Volume 9 Number 4 3 June 2021

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

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

Review of Montana Euphilotes Mattoni, [1978],

with descriptions of new taxa

(Lepidoptera: Lycaenidae: Polyommatinae)

Steve Kohler

2617 Glen Drive, Missoula, MT 59804

and

Andrew D. Warren

McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History,

University of Florida, 3215 Hull Rd., UF Cultural Plaza, Gainesville, FL 32611-2710

ABSTRACT: The genus Fuphilotes at the northern end of its distribution 1s reviewed, with special emphasis on

populations in Montana. Also discussed are some Euphilotes populations in Oregon, Washington, Idaho and southwestern

Canada that have affinities with Montana. This region has until recently been essentially neglected in studies of the genus. We

characterize and discuss six species (two newly described) and twenty-five subspecies (six newly described) of Euphilotes,

including a number of yet undescribed populations. New species include Euphilotes heracleoides Kohler & A. Warren (type

locality in Sanders County, Montana) and Euphilotes oakleyi Kohler (type locality in Missoula County, Montana). New

subspecies include Euphilotes baueri borealis Kohler (type locality in Sanders County, Montana), Euphilotes baueri shoshone

Kohler (type locality in Butte County, Idaho), Euphilotes oakleyi madisonensis Kohler (type locality in Madison County,

Montana), Euphilotes ancilla montosa Kohler (type locality in Missoula County, Montana), Euphilotes ancilla campestris

Kohler (type locality in Powder River County, Montana), and Euphilotes rita montanensis Kohler (type locality in Carbon

County, Montana). Adults and male and female genitalia are figured for all newly described taxa, and compared to various

congeners.

Additional key words: Biogeography, distribution, genitalia, larval food plant associations, phenology.

ZooBank registration: urn:lsid:zoobank.org:pub:47038BBD-E62F-4390-A885-C491 B8E99ECE

INTRODUCTION

Euphilotes Mattoni, [1978] blues are among the least well-known of North American butterflies.

Since the original description of the genus, myriad subspecific taxa have been described, and the most

recent review of the genus (Pratt & Emmel, 1998) only partially resolved species-level taxonomic issues.

Much of the confusion surrounding Euphilotes stems from the very similar superficial appearances of

several taxa. Knowledge of larval food plants, adult genitalia, and local geographical and temporal

distributions are often necessary for identification. The life cycles of Euphilotes are closely coordinated

with those of their larval food plants, which are various Eriogonum Michx. species (Polygonaceae), wild

buckwheat. More than one taxon of Euphilotes may co-occur, either synchronously or not, but most co-

occurring species use different food plants (Austin et al., 2008).

Kohler (1980) listed only two species of Euphilotes as occurring in Montana, EF. battoides glaucon

(W. H. Edwards, 1871) and E. enoptes ancilla (W. Barnes & McDunnough, 1918). Pratt & Emmel’s

(1998) revision of the Euphilotes enoptes and E. battoides complexes elevated E. ancilla to the species-

level, but otherwise provided little information relative to Montana. This last and earlier works with a

1

primary focus on California and Nevada have essentially neglected the northern end of the distribution of

the genus, including Oregon, Washington, Idaho, Montana and southwestern Canada. Warren (2005)

presented considerable information on Euphilotes in Oregon, including some discussions of a few

populations in other northwestern states. Pelham (2008) presented a complete synonymic list of

Euphilotes, including all of the then known 57 described subspecies-level taxa (except E. bernardino

garthi Matton, 1989, endemic to Isla de Cedros, Baja California, Mexico), arranged into thirteen species.

At the time Pelham (2008) was published, several of the taxa described by Pratt & Emmel (1998) as

subspecies of FE. battoides (Behr, 1867) had not been formally aligned with species-level taxa in that

complex. As a result, the arrangement of subspecific taxa under EF. battoides in Pelham (2008) apparently

represented a polyphyletic assemblage. The listing of Euphilotes taxa by Warren et al. (2009) followed

Pelham’s (2008) arrangement, except that it also indicated a number of undescribed segregates among

then recognized species, included the recently described Euphilotes stanfordorum Opler & A. Warren,

2009, and included the Mexican endemic E. bernardino garthi. Pelham’s (2021) revised online catalog

lists 60 described subspecies-level Euphilotes taxa, arranged into 16 species. Color images of adults of

essentially all taxa discussed herein, including a large number of primary and secondary types, are

provided in Warren et al. (2016).

Males of most Euphilotes taxa may travel a considerable distance from their larval food plants in

search of mud, where they sometimes congregate in large numbers. In such situations, determining the

identities of these males in the absence of larval food plant information can be difficult. Rarely, females

also visit mud. Without prior detailed knowledge of what Euphilotes segregates in the different species

groups occur in any given area, Euphilotes adults found at mud sometimes cannot be reliably identified in

the field, even when voucher specimens are sampled (Warren 2005). We have used caution in applying

and citing information gleaned from the literature on Euphilotes taxa due to confusion between similar

species, the complex taxonomy of their Eviogonum food plants, and the large number of museum

specimens that are incorrectly determined to species-group (in these cases, male genitalia usually have not

been examined). Considerable variation exists between individuals in any given population of Euphilotes,

both in details of the configuration of the male and female genitalia, and wing characters in the adult

butterflies. In addition, some adults of the different described species can be very similar to one another.

To define species boundaries, recent authors have used a combination of food plant associations,

biological studies of the immature stages, geographic isolation, isolation due to seasonal differences in

adult flight periods, and adult morphology (various wing characters including ventral forewing suffusion,

maculation, the aurora of the hind wing, wing borders and fringes, the amount and shade of blue dorsally,

and male and female genitalia). Often, the food plant species for the taxon is more important in

identifying the taxon to species or subspecies than is the external morphology of the butterfly.

MATERIALS AND METHODS

Specimens examined are deposited in the collections of the McGuire Center for Lepidoptera and

Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville (MGCL); Oregon

State Arthropod Collection, Oregon State University, Corvallis (OSAC); research material of Andrew D.

