Volume 11 Number 4 October 15, 2023

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

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

Examination of the status of Phyciodes tharos distincta

Bauer, 1975, confirming it as a valid subspecies.

Harry Pavulaan

606 Hunton Place NE, Leesburg, Virginia, USA, 20176

ABSTRACT. The subspecific status of Phyciodes tharos distincta is reexamined. A specimen series from

southern Arizona reveals that distincta shows a distinct, unique phenotype dissimilar from eastern North American

nominotypical P. t. tharos. Subspecies distincta occupies a very limited range in extreme southeastern California,

southern Arizona, and northwestern Mexico. A lectotype is designated.

Additional key words: Lectotype, subspecies, RGB and HSB color analysis.

INTRODUCTION

From its original description (Bauer, 1975), Phyciodes tharos distincta was recognized as

a subspecies inhabiting the far southwestern corner of the United States and parts of northwestern

Mexico. In recent treatments on the Phyciodes tharos complex, Scott (1992, 1994, 1998) asserted

that distincta is a mere synonym of eastern, nominotypical P. tharos tharos, or at best, a “weak

subspecies”. Having myself spent a lifetime among hordes of tharos in field habitats throughout

the eastern U.S., I dispute Scott’s assertion that distincta is nothing more than a synonym of

nominotypical tharos. Specimens in my possession from southern Arizona show a unique

phenotype, clearly differentiated from nominotypical tharos by size, ground color and expression

of wing pattern. This paper makes the comparison and demonstrates the difference between

nominotypical tharos and distincta (Fig. 5). The suggestion that distincta is a “weak” subspecies

(Scott, 1994) appears to be one of personal interpretation.

PRIOR PUBLISHED OBSERVATION OF INTEREST

Emmel & Emmel (1973), prior to the description of distincta, provided some interesting

observations on tharos in southeastern California. The authors list tharos instead as “near

pulchella”, noting that Gunder (circa 1930) found the species in the Imperial Valley being

phenotypically different from typical tharos. There are no published records of pulchella in the

Imperial Valley to date, thus this previous reference applies to P. t. distincta. The historical

presence of tharos in the Imperial Valley is attributed to the introduction of irrigation and tharos

doubtfully previously resided in the area’s natural desert habitat.

PHYCIODES THAROS DISTINCTA BAUER (1975), ORIGINAL DESCRIPTION

Miller & Brown (1981) stated, quite eloquently: “This name is another proposed by Bauer

in a rather unsatisfactory manner. In this instance the type-locality is given, but no types are

designated, nor are they figured.” The original description by Bauer (in Howe, 1975) follows:

(b) distincta Bauer. This subspecies has a complete pattern of fine lines on the upper

surface, is very constant in wing pattern above, and rarely has the blotchy, irregular appearance so

common in individuals of tharos tharos. The cool weather form is scarce and appears only during

midwinter over most of its range. Similar individuals appear in colonies of typical tharos, and there

is a broad blend zone of the two populations in Texas and northeastern Mexico.

Distribution: The type locality is Calexico, Imperial county, California. This is a

predominantly Mexican subspecies, ranging south to the Sierra Madre del Sur and occurring in the

United States along the lower Colorado River north to Moab, Utah, and in a blend zone with nominate

tharos from southeastern Arizona into Texas.

A holotype was not designated in the original description of P. tharos distincta (ref. ICZN

Article 73.1.3). The supposed holotype reportedly residing in the collection of the Natural a stony

Museum of Los Angeles County (Ferris, 1989,

Scott, 1998) was not found per Weiping Xie,

Collections Manager, Natural History Museum of

Los Angeles County (pers. comm.), though there is

a unit tray labelled as all “paratypes” (Fig. 1).

However, these cannot be treated as paratypes as the

yellow note indicates they were designated by

Bauer on 4-13-1982; and at least 8 of the males

(rows 2-4 from top) cannot be paratypes as they

were collected during the period of Sept. 12-15,

1982. The older specimens in row | may be

syntypes. No specimens in the unit tray actually

bear “paratype” labels. However, Nick Grishin

(pers. corr.) photographed two specimens labelled

as “paratypes” (Figs. 2 & 3) in the collection of the

McGuire Center for Lepidoptera and Biodiversity.

