taxonID	type	description	language	source
03F587D11461FFD4330CFF6FFB2E6309.taxon	description	Euphydryas phaetoneta. Herrich-Schäffer, G. A. W. (1865) misspelled the species as Melitaea phaedon, a synonym. Holland (1889) misspelled the species as Melitaea phaëtona, a synonym.	en	Pavulaan, Harry (2021): Reevaluation of the described subspecies of Euphydryas phaeton (Drury, 1773) with a replacement name for Melitaea phaeton schausi (Clark, 1927). The Taxonomic Report of the International Lepidoptera Survey 9 (10): 1-21
03F587D11467FFD23343FF52FB916581.taxon	description	“ Characters. - Closely resembling E. phaëton phaëton (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 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).	en	Pavulaan, Harry (2021): Reevaluation of the described subspecies of Euphydryas phaeton (Drury, 1773) with a replacement name for Melitaea phaeton schausi (Clark, 1927). The Taxonomic Report of the International Lepidoptera Survey 9 (10): 1-21
03F587D11466FFD13017FDC2FF2267FC.taxon	description	“ 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 a name. 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 phaëton. 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. phaëton phaëton for Mississippi. Forbes (1960) recognized both ssp. phaëton 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).	en	Pavulaan, Harry (2021): Reevaluation of the described subspecies of Euphydryas phaeton (Drury, 1773) with a replacement name for Melitaea phaeton schausi (Clark, 1927). The Taxonomic Report of the International Lepidoptera Survey 9 (10): 1-21
03F587D11465FFDE3352FC56FEB16057.taxon	description	“ 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. phaëton 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. 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, 2004 a, 2004 b, 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).	en	Pavulaan, Harry (2021): Reevaluation of the described subspecies of Euphydryas phaeton (Drury, 1773) with a replacement name for Melitaea phaeton schausi (Clark, 1927). The Taxonomic Report of the International Lepidoptera Survey 9 (10): 1-21
03F587D11469FFDA30FFF9ACFCAB60A1.taxon	description	Comparison of the four described subspecies 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.	en	Pavulaan, Harry (2021): Reevaluation of the described subspecies of Euphydryas phaeton (Drury, 1773) with a replacement name for Melitaea phaeton schausi (Clark, 1927). The Taxonomic Report of the International Lepidoptera Survey 9 (10): 1-21
03F587D1146DFFD931A0FA64FB0B63F6.taxon	description	= phaetona Holland (1889) [misspelling] = superba (Strecker, 1878) [ref. Pelham (2008), # 699 a, described as “ variety ”] = phaethusa (Hulst, 1881) [ref. Pelham (2008), # 699 a, described as “ aberrant ”] = streckeri (Ellsworth, 1902) [ref. Pelham (2008), # 699 a, described as “ aberration or variety ”]	en	Pavulaan, Harry (2021): Reevaluation of the described subspecies of Euphydryas phaeton (Drury, 1773) with a replacement name for Melitaea phaeton schausi (Clark, 1927). The Taxonomic Report of the International Lepidoptera Survey 9 (10): 1-21
