more or less reduced

by a greater development of black along the veins and an invasion of black on all sides; the light markings on the

under side are also purer white than in the northern form.”

Maximum wing expanse measurements (wingtip-to-wingtip of mounted specimens) of schausi

indicated males (n=99) ranged between 45.0 to 64.0 mm, averaging 52.5 mm; females (n=61) ranged

between 50.4 to 67.8 mm, averaging 60.3 mm. By comparison, males from New Jersey to Massachusetts

(n=17) ranged between 49.4 to 60.0 mm, averaging 54.5 mm; females (n=8) ranged between 54.0 to 69.8

mm, averaging 59.5 mm).

The variety “magnifica” (Clark, 1927) was described from a specimen taken at the schausi TL in

Maryland, thus remaining under the synonymy of E. p. schausi.

Literature Treatment 1929-1940

Clark (1929, 1932) oddly listed Washington D.C. area phaeton as subspecies phaeton only one

year, then again four years, after he described subspecies schausi. It is unclear what course of events led

to this taxonomic change of heart by the author of schausi himself. One possibility is hinted at, in the

Nomenclature section of each paper. Clark indicates in each of the 1929 and 1932 papers that

nomenclature is based on Barnes & Benjamin (1926), which was published prior to the description of

schausi. It might be conjectured that Clark felt obliged to adhere to the most recent major synonymic

checklist, or this adherence was insisted upon by peer reviewers William T. M. Forbes of Cornell

University and William J. Holland of the Carnegie Museum. In a curious comment, Clark (1932) states:

“But whatever the status of the more or less unfamiliar names may be the fact remains that radical

innovations in nomenclature, whether justified or not, are wholly out of place in a local list. The object

of a local list is to make clear the relation of the local fauna to the fauna of the larger area...This can be

done only if in the local list a system of nomenclature is used which is in general agreement with the

4

nomenclature employed in similar lists...”” This change, initiated by Clark himself, is likely the reason

why schausi remained the “forgotten” subspecies and ignored by subsequent authors.

Field (1938) discussed E. phaeton in Kansas and Missouri and noted a phenotypic difference from

northeastern nom. phaeton.

McDunnough (1938) lists E. phaeton and treats schausi as a junior synonym.

The following authors treated phaeton at species level only, for various states and regions: The

Natural History Society of Maryland (1936); Saunders (1932); Wild (1939).

Euphydryas phaeton borealis (F. H. Chermock & R. L. Chermock, 1940)

In 1940, brothers Frank Chermock and Ralph Chermock described new subspecies E. phaéton

borealis. They started their description first with a conclusion:

“In 1927, Dr. Austin H. Clark, recognized two distinct races of E. phaeton, the one a northern race and the other a

southern race. He considered the northern race as typical and redescribed the southern race from Maryland specimens,

calling it schausi. The southern race represents typical phaeton, therefore schausi becomes a synonym of phaeton

leaving the northern race without aname. Dr. Clark, because of our study and extensive material on hand, has advised

us to describe this unnamed northern race.”

One can immediately see here a possible misinterpretation of Clark’s description of “southern”

(schausi) and “northern” (nominotypical phaeton) races by the Chermocks. By “southern” race, Clark

described subspecies schausi from Maryland and Virginia, that differed from the nominotypical phaeton

in the region of New York, New Jersey and Massachusetts, which Clark referred to as the “northern” race.

Clark had correctly given ample discussion of nominotypical phaeton being the “northern” race. The

Chermocks subsequently concluded the southern race (schausi) represented “typical phaeton” (despite the

fact that nominotypical phaeton was described from New York) without real analysis or justification, then

claimed this left the “northern” race without a name, which according to Clark, it did in fact: E. p. phaeton.

The best interpretation I can determine is what the Chermocks might have intended to convey is that they

considered schausi synonymous with nominotypical phaeton over the broad region from Virginia to

Massachusetts, thus representing the “southern” race. However, Clark had not addressed populations

north of Massachusetts, so the Chermocks defined a new geographic region for the “northern race’. The

description of ssp. borealis follows:

“Upper side: the ground color of this race is a jet black, almost glossy, in contrast to the dull sooty black of typical

phaeton; the orange marginal spots of phaeton are replaced by large, almost red markings which form a rather wide

band intersected only by the black veins. The red spots in the cells of both wings are large and pronounced; white

markings similar to phaeton.