Discussion with Crispin Guppy (pers. corr.)

suggests that, since no holotype was designated in

the original description of distincta, these are all

syntypes (ICZN Article 73.2) from which a

lectotype can be designated (ICZN Article 74.1).

Fig. 1. Unit tray of P. ¢. distincta in the Natural

History Museum of Los Angeles County. Photo

courtesy of Weiping Xei, Collections Manager of

Entomology.

12) Tunser eal

Ex DD, Baier call

5

pve 21057423

cio Mick 4. Grishin

Fig. 2. Phyciodes tharos distincta male “paratype”. Now designated as lectotype. Photo courtesy Nick Grishin

and McGuire Center for Lepidoptera and Biodiversity.

Fig. 3. Phyciodes tharos distincta female “paratype”. Now designated as paralectotype. Photo courtesy Nick

Grishin and McGuire Center for Lepidoptera and Biodiversity.

I designate the specimen in Fig. 2 as LECTOTYPE of Phyciodes tharos distincta Bauer,

1975 with the following data: California, Imperial County, Calexico, May 5, 1934, leg Lloyd M.

Martin; det. D. L. Bauer; ex-D. Bauer collection via J.D. Turner collection; MGCL Accession

#2010-29; DNA sample ID: NVG-21067H12, c/o Nick V. Grishin. The specimen resides in the

McGuire Center for Lepidoptera and Biodiversity.

SUBSEQUENT (POST-1975) TREATMENT IN THE LITERATURE

Miller & Brown (1981) treated distincta at subspecies rank.

Hodges (1983) treated distincta at subspecies rank but emended the name to distinctus per

the ICZN.

Tilden & Smith (1986) interestingly, did not include P. tharos for the western states.

Rather, they treated populations in southeastern California east to western Texas and Mexico as

subspecies P. pascoensis distinctus.

Ferris (1989), under species entry 161:623c commented: “To conform to the CODE,

emend spelling to: distinctus.” [This emendation has not been adopted by some subsequent

authors.| Ferris treated distinctus at subspecific rank. However, Ferris made an interesting

comment: “In south-central Wyoming and along the Arizona-New Mexico border, there are tharos

populations that will probably prove to be distinct species (C. G. Oliver, in litt.).”

Bailowitz & Brock (1991) treated southeastern Arizona tharos as ssp. tharos, but

commented: “True tharos, which occurs at least in southern Arizona, has its subspecific taxonomy

in confusion as well. Material from the region appears most like nominotypical tharos and less

like distincta Bauer.” They give the habitat as “agricultural and riparian areas, occasionally from

the more wooded canyons”. Early/late flight dates are Mar. 22 to 13 Nov.

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Brown, et al. (1992) treated distincta at subspecies rank.

Scott (1992) stated: “...distincta Bauer (type locality Calexico, Imperial Co., Calif.) is a

synonym of tharos (unless distincta is used for the southern populations with white-and-black

antenna clubs, tharos for northern populations with orange-and-black antenna clubs), and even

tharos from central Mexico do not differ appreciably from SE U.S. tharos.” [The present paper

opts to follow Scott’s assertion: “...distincta is used for the southern populations with white-and-

black antenna clubs’, though the range is more restricted to the SW U.S.].

Scott (1994) stated: “The name distincta Bauer (1975) (figs. 86-89), type loc. Calexico,

Imperial Valley Calif., is a synonym of tharos based on the series of adults I examined. The unf

black spots appear to be a little smaller than typical tharos, and one female (fig. 89) has somewhat

less black ups markings than typical tharos, and one female has the tip of lamella paraostialis a

little more rounded than tharos and more like riocolorado, but distincta is closer to tharos than to

riocolorado in wing pattern and genitalia, so I treat distincta as a synonym of tharos; at best it is a

weak ssp. for splitters.” Scott illustrated “syn. “distincta’”’ with specimens from Calexico and

Brawley, CA., on page 113. [The few male specimens of Utah riocolorado in my possession are

practically identical to ssp. tharos males (Fig. 7), contrary to Scott’s assertion, whereas distincta

is recognizably different from both. See comments below.] Scott, citing Bailowitz & Brock (1991)

listed the host as A. subulatus var. ligulatus in Arizona. However, Bailowitz & Brock (1991) listed

the host as Aster exilis, a nomen dubium revised as Symphyotrichum divaricatum (USDA, 2022).