Warren, Castle Rock, Colorado (ADW); private collections of Steve Kohler, Missoula, Montana (SK),

and Dana Ross, Corvallis, Oregon (DR). Specimens that form the basis for other records cited herein are

housed in American Museum of Natural History, New York, New York (AMNH); Essig Museum of

Entomology, University of California, Berkeley, California (EMEC); California Academy of Sciences,

San Francisco, California (MCAS); Museum of Comparative Zoology, Harvard University, Cambridge,

Massachusetts (MCZC); Natural History Museum of Los Angeles County, Los Angeles, California

(NHMLA); Peabody Museum of Natural History, Yale University, New Haven, Connecticut (PMNH);

National Museum of Natural History, Smithsonian Institution, Washington, D. C. (USNM); University of

Washington Burke Museum, University of Washington, Seattle, Washington (UWBM); William F. Barr

Entomological Museum, University of Idaho, Moscow, Idaho (WFBM); Washington State University,

Pullman, Washington (WSU).

2

Full data are provided for all Euphilotes specimens examined from Montana. In addition, full data

are provided from other states and provinces for all specimens examined of newly described taxa. For

most other taxa discussed herein, complete specimen data from MGCL, OSAC, and private collections

have not been listed. Partial listings of specimens examined, with an emphasis on those examined by the

senior author (SK), are provided for taxa occurring adjacent to those that exist in Montana or to

accompany specimens figured in captions; in most cases, data from specimens examined of California-

Nevada taxa described since 1998 have not been listed. For taxa discussed herein that also occur in

Oregon, data from specimens examined leading up to the publication of Warren (2005) are provided; data

obtained since 2005 generally are not.

Distribution maps were constructed using base topographical maps, to which were added the

captions, legends and dots, visually placed in the approximate collection localities. Each finished map

was then saved as a Word file. In addition to specimens we have personally examined, maps are

populated with locality data provided by Shields (1975, 1977), Stanford & Opler (1993), Austin (1998),

Layberry et al. (1998), Pratt & Emmel (1998, 2008), Davenport (2004a, b), Warren (2005) and Austin et

al. (2008), when unambiguous.

Figures of the adult butterflies are shown at 1.6X life size and to scale. Both dorsal and ventral

surfaces are shown for each specimen. Full data for each specimen is provided. Figures of the male and

female genitalia are shown at approximately 50X magnification. For comparison, the entire genital

capsule of each male is shown in the same position, with the left valva in lateral view, anterior left and

posterior right. For the female genitalia, the sclerotized ventral portion (lodix) of the terminal segments of

the female abdomen is shown in ventral view with the posterior end up. Previous authors of Euphilotes

studies have used a variety of terms to describe the external structures and configuration of the male and

female genitalia, at times using different terminology for the same structure. To avoid confusion when

describing and comparing the various structures for the purpose of this review, we list below in glossary

form the definitions upon which these comparisons will be made. The definitions are largely based on

those from Ehrlich & Ehrlich (1961), Klots (1970) and Tuxen (1970). The locations of these structures

are indicated on photos of the genitalia below.

--- inner process

- valva (bifurcate)

----.cucullus

SaccuS ---

4

Male genitalia of Euphilotes showing the shape of the left valva and the location of the associated structures. 1. Euphilotes

battoides complex. 2. Euphilotes oakleyi. 3. Euphilotes enoptes complex. 4. Euphilotes rita complex. All shown in left

lateral view.

MALE GENITALIA

Aedeagus: Penis or phallus, the male intromittent organ.

Anal angle: An angle of the cucullus, forming the ventrocaudal angle of the valva.

Crista obliqua (pl. cristae obliquae): Oblique, dentate process of the median, basal region of the valva.

Cucullus: Distal portion of the valva.

Falces (= gnathos): Paired, heavily sclerotized, curved or angulate arms, articulated with the caudal

margin of the tegumen ventrad of the base of the uncus and extending ventro-caudad.

Labides: Lobes of the uncus.

Pedunculus: Lateral part of the tegumen, articulating with the dorsal part of the vinculum.

Saccus: Anterior projection from the base of the vinculum.

Tegumen: The dorsal basal structure of the male genitalia, roof-like or hood-like.

Uncus: The dorsal distal structure extending caudad from the tegumen.

Valva (pl. valvae): Paired, presumably clasping organs of the male genitalia.

Vinculum: The U-shaped sclerite to which the valvae are attached.

lamella postvaginalis

- lamella postvagine

ostium bursae -- ostium bursae

m

&g lathella antevaginalis

amella antevarinag

'

- ductus bursae

5

lal ul

lamella pos " --- lamella postvaginalis “24 teil HAS

m bursae ee he le

ostium bursae ostium bursae --

antrum -antrum -

lamella antevaginalis ----

i ole

ductus bursae ---

3 Ba th ’ 1

Female genitalia of Euphilotes showing the shape of the lodix and the location of the associated structures. 1. Euphilotes

battoides complex, ventral view. 2. Euphilotes oakleyi, ventral view. 3. Euphilotes enoptes complex, ventral view. 4. Same,

lateral view. 5. Euphilotes rita complex, ventral view. 6. Same, lateral view.

FEMALE GENITALIA

Antrum: A bulbous, ribbed mating tube, the most caudal part of the ductus bursae when more heavily

sclerotized and differentiated from the remainder of the ductus bursae.

Bursa copulatrix: A collective term for the structure comprised of the ostium bursae, antrum, ductus

bursae and corpus bursae.

Corpus bursae: The enlarged sac-like distal part of the bursa copulatrix.

Ductus bursae: A tube of varying length, most commonly relatively narrow, and connecting the ostium

bursae with the corpus bursae, including the antrum. Part of the bursa copulatrix.

Lamella antevaginalis: The part of the lodix which is cephalad and ventral of the ostium bursae.

Lamella postvaginalis: the part of the lodix which is caudad and dorsal of the ostium bursae.

Lodix: The entire complex of sclerotized structures surrounding the ostium bursae, consisting of the

lamella antevaginalis and lamella postvaginalis, located mid-ventrally and caudad of the seventh sternite,

which fit corresponding structures of the male genitalia and presumably serve to hold the mating

individuals together. It is also called the genital plate or sterigma.

Ostium bursae: The copulatory entrance into the ductus bursae.

Note: No sharp distinction between the lamella antevaginalis and lamella postvaginalis is possible in Euphilotes, with the two

being joined and surrounding the ostium bursae and antrum. Mattoni [1978], in erecting the genus stated “Lodix large

subcuboid formed as a single apparent sclerite. Deeply invaginated into the seventh segment”.