Lower surface: The red markings on this surface are again large and very pronounced; white markings similar to

the typical form.

Male wingspread averages about 42 mm.; female wingspread averages about 47 mm. Generally, this race is

smaller than typical phaeton.”

The Chermocks list holotype and paratype locations from the following areas: Enfield, Lincoln,

and Portland, ME; Hamilton, and Mer Bleue, ON; Georgeville, Knowiton, and Lanoroie QC; Baddeck,

NS. This defines, in part, the range of borealis as determined by the Chermocks. Masters (1968) describes

the range as “Maine and Quebec, into the Maritime Provinces — and interestingly, also in western

Wisconsin and Minnesota...separated from the nominate subspecies by a sharp cline.”

Literature Treatment 1941-1968

Clark (1951) interestingly, dropped use of the trinomial name schausi in The Butterflies of

Virginia and simply applied the species name phaéton.

Klots (1951) recognized both subspecies phaeton and borealis, but noted: “These poorly

differentiated subspecies are really statistical gradations in a cline.”

Tietz (1952) lists E. phaeton at species rank for Pennsylvania, with schausi as a synonym.

Furguson (1953) recognized Nova Scotia populations as E. phaeton borealis.

Mather & Mather (1958) list E. phaéton phaéton for Mississippi.

Forbes (1960) recognized both ssp. phaéton for New York and borealis as the northern race.

dos Passos (1964) listed subspecies phaeton and borealis.

The following authors also treated phaeton at species level only, for various states and regions:

Kimball & Jones (1943); Macy & Shepard (1941); Moore (1960), Ehrlich & Ehrlich (1961); Shapiro

(1966).

Euphydryas phaeton ozarkae (Masters, 1968)

Prior to the description of ozarkae, W. Hoffmeister (1881) described larvae in Lee County, IA

(later determined to be ssp. ozarkae) feeding on Aureolaria pedicularia. Field (1938) first noted that

specimens from Kansas and Missouri differed from phaeton in places such as Pennsylvania, New York,

Maine, Wisconsin...” and suggested “Whether this material represents a new subspecies ...has not yet

been determined.” Masters (1968) described subspecies ozarkae from interior North American

populations that inhabit a different habitat type: dry valleys, hillsides, high well-drained hilltops, and

thinly wooded ridges; and that feed primarily on Aureolaria grandiflora. His description follows:

“Male (Figs. 1, 2). — The same general appearance as nominate E. phaeton (Figs. 3, 4) but the red coloring is paler

and of a more yellow cast. With an expanse of one forewing (base to apex) of 28 to 32 mm it is somewhat larger.

Upperside (Fig. 1): Marginal red spots are reduced in size. Black lines over veins are wider and the black marginal

band invades the red band, resulting in a wider spacing of the red spots. Red spots at apex of the forewing tend to be

narrow, in no case are they wider than high. Red spots in forewing cell are not well developed — 75% of specimens

have only one poorly defined spot; in the remainder one spot is weakly developed and there is a faint suggestion of

the second.

Underside (Fig. 2): White coloring tends to be “whiter” than on the nominate subspecies. Discal cluster of red

spots are more broken and separated by black.

The genitalia (Fig. 9) do not differ from the nominate subspecies.

Female (Figs. 5, 6). — The same general appearance as nominate phaeton (Figs. 7, 8) but the red coloring is

reduced — often wanting altogether on upper surfaces — and is of a paler, yellower cast. Very large size — expanse of

one forewing (base to apex) 31 to 38 mm.

Upperside (Fig. 5): Forewing discal red is not present. Marginal red spots, if present, have a distinctly triangular

shape and are reduced in size so that the space between them is as large as the spots themselves. White areas tend to

be larger and “whiter” — four white bands are present on the forewing, fusing to three near the anal angle. Outer row

of white spots are larger than marginal row of red spots on forewing.

Underside (Fig. 6): Discal red pattern appears to be more broken because of the wider separation of the spots.

White rows tend to be wider and more regular.”

At the time of the description of ozarkae, Masters (1968) gave the range of ozarkae as: Springfield,

and vicinity of St. Louis, MO; Brown Co., IL; Lawrence and vic., KS; Ottawa Co., OK; and northern

Arkansas. Masters commented: “While Chelone glabra is found throughout the Ozarks, I never found E.

phaeton in association with it but rather with Aureolaria.” Dole, et al. (2004) indicate the range of phaeton

extending into northeast Texas, and Plantago lanceolata is listed as an additional host for that region.