Scott (1998) simply listed, in the synonymy of tharos species-group: “=distincta Bauer,

1975, type LACM (=Natural History Museum Los Angeles County) (Ferris, ed., 1989).”

Emmel, et al. (1998) listed distinctus at subspecific rank in their California checklist.

Austin (1998) listed distinctus at subspecific rank in the Nevada checklist.

Bailowitz & Brodkin (2007) gave the following habitat description for P. tharos: “This is

a species of riparian areas, predominantly in the southern reaches of the state. It frequents

permanent watercourse edges, lakesides, and cienegas [permanently saturated, alkaline,

freshwater, spongy, wet meadows], especially those with asters, beggarticks, and buttonbush.”

Pelham (2008-2023) listed distincta as a junior synonym of P. t. tharos and comments:

“Location of holotype not known”’.

METHODOLOGY

Color analysis was performed using the Color Grab™ cellphone application

(www.loomatix.com), version 3.9.2, to establish exacting RGB and HSB color codes under

“daylight” fluorescent lighting, in combination with the Colblindor™ application (www.color-

blindness.com/color-name-hue/) to produce refined color swatches rather than giving generalized

color descriptions as is traditional with taxon descriptions. Two different areas of the wings were

measured for their red/green/blue (RGB) and hue/saturation/brightness (HSB) color codes.

Ground color was measured on the dorsal hindwings, which showed a more consistent, stable color

in each of the two series (dorsal forewing color showed slight variation between the postmedian

ground color vs. remainder of the dorsal wings, on different specimens). Darkness of the black

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pattern in each series was performed on the outer portion of the forewings either near the apex or

tornus. The ventral surfaces were not analyzed, due to considerable variation across both

subspecies. 300 males and 196 females of subspecies tharos from many areas of eastern North

America (N.C., VA., W.V., MD., N.Y. and R.I.) were analyzed, while 50 males and 31 females of

subspecies distincta (all from Santa Cruz Co., AZ.) were analyzed. All specimens were of summer

phenotypes [sufficient numbers of spring specimens of distincta were not available]. Color codes

of individual specimens were then averaged to produce results for each subspecies (Fig. 4). Color

names in the description of each of the four species references the color names given in the Color

Grab™ and Colblindor™ applications. Of particular interest are the differing results of the color

analysis vs. visual comparisons. While the human eye perceives a decidedly paler overall

appearance in distincta (Fig. 5), the color analysis tools show fairly similar colors in the two areas

examined (Fig. 4). The color analysis tools do, however, reveal a difference in ground color and

slight difference in “blackness” of the wing patterns, between series of females. Males of both

subspecies are fairly similar. The comparisons are made between specimens of “fresh” condition.

Additionally, wing measurements were made from the examined series. Measurements

were made of forewing length (Fig. 4). These were then averaged and a range and mode for each

was determined. Interestingly, specimens of distincta showed more consistent measurements,

while tharos showed considerably more variation in wing measurements.

PHENOTYPIC COMPARISON

Southern Arizona distincta differs from eastern nominotypical tharos as follows:

Size. Male distincta are slightly smaller than nominotypical tharos (Fig. 4). The length

of the male forewings of the examined distincta series ranges 13-16 mm, averaging 14.5 mm,

whereas eastern U.S. tharos males averaged larger, ranging 12-19 mm, averaging 15.6 mm. The

mode was, interestingly, similar at 15 mm for both subspecies. Female distincta are larger than

males, ranging 15-18 mm, averaging 16.5 mm, whereas eastern U.S. tharos females averaged

slightly larger, ranging 15-20 mm, averaging 17 mm. The mode was also similar at 17 mm for

both subspecies.