RESULTS AND CONCLUSIONS

Below, we review described and known undescribed populations of Euphilotes, with a special

emphasis on those occurring in Montana and other northwestern states and Canadian provinces, at the

northern distributional limit of the genus. We document the distributions of all Euphilotes populations

known to occur in Montana, based on nearly five decades of extensive field work researching the butterfly

fauna of Montana, by the senior author. We describe two new species and six new subspecies of

Euphilotes, and figure adults and male and female genitalia of newly named taxa, as well as of other taxa

as relevant in comparisons with new taxa. Full synonymies for all taxa discussed are provided by Pelham

(2021).

Pratt & Emmel (1998) recognized three species complexes in Euphilotes and outlined characters

of the male and female genitalia used to separate them. As a result of the current study, we have

confirmed that the conformation of the valvae in the male genitalia and the lodix in the female genitalia

have highly significant taxonomic value for specific, and in some cases, subspecific determination. In this

study, we have refined the use of these differences in defining populations discussed below. The taxa we

discuss are organized according to the three species complexes as defined by Pratt & Emmel (1998).

THE EUPHILOTES BATTOIDES COMPLEX

Pelham (2021) included the following species in the Euphilotes battoides complex: E. battoides

(Behr) (Figs. 235, 316), E. bernardino (W. Barnes & McDunnough) (Figs 236-237, 317-318), E. allyni

(Shields), E. glaucon (W. H. Edwards) (Figs. 238-249, 319-323), E. baueri (Shields) (Figs. 261-271, 332-

340), FE. centralis (W. Barnes & McDunnough) (Fig. 272) and E. ellisii (Shields). Warren (2005)

discussed two additional undescribed taxa in the E. battoides complex, treated at the species-level, in the

Pacific Northwest. One of these, which is associated with Eriogonum heracleoides Nutt., is described

below. The second taxon, endemic to high elevations in the Cascade Range and associated with

Eriogonum marifolium Torr. & A. Gray, will be described by the authors in a subsequent publication. In

this species complex, the male genitalia have bifurcate valvae, consisting of a broad lower portion and a

narrow, usually curved upper prong of approximately equal length. In the female genitalia, the lodix is

considerably reduced, being more ring-shaped.

Euphilotes glaucon glaucon (W. H. Edwards, 1871)

(Figs. 1-22, 238-245, 319-321)

Lycaena Glaucon W. H. Edwards, 1871, Trans. Am. Entomol. Soc. 3(3/4): 205, 210, no. 9.

Philotes glaucon glaucon Edw.; McDunnough, 1938, Mem. S. Calif. Acad. Sci. 1: 28.

Philotes battoides glaucon (Edwards); dos Passos, 1964, Mem. Lepid. Soc. (1): 67; Langston, 1969, J.

Lepid. Soc. 23(1): 50.

Shijimaeoides battoides glaucon (Edwards, 1871); Shields, 1977, J. Res. Lepid. 16(1):31.

Euphilotes battoides glaucon (W. H. Edwards); Miller & Brown, 1981, Mem. Lepid. Soc. (2): 118.

Euphilotes battoides glaucon (W. H. Edws.); Mattoni, 1989, J. Res. Lepid. 27(3/4): 175.

Euphilotes battoides glaucon (Edwards, 1871); Pratt & Emmel, 1998, Syst. W. N. Am. Butterflies

(16): 216.

Euphilotes glaucon glaucon (W. H. Edwards, 1871); Warren, 2005, Butts. Ore.: 173; Pelham, 2008, J.

Res. Lepid. 40: 247; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Nevada: Storey County; near Virginia City.

Mattoni (1989) was the first to suggest that E. glaucon may represent a species-level taxon.

Warren (2005) elaborated on this hypothesis and applied the name broadly, as a wide-ranging polytypic

species, to all populations in Oregon (then mostly known from the western half of the state) feeding on

Eriogonum umbellatum Torr. This application was supported by adult morphology and the timing of

6

adult flight in populations occurring in sympatry and parapatry with other members of the EF. battoides

complex. Additional fieldwork in the Pacific Northwest since Warren (2005) has documented Er.

umbellatum-feeding populations of E. glaucon in northeastern Oregon, on the east side of the Cascades in

Washington, and at several sites in Montana. Eriogonum umbellatum-feeding populations of E.

“battoides glaucon”’ in western Montana were discussed by Shields (1977), but he listed only one

record—Ravalli County. Kohler (1980) listed records of E. “battoides glaucon” for only three Montana

counties (Madison, Missoula [in error], and Ravalli). As discussed above, E. glaucon is now considered a

species-level taxon. Since Kohler (1980), we have documented additional Montana localities for E.

glaucon, as detailed below. The morphology of Montana populations of £. glaucon does not differ

significantly from that of the nominotypical populations of the Great Basin.

Because of its use of (early blooming) Er. umbellatum, Davenport (2004a) treated Euphilotes

comstocki (Shields, 1975) as a subspecies of Euphilotes intermedia (W. Barnes & McDunnough, 1917)

(although Pelham (2021) includes intermedia as a subspecies of E. glaucon). This circumscription of E.

glaucon also implies that some additional taxa (outside the range of taxa treated by Davenport), described

as subspecies of FE. battoides by Pratt & Emmel (1998) and so treated by Pelham (2008), should be

associated with E. glaucon.

The known distribution of E. g. glaucon includes northern California, northern Nevada, western

Oregon, southern Idaho, southwestern Montana and northern Utah (Fig. 383).

Specimens Examined: USA: IDAHO: Lemhi County: W. side Bitterroot Range, upper Cow Creek Rd. fork, off Lemhi Pass

Rd., E. of Tendoy, 6205’, 18 July 2006, T.W. Ortenburger, 13 (at mud) (ADW); MONTANA: Beaverhead County: near

Lemhi Pass, 20 July 1982, S. Kohler, 43 (SK); Gold Cr., 6000’, FR 73, 15 mi. S. Wise River, 30 June 2017, S. Kohler, 1a 12

(SK); Ravalli County: Railroad Creek, 4950’, off Skalkaho-Rye Road, 25 June 1981, S. Kohler, 13% (SK); same locality, 25

June 2001, S. Kohler, 123 32 (SK); Silver Bow County: Barrel Spring trail, 6100’, 4-5 mi. NE of Melrose, 7 June 2004, S.