Schlicht et al. (2007) show ozarkae in extreme southeast Iowa. Interestingly, Harris (1972) writes that all

Georgia specimens to his knowledge were collected on “hillsides and mountain slopes” with the host

unknown and with no evidence of Chelone glabra. This highlights the need for more detailed fieldwork

to define the eastern range of ozarkae. Due to phenotypic similarity to schausi from the present analysis,

the conclusion is that ozarkae can be more reliably defined by habitat and primary hostplant association.

Differentiating populations of ozarkae from schausi in the intervening region of the Ohio River watershed

will rely heavily on host and habitat associations rather than phenotype alone.

Literature Treatment 1969-2021

Harris (1972) refers to Georgia populations as nominotypical E. p. phaeton. [Interestingly, all

cited reports are from upland habitats, suggestive of eastward influence of ozarkae.|

Irwin & Downey (1973) recognized E. p. phaeton and E. p. ozarkae separately as subspecies, and

indicated their separate distributions in Illinois.

Brower (1974) treats Maine populations as E. p. borealis.

Howe (1975) lists E. p. phaeton and E. p. ozarkae at full subspecific rank, but discusses regional

variation in E. phaeton with great clarity: “Through the years several names have been proposed for

variations among northern, central and southern populations. E. phaeton, described from New York, has

the intermediate central coloring and pattern. The name borealis...was given to the northern color

variation with larger, redder marginal spots and glossy jet black coloring above. The name schausi...is

characterized as being blacker in ground color, with whiter light spots and narrower orange markings.

The type locality [phaeton] is in a transitional area.” Under the entry for subspecies E. p. phaeton: “If

names are desired for these variations phaeton, borealis and schausi are available, but...these names do

not represent separate populations, only the two extremes and middle of a cline.”

Opler & Krizek (1984) simply commented: “Several subspecies of uncertain merit have been

proposed. The most valid of these seem to be E. phaeton phaeton and E. phaeton ozarkae Masters. These

two subspecies may be distinguished on the basis of adult coloration, habitat, and food plant.”

Mather & Mather (1976, 1985) list E. phaéton ozarkae for Mississippi (1976), with a

grammatical name correction to phaeton (1985).

Miller & Brown (1981) listed E. p. phaeton and E. p. ozarkae as subspecies; with schausi and

borealis as junior synonyms of ssp. phaeton.

Hodges (1983) listed E. p. phaeton and E. p. ozarkae as subspecies; with schausi and borealis as

junior synonyms of ssp. phaeton.

Sedman & Hess (1985) treat west central Illinois populations as subspecies ozarkae.

Scott (1986) recognized only E. p. phaeton and E. p. ozarkae as subspecies.

7

Vawter & Wright (1986) conducted a study of genetic differentiation between E. p. phaeton and

E. p. ozarkae and found a lack of allozyme differentiation between New York and Missouri population

samples. They concluded that populations so genetically similar are unlikely to be separate species. The

authors erroneously stated that only “two subspecies have been described’.

Heitzman & Heitzman (1987) treat Missouri populations as subspecies ozarkae.

Shull (1987) recognized E. p. phaeton and E. p. ozarkae separately as subspecies, and noted that

only nominotypical phaeton has been found in Indiana

Klassen, et al. (1989) suggest that Manitoba populations are nominotypical E. p. phaeton, but

state: “The number of Baltimore subspecies is still under investigation.”

Ferris, C. D. (1989) listed E. p. phaeton and E. p. ozarkae as subspecies.

Iftner, et al. (1992) treat Ohio populations as nominotypical E. p. phaeton.

Miller (1992) recognized two subspecies, E. p. phaeton and E. p. ozarkae.

Poole & Gentili (1996) do not recognize subspecies for E. phaeton, and list schausi, borealis and

ozarkae as junior synonyms.

Neck (1996) indicates the Texas records are subspecies ozarkae.

Allen (1997) treats West Virginia populations as nominotypical E. p. phaeton. However, the

specimens illustrated from Elkins (plate 15, row 5) align with the schausi phenotype.

Layberry, et al. (1998) state that “only the nom nominotypical inate subspecies is found in

Canada” and do not recognize borealis. However, the specimen illustrated from Ottawa (plate 15, no. 28)

is clearly borealis.