Dorsal ground color. The ground color of distincta males and females is generally a

concolorous brown-orange (“Peru” in males, “Golden Bell” in females) (Fig. 4), whereas the

ground color of tharos males and females is generally more orange than brown (“Ochre” in males,

“Fire Bush” in females). Some females show a very slight amount of lighter ground coloration in

the postmedian area of the forewings. Visually, the ground color of nominotypical tharos in the

eastern U.S. is a brighter orange than distincta, which has a paler look (Fig. 5). This is less

apparent in the males, but more apparent between females of both subspecies. [Interestingly, the

color analysis tools did not pick up the intensity of the dark patterns, rather just the base color.

Dorsal pattern and color of markings. The wing markings of distincta males and females

are dark (“Cocoa”) brown (Fig. 4). Males of nominotypical tharos are similarly dark (“Cocoa’’)

brown, whereas females are a blackish (“Livid”) brown. Visual comparison of the wings reveals

that the wing markings of distincta are distinctly paler, giving a more brownish look compared to

nominotypical tharos, on which the wing markings are black and sharp. In distincta, the

postmedian orange band in the males is generally more broken than in tharos, whereas tharos

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males have a more continuous band. In distincta the pattern of markings is more consistent, and

not as variable as in tharos. In nominotypical tharos, the postmedian line on the male hindwings

is variably developed and often broken in mid-section, washed out by the orange ground color.

Distincta males always have this postmedian line fully-developed. In nominotypical tharos, the

males have a wider marginal wing pattern, giving a darker appearance, whereas, in distincta the

outer wing pattern is narrower, giving specimens a lighter overall appearance. The contrast

between females of both subspecies is striking (Fig. 5). Nominotypical tharos females generally

have heavier infuscation on the basal portion of the forewings, giving specimens a considerably

darker appearance, whereas in distincta females, this infuscation is generally lacking and the

underlying pattern is more visible, giving them a lighter appearance. In general, distincta has a

pallid appearance, compared to nominotypical tharos.

Ventral color and pattern. The general appearance of the ventral hindwings of distincta

males is somewhat paler than in nominotypical tharos, though coloration is highly variable in both

(Fig. 6). Male tharos have a more extensive, brighter orange wash on the forewings, whereas in

distincta this feature is somewhat broken up by a subdued, weblike pattern. The black pattern on

the outer portion of the ventral forewings is darker and more developed in nominotypical tharos.

The females of both subspecies display similar, highly variable coloration and pattern. The only

appreciable difference between adults of nominotypical tharos and distincta is in the brown

marginal patch of the hindwings. In distincta this patch is of a paler browner shade, whereas in

nominotypical tharos it is generally darker brown. This is more evident in the males.

Dorsal HW} Dorsal HW Dorsal FW

ground color patch

RGB color code

HSB color code

33, 84,81

32, 71, 80

news 15.6 12-19 15.0

18, 47,8

core 14.5 13-16 15.0

21, 44,19

P. t. tharos 226, 157, 62 49,41, 39 17.0 15-20 17.0

females 34, 72, 88 12, 20, 19

P. t. distincta 201, 134, 54 56, 39, 31 16.5 15-18 17.0

females 32, 73, 78 19, 44,21

Fig. 4. Color analysis results (average colors) and wing measurements of P. t. tharos and P. t. distincta.

Wing shape. While variable, the outer margin of the forewings of nominotypical tharos

tend to be more rounded, whereas the outer margin of distincta is straighter, appearing slightly

concave in some specimens. This is more noticeable in series of males (Fig. 5).