Kohler, 83 12 (SK); same locality, 1 June 2006, S. Kohler, 53 (SK); 5 June 2006, S. Kohler, 13a (SK); 6 June 2006, S.

Kohler, 123 (SK); 30 May 2007, S. Kohler, 14 (SK); Barrel Spring trail, 5680’, off Camp Creek Rd., 5 mi. NE Melrose, 15

June 2015, S. Kohler, 1a (SK); Camp Creek Road, 4-5 mi. E. of Melrose, 4 June 2004, S. Kohler, 13 (SK); same locality, 14

June 2004, S. Kohler, 33 (SK); Soap Gulch, 6980’, 7 mi. up, NE Melrose, 15 June 2015, S. Kohler, 43 (SK); Soap Gulch,

6850-7150’, 7-8 mi. up, NE Melrose, 17 June 2015, S. Kohler, 243 (SK); Soap Gulch, 6900’, 6 mi. up, NE Melrose, 21 June

2016, S. Kohler, 43 (SK); Same locality, 23 June 2016, S. Kohler 23 22 (SK); Soap Gulch, 6875-7300’, 6-7 mi. up, NE

Melrose, 24 June 2015, S. Kohler, 143 (SK); Same locality, 25 June 2015, S. Kohler, 43, (SK); Soap Gulch, 7300’, 7 mi. up,

NE Melrose, 3 July 2020, S. Kohler, 23, (SK); USA: NEVADA: Douglas County: US 395, 2.1 mi. S. Carson City line, 30

May 1980, G. T Austin, 43 42 (SK); Clear Creek Canyon, 2 mi. NW US 395, 29 May 1980, G. T. Austin, 23 22 (SK);

Washoe County: 3 mi. E. US 395 on Hwy. to Virginia City, 19 June 1980, J. B. Vernon, 42 (SK); OREGON: Deschutes

County: Hwy. 20, 1.5 mi. E. Millican, 8 June 2003, assoc. Er. umbellatum, A.D. Warren, 1¢ 12 (ADW); Rd.16, 1-2 mi. S.

Sisters, 7 June 2003, assoc. Er. umbellatum, A.D. Warren, 233 112 (ADW) (some males in the population have widened dark

wing margins, approaching the phenotype seen in EF. g. oregonensis); Sisters, 8 July 1953, S. Jewett, 23 32 (OSAC); same

locality, 24 June 1954, J. C. Downey, 12? (MGCL); Grant/Baker counties: Elkhorn Mts., saddle at 8000’, 24 July 2005, D.

Trochlell, 23 32 (ADW); Harney County: Hwy. 20, mi.85, just W. of Lake Co. line, 10 June 2003, assoc. Er. umbellatum,

A.D. Warren, 33 12 (ADW); Steens Cirque, 26 July 2000, D. Ross, 1¢ 12 (DR); Steens Mtn., Wildhorse Lake, 8450’, 21

June 1992, D. Ross, 13 12 (DR); Steens Mt., 25 July 1936, S. Jewett, 1a (OSAC); same locality, 19 July 1957, R. Albright,

13 12 (OSAC); 18 July 1958, 16 22 (OSAC); Steens Mt., E. rim overlook, 11 July 2000, D. Ross, 12 (DR); Steens Mtn.,

trailhead to summit, 25 July 1997, D. Ross, 23 12 (DR); summit of Steens Mt., 9600’, 28 July 1958, D Jewett, 1 2 (OSAC);

same locality, 17 July 1968, S. Dornfeld, 1 2 (MGCL); 1a 22 (OSAC); 15 August 1971, S. Dornfeld, 1 ¢ (MGCL); 2 August

1975, Hinchliff, 1a 22 (OSAC); Jackson County: Burnt Ck. Ranch Rd., 2 mi. S. Dead Indian Rd., 5400’, 1 July 1991, 12

(OSAC); Hyatt Lake, 9 June 1940, 16 (SK); Hyatt Prairie Rd., 1.2 mi. N. Hwy. 66, ca. 4750’, on Er. umbellatum, 2 June 2000,

A.D. Warren, 13 12 (ADW); near summit, Mt. Ashland, 6600’, assoc. Er. umbellatum, 23 June 2000, A. D. Warren, 3d

(ADW); same locality and plant association, 19 July 1997, D. Ross, 12 (DR); many more from this locality (OSAC); N. Fork

Scotch Ck. Cyn. and Lone Pine Ridge, 3800-4600’, on Er. umbellatum, 27 June 2000, A.D.Warren, 13 12 (ADW); meadow

above Soda Mtn. Rd., 3.8 mi. S. of jct. with Hwy. 66, 4900’, 21 June 2000, R.L. Romeyn 23 2? (SK); Soda Mtn. Rd. to

Hobart Peak (ca. 3 mi. S. Hwy. 66), 5200-5450’, on Er. umbellatum, 22 June 2000, A.D. Warren, 33 42 (ADW); Jefferson

County: along Metolius River at lower bridge, ca. 8 mi. N. Metolius Spring, 2300’, 31 May 2002, at mud but assoc. Er.

umbellatum, 13 (ADW); Canyon Creek Meadows, 30 July 1978, [V. McHenry], 3 3 (OSAC); Metolius River, 10 July 1952, S.

Jewett, 13 (OSAC); Rd. 11 at jct. Meadow Creek, E. side of Green Ridge, ca. 3200’, 2 June 2003, assoc. Er. umbellatum, A.D.

Warren, 43 22 (ADW); Klamath County: Bly Mtn., 5000’, 16 June 1963, E. Dornfeld, 33 (OSAC); same locality, 17 June

1963, E. Dornfeld, 13 12 (OSAC); 22 June 1959, E. Dornfeld, 23 (OSAC); 21 June 1958, R. Woodly, 3 ¢ (OSAC); Keno, 30

i.

May 1939, S. G. Jewett, 1a (MGCL); 83 82 (OSAC); Hwy. 66, at W. end jct. Hwy. 97, SW edge Klamath Falls, 4000’, 18

June 2003, assoc. Er. umbellatum, A.D. Warren, 33 62 (ADW); Lake County: Chandler State Park, 12 mi. N. Lakeview, 16

June 1958, E. Dornfeld, 1 ¢ (OSAC); Hwy. 20, mi. 81, ENE Glass Buttes, ca. 4500’, 9 June 2003, assoc. Er. umbellatum, A.D.