Bouseman & Sternburg (2001) recognized E. p. phaeton and E. p. ozarkae separately as

subspecies, and indicated their separate distributions in Illinois.

Cech & Tudor (2005) treat E. phaeton at species rank and comment: “Baltimores living in dry,

upland forest of the Ozark Mountains were formerly considered a separate race...but more recent

investigations failed to support this distinction (Vawter & Wright, 1986). Indeed, “Ozark-like” upland

populations are also now known from New England and New York.” [The authors clearly do not

recognize phenotypic differences for E. phaeton as qualifying for subspecific status, and no subsequent

study has been done on purported dry-habitat phaeton in the northeast other than anecdotal references. |

Schlicht, et al. (2007) recognized E. p. phaeton and E. p. ozarkae separately as subspecies, and

indicated their separate distributions in Iowa.

Scott (2008) recognizes only subspecies E. p. phaeton and E. p. ozarkae, then lists borealis as a

synonym of E. p. phaeton, and does not recognize schausi.

Belth (2013) recognized E. p. phaeton and E. p. ozarkae separately as subspecies, and noted that

only nominotypical phaeton has been found in Indiana, but that ozarkae may eventually be found in

southern Indiana.

Spencer (2014) treats Arkansas populations as subspecies ozarkae.

Jeffords, et al. (2014) recognized E. p. phaeton and E. p. ozarkae separately as subspecies, and

indicated their separate distributions in Illinois.

Monroe & Wright (2017) recognize Pennsylvania populations as nominotypical subspecies E. p.

phaeton.

Pohl, et al. (2018) list only subspecies E. p. phaeton for Canada.

The following authors treat phaeton at species level only, for various states and regions: Acorn &

Sheldon (2016); Allard (2013); Allen, et al. (2005); Betros (2008); Blakney (2015); Blakney & Gallagher

(2020); Brock & Kaufman (2003); Carmichael & Vance (2003); Cossey (2016, 2017); Covell (1999);

Daniels (2003, 2004a, 2004b, 2005); Douglas & Douglas (2005); Ebner (1970); Ely, et al. (1986); Feltwell

& Hargreaves (1992); Glassberg (1993, 1999, 2017); Gochfeld & Burger (1997); Grehan, et al. (1995);

Hall, et al. (2014); Handfield (2011); Holmes, et al. (1991); Howell & Charny (2010); Jones & Schaeffer

(2012); Kiel (2003); Kimball & Jones (1943); Leboeuf & Le Tirant (2012); Mello & Hansen (2004);

Nielsen (1999); O’Donnell, et al. (2007); Ogard & Bright (2010); Opler & Malikul (1998); Patterson

(2011); Pyle (1981); Riotte (1992); Shapiro (1974); Shapiro & Shapiro (1973); Smith & Domingue

(2019); Stichter (2015); Veilleux & Prévost (1976); Venable (2014); Wagner (2005); Weber (2002, 2006);

Woodbury (1994).

COMMENTS ON DESCRIBED E. PHAETON SUBSPECIES

The recent traditional treatment has been to recognize either phaeton at species rank only (mainly

for publications covering regions in the north and east), while others include ozarkae as distinct for its life

history aspects. There is scant mention of borealis at subspecies rank in the literature (Klots, 1951;

Ferguson, 1953; Forbes, 1960; dos Passos, 1964; Brower, 1974; Howe, 1975). What is interesting to note

is that subspecies schausi has been nearly completely ignored by authors subsequent to its description by

Clark in 1927 (with the exception of discussion in Howe, 1975) and considered simply part of the

northeastern nominotypical subspecies populations. Subspecies ozarkae, was recognized immediately by

authors after its original description by Masters in 1968, and despite habitat and primary host differences

from eastern E. phaeton populations, appears phenotypically similar to subspecies schausi. Southern

Appalachian Mountain records of phaeton are phenotypically closer to ozarkae. Harris (1972) describes

dry upland populations in Georgia which might be considered ozarkae. Were it not for habitat and host

differences, the two might even be considered consubspecific based on phenotype alone. An unresolved

issue is the lack of published habitat, host and life history observations specific to the intervening region

between the Appalachian Mountains and the Ozark Region. Presently, there is no information suggesting

where schausi grades or transitions over to ozarkae. Images of E. phaeton photographed in Tennessee

that are posted to butterfliesandmoths.org show distinct ozarkae phenotypes throughout much of that state,

but host and habitat information are lacking. Several observations in the eastern United States indicate

that isolated dry, upland populations that feed on Aureolaria occur near Chelone glabra in wet habitats in

the surrounding region with no evidence of feeding on Chelone glabra, thus suggesting that, at least,

nominotypical phaeton and subspecies schausi may be capable of adapting to habitat and host changes

9

that would be suggestive of ozarkae. These have been reported from Connecticut (Saunders, 1932) and

western North Carolina.