Fig. 5. Comparison of P. tharos tharos and P. tharos distincta. Males in left panel, females in right panel. P. ¢.

tharos males (column 1), top to bottom: Pendleton Co., W.V.; Tucker Co., W.V.; Union Co., N.C.; Fauquier Co.,

VA.; Frederick Co., VA.; Frederick Co., VA. P. t. distincta males (column 2): all Santa Cruz Co, AZ. P. t. tharos

females (column 3), top to bottom: Randolph Co., W.V.; Fauquier Co., VA.; Fairfax Co., VA.; Randolph Co., W.V.;

Randolph Co., W.V.; Fauquier Co., VA. P. t. distincta females (column 4): all Santa Cruz Co, AZ.

Fig. 6. Comparison of P. tharos distincta (left side) and P.

tharos tharos (right side) venters. Males in upper row, females

in lower row. Male and female P. ¢. distincta from Santa Cruz

Co., AZ. Male P. t. tharos from Haywood Co., N.C.; female

from Fairfax Co., VA.

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DISTRIBUTION

The range of distincta has yet to be refined and requires further study. Based on literature

sources, specimen series and images posted to butterfliesofamerica.com (accessed 16 Jan. 2023)

and iNaturalist.org (accessed 16 Jan. 2023), the range in the United States can be defined primarily

as extreme southeastern California (Imperial, Riverside and San Bernardino Counties) and south-

central Arizona (Pima, Santa Cruz and Yuma Counties). Most records are clustered south of Tucson

to the Mexican border. In Arizona, individuals in Cochise and Graham Counties show

intergradation to nominotypical tharos. All New Mexico records appear to be nominotypical tharos

with some individuals appearing slightly intermediate to distincta. All of Texas appears to be

within the range of nominotypical tharos. In Mexico, distincta is reported from the state of Baja

California (Baja California Norte). Images posted to butterfliesofamerica.com show candidate

distincta specimens down the west side of the northern half of Mexico, in Sonora and Nayarit states,

essentially west of the Sierra Madre Occidental. However, examined images in iNaturalist indicate

all other Mexican populations east of the Sierra Madre Occidental and in the southern half of

Mexico are closer to nominotypical tharos. Thus, distincta is confined to a very small range along

the California and Arizona border with Mexico, and possibly south along the Mexican Pacific Coast

region, and may be considered an endemic of the region.

COMMENT ON P. THAROS RIOCOLORADO

Scott (1992) described Phyciodes tharos riocolorado (initially as Phyciodes

tharos/morpheus riocolorado): “Adults...are characterized by having the same pattern of dark

lines as typical Phyciodes tharos/morpheus tharos but the black borders are narrower, and the

overall upperside color is more ochre-orange (less reddish-orange) than other tharos populations

(adults vary little in upperside color)...This subspecies is not enormously different from ssp.

tharos.” In agreeing with Scott’s latter point, a series of reared male specimens from Grand

County, UT. in my possession, as well as images in iNaturalist from Utah, are essentially

indistinguishable from reared nominotypical male tharos (Fig. 7), by shade of orange and by extent

of markings, and fall well within the range of variation of nominotypical tharos. Females in the

Utah series differ from nom. tharos mainly by having less extensive and less heavy black

infuscation on the basal half of the dorsal forewing (Fig. 7), as Scott (1994) noted, which

characterizes many tharos females. This gives riocolorado females a “paler” look, and fairly

similar to distincta. Thus, I am inclined to view riocolorado as a weak subspecies based on a

single female character (basal dorsal infuscation). A better set of character differences need to be

identified, especially for the males [outside the scope of this paper].

Scott (1994) asserted: “This ssp. has blackish antenna clubs, whereas the clubs are orange

on E. Colo. tharos, indicating that riocolorado derived from lower Colorado River (W Arizona/SE

Calif.) stock rather than western Great Plains stock which has orange antennae.” [Western Great

Plains orange-antenna populations are now deemed to be species-level Phyciodes orantain (Zhang,

et. al., 2022)]. Due to extreme similarity to nominotypical tharos males (Fig. 7), I contend that

riocolorado is likely derived from nominotypical tharos stock, with females appearing more like

distincta. Thus, riocolorado can be viewed as an intermediate population. Scott continued: “When

I named riocolorado I described the wings are oranger, but actually the orange seems to be about

the same tint as ssp. tharos but the wings appear paler because the black markings are much

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smaller, even on unf the black spots are smaller.” [See comments above, regarding dorsal

differences in females.] Scott goes on to describe other minor differences from nom. tharos,

mainly in the shape of the genitalia and differences in the larvae and pupae. I consider riocolorado

a weak subspecies but propose no changes to nomenclature here.