Warren, 33 12 (ADW); same locality, 10 June 2003, assoc. Er. umbellatum, A.D. Warren, 33 42 (ADW); Lost Forest nr.

Christmas Valley, 16 June 1970, 13 (OSAC); Warner Mts., Kelly Creek, T40S R21E S9, 4810’, 13 June 1999, V. Covlin, 16

(ADW).

Additional Records: The following include published records or others known to us. Specimens have not been examined.

USA: IDAHO: Lemhi County: Fourth of July Cr. at Blacktail Cr., 17 June 1999, L. May (WFBM); Salmon River Rd., 41 mi.

SW North Fork, 26 June 1999, L. May (WFBM); MONTANA: Madison County: C. Durden (reported to R. E. Stanford,

1978; these could be the new taxon described from Madison County, below); Ravalli County: Sula, 4700’, 23 June 1929, 12

(WFBM) (Shields, 1977).

Euphilotes glaucon oregonensis (W. Barnes & McDunnough, 1917)

(Figs. 23-26, 246)

Philotes battoides oregonensis B. & McD., 1917, Contrib. Nat. Hist. Lepid. N. Am. 3(4): 214;

McDunnough, 1938, Mem. S. Calif. Acad. Sci. 1: 28.

Philotes battoides oregonensis Barnes & McDunnough, 1917; dos Passos, 1964, Mem. Lepid. Soc.

(1): 67; Langston, 1969, J. Lepid. Soc. 23(1): 50.

Shijimaeoides battoides oregonensis (Barnes & McDunnough, 1917); Shields, 1977, J. Res. Lepid.

16(1):44.

Euphilotes battoides oregonensis (Barnes and McDunnough); Miller & Brown, 1981, Mem. Lepid.

Soc. (2): 118.

Euphilotes battoides oregonensis (B. & McD.); Mattoni, 1989, J. Res. Lepid. 27(3/4): 175.

Euphilotes battoides oregonensis (Barnes & McDunnough, 1917); Pratt & Emmel, 1998, Syst. W.N.

Am. Butts. (16): 220.

Euphilotes glaucon oregonensis (Barnes & McDunnough, 1917); Warren, 2005, Butts. Ore.: 173;

Pelham, 2008, J. Res. Lepid. 40: 247; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-

Cat.htm.

Type Locality: Oregon: [Klamath County]; Crater Lake.

Typical E. g. oregonensis occurs in the pumice flats and low hills of central Klamath County,

Oregon (e.g., Sand Creek area, ca. 4800’; south entrance to Crater Lake National Park), north to extreme

southwestern Deschutes County (Fig. 383). As with other E. glaucon populations, it is associated with

Er. umbellatum. Adults of E. g. oregonensis fly mostly from mid-June to mid-July. Males are dark blue

above, usually with very wide dark wing borders (Fig. 23). Above, females are dark brown, with a

narrow orange aurora (Fig. 25). Below, both sexes have a ground color similar to E. g. glaucon, and the

spot size averages somewhat larger (Warren, 2005) (Figs. 24, 26).

Specimens Examined: USA: OREGON: Klamath County: Antelope Desert, W. of Hwy 97 at mi. 226.6, 22 June 2017, B.

A. O’Hara, 13 2? (SK); Antelope Desert, Sand Creek area, flats W. Hwy. 97 at mi. 226, 4557’, 29 June 2017, R. L. Romeyn,

52 (SK); Crater Lake, 11 July 1925, 16 32 (OSAC); same locality, 12 July 1937, 43 (SK); 27 July 1933, 1d (SK); 11 July

1930, 22 (MGCL); 23 July 1944, 24 (OSAC); 26 July 1963, Hinchliff, 2¢ 2% (OSAC); 18 July 1970, D. W. Jenkins, 13 2°

(MGCL); 22 July [??], 1a (MGCL); Crater Lake N. P., 17 July 1944, B. Weber, 23 102 (MGCL); same locality, 23 July

1944, B. Weber, 53 42 (MGCL); Hwy. 97, vic. mi. 226.4, ca. 4700’, on Er. umbellatum, 21, 28 June 2000, A. D. Warren,

483 252 (ADW); same locality and plant association, 23 June 2001, A. D. Warren, 23 22 (ADW); same locality, association

and date, D. Ross, 63 5? (DR); 18 June 1992, D. Ross, 53 32 (DR); 17 June 2000, D. Ross, 73 2? (DR); Sand Creek, Hwy.

232, 3 July 1968, E. J. Dornfeld, 1¢ (MGCL); 0.9 mi. N. Sand Creek, and W. of US 97, 4500’, 18 June 2006, J. & F. Preston,

13 (MGCL); 1.6 mi. E. Hwy. 97, 9 mi. N. Kirk, 4650’, 27 July 1964, E. M. & S. F. Perkins, 23 (MGCL); 5 mi. N. Sun Pass,

Hwy. 232, 4750’, 10 July 1962, 12 (MGCL).

Euphilotes glaucon australoglaucon Pratt & J. Emmel, 1998

(Figs. 27-30)

Euphilotes battoides austroglaucon Pratt & J. Emmel, 1998, Syst. W. N. Am. Butts. (16): 216-217.

Euphilotes glaucon australoglaucon Pratt & J. Emmel, 1998; Pelham, 2008, J. Res. Lepid. 40: 247;

Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: California: Inyo County; east slope Sierra Nevada, Independence Creek, 6300’ elevation.

Pratt & Emmel (1998) considered this taxon to be “ecologically very close to subspecies glaucon’’,

but treated both as subspecies of E. battoides. Davenport (2004b), who stated, “The occurrence of

battoides on Glass Mountain well east of the Sierra Nevada within the range of glaucon suggests that

these may be two different species”, aligned astraloglaucon with E. glaucon, an arrangement followed by

Pellham (2008). This taxon differs from nominate glaucon mainly in the presence of extensive bluish

scaling above on many (but not all) females, and a somewhat paler blue color above in males. As

reported in its original description, E. g. astraloglaucon uses Eriogonum umbellatum Torr. var. nevadense

Gand. as its larval food plant. This taxon was reported (Pratt & Emmel, 1998) to occur along the east

slope of the Sierra Nevada in California, from about the latitude of Lone Pine in Inyo County, north to the

Sherwin Grade of the south part of Mono County (Fig. 383).