HOSTS

E. phaeton of the northeastern U.S. and eastern Canada is historically known to dwell primarily in

marshy habitats, wet meadows, brushy swamps, fens, bogs, sphagnum bogs, boggy ditches, boggy swales,

mesic pastures, poorly drained pastureland, open woodland seeps, oak-pine barrens, streamsides and lake

edges where the primary host Chelone glabra (White Turtlehead) occurs. Host use of Chelone glabra

was first reported by W. H. Edwards (1884), in West Virginia, then by Scudder (1889), in Massachusetts.

Later observations by multiple authors reported that larvae have been found on the secondary hosts Aster

sp. (Aster), Aureolaria flava (Smooth Yellow False Foxglove), Aureolaria grandiflora (Largeflower False

Foxglove), Aureolaria pedicularia (Fernleaf False Foxglove), Camissonia campestris (=Oenothera

dentata var. parishii) (Mojave Suncup), Corylus sp. (Hazelnut), Crataegus acrosperma (Bigfruit

Hawthorn), Dasistoma macrophylla (Mullein Foxglove), Fraxinus americana (=biltmoreana) (White

Ash), Fraxinus pennsylvanica (Green Ash), Galeopsis tetrahit (Brittlestem Hemp Nettle), Lonicera

canadensis (=ciliata) (American Fly Honeysuckle), Lonicera japonica (Japanese Honeysuckle), Lonicera

oblongifolia (Swamp Fly Honeysuckle), Lonicera tatarica (Tatarian Honeysuckle), Lonicera xylosteum

(Dwarf Honeysuckle), Mimulus ringens (Allegheny or Square-stemmed Monkey Flower), Pedicularis

canadensis (Canadian Lousewort or Wood Betony), Penstemon digitalis (Foxglove Beardtongue),

Penstemon hirsutus (Hairy Beardtongue), Plantago lanceolata (English or Narrowleaf Plantain), Plantago

rugelii (Pale Plantain), Rhinanthus minor (=crista-galli) (Little Yellow Rattle), Ribes nigrum (European

Black Currant), Sagitaria sp. (Arrowhead), Salix sp. (Willow), Scrophularia marilandica (Carpenter’s

Square), Scrophularia nodosa (Woodland Figwort), Solidago sp. (Goldenrod), Symphoricarpos albus

(Common Snowberry), Symphoricarpos orbiculatus (Coralberry), Typha latifolia (Broadleaf Cattail),

Valeriana edulis var. ciliata (Tobacco Root), Valerianella radiata (Beaked Corn Salad), Verbesina

alternifolia (Wingstem), Veronica sp. (Speedwell), Viburnum dentatum (Southern Arrowwood),

Viburnum opulus (=trilobum) (American Cranberry Bush), and Viburnum recognitum (Smooth

Arrowwood). Some of the listed hosts might be in error, misidentified, or larvae were simply found on

them, but not feeding. Southern New England populations have recently switched their primary host to

Plantago lanceolata, with an associated switch to dry, open field habitats; resulting in frequent explosive

population irruptions. Masters (1968) first documented populations in the U.S. interior feeding on

Aureolaria grandiflora, which he described as subspecies ozarkae. Interestingly there is an account of

larvae selecting Aureolaria flava on a “high, dry rocky ridge” in Connecticut (Saunders, 1932; O’ Donnell,

et al., 2007) but this was never further researched. Clark (1927) indicated that captive larvae will not

accept Wisteria (Wisteria). Saunders (1932) indicates captive larvae will not eat Viburnum plicatum var.

tomentosum (Japanese Snowball).