Fig. 7. Comparison of three subspecies of P. tharos. Subspecies distincta (left side), Santa Cruz Co., AZ.; subspecies

riocolorado (middle), ex-ova, Grand Co., UT.; subspecies tharos (right side), Pendleton Co., W.V. Males in top row,

females in bottom row.

ACKNOWLEDGEMENTS

Thanks go to Tom Kral for manuscript review; to Crispin Guppy for initial review and

discussions concerning type specimens; Tom Kral and Ken Davenport for distincta specimens

used in this study; Todd Stout for riocolorado specimens; Nick Grishin for information and photos

of distincta lectotype and paralectotype.

LITERATURE CITED

Austin, G. T. 1998. Checklist of Nevada Butterflies. Jn: T. C. Emmel (ed.). Systematics of

Western North American Butterflies. Mariposa Press, Gainesville, FL. Chapter 73: 837-

844.

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

Studies, Inc., Tucson, AZ.: 1x + 342 pp.

Bailowitz, R. A. & H. Brodkin. 2007. Finding Butterflies in Arizona, A Guide to the Best

Sites. Johnson Books, Boulder, CO.: xxi + 233 pp. + 8 pl.

Bauer, D. L. 1975. Tribe Melitaeini. /n: Howe (ed.) The Butterflies of North America.

Doubleday & Company, Garden City, NY: 139-195.

Brown, J. W., H. G. Real & D. K. Faulkner. 1992. Butterflies of Baja California. Lepidoptera

Research Foundation, Beverly Hills, CA.: iv + 129 pp. + 8 pl.

Emmel, T. C. & J. F. Emmel. 1973. The Butterflies of Southern California. Natural History

Museum of Los Angeles County. Science Series No. 26: xi + 148 pp.

Emmel, T. C., J. F. Emmel & S. Mattoon. 1998. A Checklist the Butterflies and Skippers of

California. Jn: T. C. Emmel (ed.). Systematics of Western North American Butterflies.

Mariposa Press, Gainesville, FL. Chapter 72: 825-836.

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.

Hodges, R. W. (ed.). 1983. Check List of the Lepidoptera of America North of Mexico. E.

W. Classey Limited and The Wedge Entomological Research Foundation. London,

England: xxiv + 284 pp.

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.

Pelham, J.P. 2008. A Catalogue of the Butterflies of the United States and Canada. Journal

of Research on the Lepidoptera, Vol. 40: xiv + 658 pp.

Scott, J. A. 1992. Hostplant records for butterflies and skippers (mostly from Colorado) 1959-

1992, with new life histories and notes on oviposition, immatures, and ecology. Papilio

(New Series) No. 6: 185 pp.

Scott, J. A. 1994. Biology and systematics of Phyciodes (Phyciodes). Papilio (New Series)

No. 7: 120 pp.

Scott, J. A. 1998. Phyciodes (Phyciodes): New discoveries, new subspecies, and

convergence. Papilio (New Series) No. 10: 42 pp.

Tilden, J. W. & A. C. Smith. 1986. A Field Guide to Western Butterflies. Peterson Field

Guide Series. Houghton Mifflin Company, Boston, MA.: 1x + 370 pp.

USDA. Accessed Jan. 15,2023. PLANTS Web Site. United States Dept. of Agriculture,

Natural Resources Conservation Service. https://plants.usda.gov/home

Zhang, J., Q. Cong, J. Shen, L. Song, R. J. Gott, P. Boyer, C. S. Guppy, S. Kohler, G. Lamas,

P. A. Opler & N. V. Grishin. 2022. Taxonomic discoveries enabled by genomic analysis

of butterflies. The Taxonomic Report 10(7): 1-60.

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