Specimens Examined: USA: CALIFORNIA: Inyo County: S. Fork Bishop Creek Canyon, S. of South Lake Rd., 0.2 mi. W.

of Mountain Glen CG, 8600’, 20 June 2017, assoc. Er. umbellatum, R. L. Romeyn, 1c (SK); flats S. FR 7S01, 4.5 mi. W.

Hwy. 168 via Buttermilk Rd., 6840’, Sierra Nevada Mts., 23 May 2017, R. L. Romeyn, 113 92 (SK); Mono County: S.

facing slope along upper Rock Creek Rd., 8500’, 2.5-3.4 mi. W. of jct. with Hwy. 395, 1 July 2011, assoc. Er. umbellatum, R.

L. Romeyn, 1d 12 (SK); NW ject. of Hwy. 395 and McGee Creek Rd., 7050’, Sierra Nevada Mts., 7 June 2016, assoc. Er.

umbellatum var. nevadense, R. L. Romeyn, 133 162 (SK).

Euphilotes glaucon comstocki (Shields, 1975)

Shijimiaeoides battoides comstocki Shields 1975, Bull. Allyn Mus. (28): 12.

Euphilotes battoides comstocki (Shields); Miller & Brown, 1981, Mem. Lepid. Soc. (2): 118; Matton1,

1989, J. Res. Lepid. 27(3/4): 175.

Euphilotes battoides comstocki (Shields, 1975); Pratt & J. Emmel, 1998, Syst. W. N. Am. Butts. (16):

220.

Euphilotes intermedia comstocki [Shields]; Davenport, 2004a, Tax. Rept. Int. Lepid. Surv. 4(7): 10.

Euphilotes glaucon comstocki (Shields, 1975); Pelham, 2008, J. Res. Lepid. 40: 248; Pelham, 2021,

http://www. butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: California: Kern County; Tehachapi.

In the original description, Shields (1975) stated that comstocki was known only from the type

locality (Tehachapi, Kern Co., California), and that the larval host was an unknown Eviogonum species.

Pratt & Emmel (1998) noted that comstocki is phenotypically much like E. g. glaucon, though it has

slightly narrower orange aurorae and smaller macules, and has a slightly later emergence, feeding on a

later-blooming Er. umbellatum. Davenport (2004a) stated that Pratt (pers. comm.) had concluded that

comstocki was better placed with intermedia than battoides. He also noted recent records expanding the

known distribution of comstocki in California as follows: Piute Mtn. Vista, Piute Mts., Kern Co. (Jim

Brock); Kennedy Meadows, Kern Plateau, Tulare Co. (James Scott); Bald Mountain, Kern Plateau, Tulare

Co., Pine Flat at the south end of the Kern Plateau, the Piute Mtn. Rd. overlooking Lake Isabella and

several sites in the Greenhorn Mts., all Kern Co. (Ken Davenport). Davenport (2004a) also noted that E.

g. comstocki is common in its localized habitats where its larval food plant is Er. umbellatum. The

distribution is shown in Fig. 383.

Euphilotes glaucon intermedia (W. Barnes & McDunnough, 1917)

(Figs. 31-34, 249)

Philotes battoides intermedia B. & McD., 1917, Contrib. Nat. Hist. Lepid. N. Am. 3(4): 214.

Philotes glaucon intermedia B. & McD.; McDunnough, 1938, Mem. S. Calif. Acad. Sci. 1: 28.

Philotes battoides intermedia Barnes & McDunnough, 1917; dos Passos, 1964, Mem. Lepid. Soc. (1):

67; Langston, 1969, J. Lepid. Soc. 23(1): 50.

Shijimaeoides battoides intermedia (Barnes & McDunnough, 1917); Shields, 1977, J. Res. Lepid.

16(1): 44.

Euphilotes battoides intermedia (Barnes and McDunnough); Miller & Brown, 1981, Mem. Lepid. Soc.

(2): 118.

Euphilotes battoides intermedia (B. & McD.); Mattoni, 1989, J. Res. Lepid. 27(3/4): 175.

Euphilotes battoides intermedia (Barnes & McDunnough, 1917); Pratt & Emmel, 1998, Syst. W. N.

Am. Butts. (16): 220.

Euphilotes intermedia (Barnes & Benjamin) ; Davenport, 2004a, Tax. Rept. Int. Lepid. Surv. 4(7): 10.

Euphilotes glaucon intermedia (Barnes & McDunnough, 1917); Warren, 2005, Butts. Ore.: 173;

Pelham, 2008, J. Res. Lepid. 40: 247; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-

Cat.htm.

Type Locality: [California]: Shasta County; (presumably in the vicinity of Castella in the Upper

Sacramento Valley).

Mattoni (1989) noted that intermedia is sympatric and synchronic with E. b. battoides at Gold

Lake, California, and Warren (2005) allied intermedia with E. glaucon, since they become

indistinguishable in southwestern Oregon. This arrangement was followed by Pelham (2008). The

distribution of E. g. intermedia includes northern and east-central California (Fig. 383). Warren (2005)

discussed Oregon populations of E. glaucon from Josephine and Curry counties which he called

Euphilotes glaucon nr. intermedia. Adults from these two counties differ from typical E. g. intermedia by

having somewhat better-developed dark wing margins and darker blue males above, and in having

slightly bolder spots below. The two taxa share the same pale ventral ground coloration. He also

discussed another unique population from along the western slope of the Cascades in Lane County,

apparently related to E. g. nr. intermedia. Adults from this population average larger than those from

Josephine and Curry counties and males are darker blue above, with somewhat wider dark wing margins.

Below, adults of the Lane County segregate have very small discal spots (some spots are occasionally

absent), and ground color is whitish, even paler than that of FE. g. nr. intermedia. However, ventral hind

wing aurorae are bold and well-developed on adults in Lane County, and contrast strongly with the

otherwise mostly immaculate ventral surface. These populations will be treated by the authors in greater

detail in a subsequent publication, but for now are mapped as E. g. intermedia (Fig. 383).

Populations of EF. glaucon in Jackson, southern Klamath, and southern Lake (Warner Mts.)

counties of southwestern Oregon are highly variable. Adults from these populations generally resemble

E. g. glaucon, though occasional males approach the appearance of EF. g. oregonensis with somewhat

widened dark wing margins. For now, all of these populations are mapped (Fig. 383) as E. g. glaucon.