TAXONOMY

Euphydryas phaeton clarki Pavulaan, 2021 nomen novum

The subspecific name Euphydryas phaeton clarki is proposed to replace Euphydryas phaeton

schausi (Clark, 1927), preoccupied by Melitaea schausi (Godman & Salvin, 1901) which is presently

considered a subjective synonym of Chlosyne definita definita (E. Aaron, 1885). The same data (i.e.,

holotype, type locality) from the description of E. p. schausi (Clark, 1927) applies to clarki (I.C.Z.N. Code

Article 60.3). The name clarki recognizes Austin H. Clark, who first described schausi.

10

Comparison of the four described subspecies

Dorsabwing borealis haeton ear ozarkae

characters P =schausi

glossy pure black dark grayish black sooty grayish black

black

FW maximum length males 20-24 mm 21-26 mm 23-29 mm 24-30 mm

FW maximum length females 25-27 mm 26-29 mm 28-34 mm 30-35 mm

FW wing cell, inner orange mark | Enlarged, well-defined | Well developed, very Mostly absent, variable | Mostly absent,

enlarged, round, faded in | with faded edges variable with

some individuals faded edges

FW wing cell, outer orange mark | Enlarged, well-defined | Well developed, variable, | Mostly absent, variable, | Variable, weakly

enlarged irregular shape with faded edges developed

FW marginal spot row alignment | Solid band, separated | Solid band, crenate Variable, mostly very Very reduced,

by black wing veins (toothed) on inner edge, reduced, spots spots separated

separated by black wing separated by wide by black

veins black wing veins

Shape of FW marginal spots Distinctly square Variable, connected U- or | Variable, mostly Very reduced,

V-shapes rectangular, faded at appearing

edges, some with U- rounded, faded

shape at edges

cream tint

FW presence of submedian Only a single white dot | Variable, mostly weakly- Absent but some have | Few white spots,

most

HW wing cell, orange mark ighly variable, mostly | Absent to mostly weakly Absent Absent

weakly present present

HW marginal spot row alignment | Broad, forming a solid | Broad, forming row of Variable, mostly very Much reduced,

band, divided by black | connected U-shapes, reduced, spots divided by broad

wing veins divided by black wing separated by wide black areas

veins black wing veins

Shape of HW marginal spots Filled U-shape. Filled U-shape. Filled U or V-shape Much reduced,

rounded

irregular shape

with faded

edges

HW submarginal spot row Very thin, line-like Broad crescents, shaped Broad crescents, Broad crescents,

crescents. like thickened V shaped like thickened V | many stretched

into V-shapes

HW presence of submedian White spots absent Variable, ghost pattern Mostly absent, ghost Mostly absent,

pattern reflective of venter pattern reflective of ghost pattern in

venter in some some

Fig. 4. Chart comparison of the four described E. phaeton subspecies.

11

Fig. 5. Dorsal phenotypic comparison of the four described E. phaeton subspecies. Males on left, females on right.

Subspecies from top to bottom: borealis (Edmundston, NB), phaeton (Pinelawn, NY), clarki (Harmans, MD),

ozarkae (Sullivan, MO). Printed specimen images are actual size.

12

Conclusion

E. phaeton appears to consist of a broad cline from northeast (borealis) to southwest (ozarkae)

(Fig. 6). The subspecies borealis (Fig. 5) is smallest, characterized by its glossy, pure black dorsal ground

color and sharply-defined, deep red markings. Subspecies ozarkae (Fig. 5) is largest, characterized

primarily by its very reduced deep orange markings, the marginal ones of which are rounded and display

faded edges. Host and habitat presently define this subspecies. Subspecies clarki (Fig. 5) is

phenotypically most similar to ozarkae. Where clarki transitions into ozarkae remains to be studied. The

two subspecies may overlap by their habitat (dry upland vs. wetland) and primary host (Chelone vs.

Aureolaria) choices. However, this may be unreliable as dry upland Aureolaria-associated populations

tentatively identified as ozarkae have been documented in the east, especially in northern Georgia and

Alabama.

Nominotypical subspecies phaeton (Fig. 5) is most similar to borealis but is clearly a transitional

form between borealis and clarki. It is highly variable, characterized by well-developed interior orange

markings. A small percentage of specimens could be assigned to either borealis or clarki. Thus, authors

who simply compare subspecies phaeton to borealis might be tempted to dismiss borealis as nothing more

than a variant, or synonym, of phaeton. Similarly, authors who simply compare subspecies phaeton to

clarki might be tempted to dismiss clarki as nothing more than a variant, or synonym, of phaeton.

However, when comparing subspecies clarki to borealis, the contrasting phenotypes are obvious. Despite

the temptation to synonymize the names of populations within clines, especially transitional phenotypes

in the middle of a cline, does this suggest dropping nominotypical phaeton from usage? Per rules of the

I.C.Z.N., once a nominotypical taxon is described and named, that name permanently applies to that taxon,

even if synonymized. But recognizing the clear differences between populations at the ends of a cline

merits their recognition as named entities.

Species E. phaeton presents a challenge to taxonomists and evolutionary biologists. The

mechanics of a cline in this species calls for further study. Relationships need to be thoroughly studied

among different habitat and host-associated populations.

DISTRIBUTION

The nominotypical subspecies phaeton ranges from southern New England west to Illinois,

primarily north of the Ohio River and into the Great Lakes region (Fig. 6). Subspecies borealis ranges

throughout the Canadian Maritimes, across northern Maine, into far eastern Ontario. Some specimens

from Wisconsin appear to be borealis, but most are phaeton or intermediate to phaeton. In eastern Ontario,

male specimens appear close to borealis, while females appear closer to phaeton. Subspecies ozarkae

ranges from Missouri and southern Illinois, east into Tennessee, Mississippi, Alabama and Georgia, and

includes an isolated population in Texas. This distribution is based primarily on populations inhabiting

dry, upland habitats with no association with Chelone. Subspecies clarki consists of Chelone-associated

populations from Maryland, southward in the Appalachian Mountains. Where clarki and ozarkae meet or

overlap remains problematic. Phenotypically, specimens from the Carolinas and Kentucky are difficult

to assign to either subspecies, and there 1s virtually no life history information from this region. Though

they may be differentiated by habitat and host associations, it remains to be determined what Aureolaria-

feeding populations in this intervening region are properly referred to.

13

ee AY’

a =

ae

, Me iy

Fig. 6. Approximate ranges of E. phaeton subspecies. Red arrows

point to range extensions. Question marks indicate region in question

where clarki transitions into ozarkae.

PROPOSED REVISION

The following revision is proposed. Euphydryas phaeton is divided into four previously described

subspecies with a replacement name for schausi (Clark, 1927). Reference is made to Pelham (2008) with

its original species numbers. Synonymic treatments (subjective synonyms, misspellings, variety and

aberration names) are all preceded by “=” with text in grey tint.

Euphydryas phaeton (Drury, 1773) [ref. Pelham (2008), #699]

Euphydryas phaeton phaeton (Drury, 1773) [ref. Pelham (2008), #699a]

= phaetaena Hiibner (1816) [ref. Pelham (2008), #699a, original description indicates

an aberrant form]

= phaetontea Godart (1819) [misspelling; original description vague and general |

= phaedon Herrich-Schaffer, G. A. W. (1865) [misspelling]

= phaetona Holland (1889) [misspelling]

= superba (Strecker, 1878) [ref. Pelham (2008), #699a, described as “variety”’]

= phaethusa (Hulst, 1881) [ref. Pelham (2008), #699a, described as “aberrant’’]

= streckeri (Ellsworth, 1902) [ref. Pelham (2008), #699a, described as “aberration or

variety”’|

= phaetoneta Barnes & McDunnough (1917) [misspelling of phaetontea Godart

(1819)]

= phaetana Barnes & McDunnough (1917) [misspelling of phaetaena Hiibner (1816)]

= phaetusa Pelham, 2008 [misspelling of “phaethusa” (Hulst, 1881)|

14

Euphydryas phaeton clarki (Pavulaan, 2021) [nomen novum|

= schausi (Clark, 1927) [ref. Pelham (2008), #699a, subjective synonym, preoccupied]

= magnifica (Clark, 1927) [ref. Pelham (2008), #699a, described as a “variety’’]

Euphydryas phaeton borealis (F. H. Chermock & R. L. Chermock, 1940) [reinstated status; ref.

Pelham (2008), #699a, subjective synonym]

Euphydryas phaeton ozarkae (Masters, 1968) [ref. Pelham (2008), #699b]

ACKNOWLEDGEMENTS

Thanks go to David Wright for review; Ricky Patterson for additional review; John Calhoun for

comments and guidance on the historical circumstances of E. phaeton’s original description.

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