Euphilotes glaucon (W. H. Edwards, 1871) (N. Oregon-Washington Cascades segregate)

(Figs. 35-38, 247-248, 322-323)

Warren et al., 2016, Illustrated lists of American butterflies, Revised 21 Nov. 2017, Accessed 19 Feb.,

2021, http://www.butterfliesofamerica.com.

Warren (2005) discussed a population of EF. g/aucon that occurs at the north end of the Cascades in

Oregon, at and above tree line on Mt. Hood (Clakamas and Hood River counties), and referred to it as

Euphilotes glaucon nr. glaucon. These populations occur in association with Eriogonum umbellatum var.

10

haussknechtii (Dammer) M.E. Jones, and adults are phenotypically most similar to E. g. glaucon.

However, adults from the Mt. Hood area differ from nominate F. glaucon in having paler blue males,

wing fringes not strongly checkered, and a somewhat darker ventral coloration (Figs. 35-38). Adults fly

well into August on Mt. Hood, and probably begin to fly shortly after the snow has melted in their

habitats. Similar populations of E. glaucon also occur in the vicinity of Mt. Adams in Yakima County,

Washington and perhaps further north on the eastern slope of the Cascades in Washington (Fig. 383).

Euphilotes glaucon may range considerably further north in the Washington Cascades than Mt. Adams, as

suggested by the additional records listed below. Whether or not some of these populations could

represent E. umbellatum-feeding populations of the new species described below, which normally utilizes

Er. heracleoides as a larval food plant, remains to be fully investigated. We will treat these populations in

greater detail in a subsequent publication.

Specimens Examined: USA: OREGON: Clackamas County: Timberline Lodge, [Mt. Hood], 21 July 1979, 12 (OSAC);

Clackamas/Hood River Counties: Mt. Hood, NW Timberline Lodge, 5700-6000’, 20 July 2003, on Er. umbellatum var.

haussknechtii, A. D. Warren, many males and females (ADW); Hood River County: Lookout Mtn., 6600’, 22 July 1978,

Hinchliff, 1¢ 22 (OSAC); S. slope of Mt. Hood, above Timberline Lodge, 6 August 2010, asssoc. Er. umbellatum, B.

O’Hara, 13 12 (SK); WASHINGTON: Yakima County: vic. Bird Meadows, Mt. Adams, 7 August 1953, S. Jewett, 13

OSAC); Bethel Ridge Microwave Tower Rd., 18 July 2009, assoc. Er. umbellatum, K. Kendall, 13 SK); same locality, 18

July 2009, assoc. Er. umbellatum, D. Nunnallee, 13 (SK); same locality, 18 July 2009, assoc. Er. umbellatum, R. M. Pyle, 1?

(SK).

Additional records: USA: WASHINGTON: Chelan County: Entiat Summit Road, 5550’ collected a large sample of

Eriogonum umbellatum var. hausknechtii for larvae, 6 August 2006, 19 larvae obtained 12 August 2006, adults pinned, 10

September 2010, D. & J. Nunnallee; Chumstick Mountain summit, 5810’, 24 July 2007, on Er. umbellatum, D. & J. Nunnallee,

12 adults including 3 2; Derby Canyon 3055-3075’, 11 July 2010, J. Pelham, D. & J. Nunnallee, A. Warren, J. Lane, on Er.

umbellatum hausknechtii; same locality, 27 July 2011, assoc. Er. umbellatum, 42; NF 7100, Derby Canyon, 27 July 2011,

assoc. Er. umbellatum; NF 7400, 4692’, 27 July 2011, assoc. Er. umbellatum, D. & J. Nunnallee, sample collected; Chumstick

Mountain summit, 5780-5829’, 6 August 2011, on Er. umbellatum hausknechtii, D. & J. Nunnallee; Kittitas County: Reecer

Creek Road, 4800’ 15 July 2010, nectaring on Er. umbellatum, D. & J. Nunnallee, 12; same locality, collected Ev.

umbellatum flowers, 5 August 2010, found 1 nearly mature larva, 7 August 2010, found 3 pupae, 20 September 2010, obtained

adults; same locality, 13 July 2011, on Er. umbellatum, D. & J. Nunnallee, 23; Stevens County: Cottonwood Divide Road,

Chewelah Mountain, 5070’, 22 July 2012, B. Yake, J. Barreca; Chewelah Mountain Summit, 5560-5773’, 22 July 2012, assoc.

Er. umbellatum, B. Yake, J. Barreca; same locality, 5760-5780’, 1 August 2012, assoc. Er. umbellatum, D. Nunnallee, mostly

?; Yakima County: Mt. Adams, south side via trail, 6000’, 11 June 2012, D. & J. Nunnallee; Bird Creek Meadows Trail,

5820’ 11 August 2012, assoc. Er. umbellatum dwarf mountain variety, D. & J. Nunnallee, 12; Bethel Ridge Microwave

Tower, 6080-6223’, 18 July 2009, strongly attracted to Er. umbellatum plants, D. & J. Nunnallee, 5 voucher adults collected.

Euphilotes heracleoides Kohler & A. Warren — new species

(Figs. 39-60, 250-260, 324-331)

ZooBank registration: urn:lsid:zoobank.org:act:F72E0453-D30A-4E39-AF2D-61DB7D595CEB

Warren (2005), who was the first to diagnose this taxon, treated it as an undescribed species, and

discussed its morphological and ecological attributes in detail. As early as 2000, Kohler found what

appeared to be two different species of Euphilotes in the battoides complex occurring at the same locality

in Sanders County, Montana. The area has an extensive stand of Eriogonum ovalifolium, Nutt. var.

pansum Reveal as well as a considerable amount of Er. heracleoides, Nutt. The first Euphilotes species

(E. baueri subspecies, see below) begins to fly when the Er. ovalifolium is in the full pre-bloom stage,

which typically occurs in early May. The Er. heracleoides in the same area blooms about one month later,

in early to mid-June, and the second battoides complex Euphilotes species, E. heracleoides 1s associated

with it.

Definition: As its name implies, Euphilotes heracleoides is found almost everywhere Eriogonum

heracleoides var. heracleoides Nutt. (Fig. 378, upper left) is abundant. While all Euphilotes are directly

associated with their larval food plants, few taxa are as reliable among random searches of unfamiliar

stands of Eriogonum as E. heracleoides. However, this species does appear to use Er. douglasii Benth. as

a larval food plant at some sites in Washington (as indicated below) and possibly in northern Oregon as

11

well (Warren, 2005). At sites where Er. heracleoides grows in sympatry with Er. umbellatum (e.g.,

summit of Pine Mountain, Deschutes County, Oregon), adults strongly orient towards and perch (e.g., for

the night) on flowers of Er. heracleoides, and show interest in Er. umbellatum only rarely, apparently to

seek nectar. At one site in Lincoln County, Montana, where Er. heracleoides grows in sympatry with Er.

flavum Nutt., adults of E. heracleoides occasionally perch or nectar on the Er. flavum. At another site in

Lincoln County, a long series of both males and females was collected on Er. flavum and no other

Eriogonum were present, strongly indicating that at this site, EF. heracleoides may be utilizing Er. flavum

as a larval food plant.

Adult E. heracleoides males are bright blue above, with fairly bold black wing margins. The

orange aurora of the dorsal hindwing is variable from none (Figs. 43, 53) to just a hint (Fig. 39), to more

extensive (Fig. 45). The majority of male individuals have little to no dorsal orange. Females are warm

brown dorsally, and usually have a somewhat broad orange aurora. Ventrally, both sexes of E.

heracleoides are quite similar to E. g. glaucon in having a bold forewing spot pattern and suffusion of

dark scales at the base of both forewings and hindwings, however, in EF. g. glaucon, the suffusion

continues out into the discal portion of both the forewing and hindwing, while in E. heracleoides the

discal area is free of suffusion, allowing the light gray ground color to be more visible, giveing the ventral

wing surfaces a brighter appearance. The genitalia are typical of the E. battoides complex, with males

having bifurcate valvae (Figs. 250-260), and females with the lodix ring-shaped (Figs. 324-331).

The life history of E. heracleoides was recorded by James & Nunnallee (2011), as “Cascadia

Blue”, Euphilotes on Eriogonum heracleoides (undescribed). They reared or partially reared E.

heracleoides on Er. heracleoides on several occasions, most frequently from partially grown larvae found

on buckwheat flowers, which pupated and eclosed as adults the following spring. They estimated the

development period from egg-hatch to pupation to be approximately 25 days. During development, the

larvae fed first on flowers and later on the developing seeds after the petals had senesced. As detailed by

James & Nunnallee (2011), the most closely related species, E. glaucon, is fairly distinct from £.

heracleoides in several immature stages. The egg of FE. glaucon has a much less conspicuous micropyle,

and the first instar larva is much lighter in color and has more extensive dorsal setae. In E. glaucon the

second instar larva is pale off-white and the third instar larva is generally pale, while the FE. heracleoides

second instar larva is red and white, and it is strongly colored and maculated in the third instar. The

fourth instar of E. heracleoides is variable, ranging from pale gray with diffused red markings to having

much brighter red markings, while this stage in E. glaucon is less brightly marked.

Etymology: This species is named after its primary food plant, Eriogonum heracleoides. While it may

use other species of Eriogonum food plants in some areas (see below), any Euphilotes found in close

association with Er. heracleoides in Oregon, Washington, British Columbia, Idaho and Montana, is very

likely to be Euphilotes heracleoides (although Euphilotes a. ancilla is associated with this plant in

Nevada (see below), and might be associated with it in far southern Idaho.

Distribution and Phenology: Euphilotes heracleoides is known from essentially all sites north of Nevada

where Eriogonum heracleoides is abundant (Fig. 384). This range includes most of northeastern Oregon,

including Baker, Crook, Deschutes, Gilliam, Grant, Harney, Jefferson, Malheur, Morrow, Sherman,

Umatilla, Union, Walllowa and Wasco counties. The report by Hinchliff (1994) of Euphilotes

“battoides”’ from Wheeler County, Oregon, almost certainly refers to E. heracleoides as well. The

species flies across most of eastern Washington; specimens were examined from Chelan, Columbia,

Klickitat, Kittitas and Okanogan counties. Records for E. “battoides” in Hinchliff (1996) suggest that

this taxon also occurs in Adams, Asotin, Benton, Douglas, Ferry, Franklin, Garfield, Grant, Lincoln, Pend

Orielle, Spokane, Stevens, Walla Walla, Whitman and Yakima counties. Washington records recently

received from Jonathan Pelham (pers. comm., 2021; see below) confirm all of these counties (Fig. 384).

As study of museum collections has shown, the great majority of records for E. “battoides glaucon” from

Washington (e.g., Pratt & Emmel, 2008) represent E. heracleoides, although the distribution of E.

glaucon there remains poorly known (see above). Euphilotes heracleoides appears to be common in

much of western Idaho, where Er. heracleoides is widespread, at least including Boise, Camas, Elmore,

Latah and Lewis counties, based on specimens we examined (Fig. 384). Few specimens from the £.

12

battoides complex are available from far southern and eastern Idaho, and almost no information on food

plant associations from that region is known. Thus, the southeastern distribution of E. heracleoides

remains to be determined, in conjunction with that of E. glaucon.

The distribution of E. heracleoides in Montana includes only the far northwestern portion of the

state, in Flathead, Lake, Lincoln and Sanders counties (Fig. 384). However, Er. heracleoides is found

farther south in the western end of Montana (Beaverhead, Gallatin and Madison counties) and also in

north central (Chouteau County) areas of the state (Lesica, 2012). Additional field work is needed to

determine if any populations of E. heracleoides are present in these additional areas where the food plant

occurs. Apparently, all records for the E. battoides complex from British Columbia, all from the extreme

south-central part of the province, represent E. heracleoides.

It remains somewhat unclear how far south the distribution of Euphilotes heracleoides extends.

The apparent occurrence of this species in the Warner Mountains of Lake County, Oregon (see additional

records below and Warren, 2005) suggests that it may also occur in parts of the same range in Modoc

County, California. Reports of both Euphilotes g. glaucon and E. g. oregonensis (as a subspecies of

intermedia) from the Warner Mountains of Modoc County, California, by Pratt & Emmel (1998, 2008)

may imply that their “oregonensis”’ refers to the same taxon we herein consider to be FE. heracleoides; we

do not consider the Warner Mountains to be part of the range of E. g. oregonensis (see above), although

acknowledge that populations of £. glaucon in this range appear to be highly variable, with some adults

approaching the dark-margined phenotype of E. g. oregonensis. Obviously, further research on

Euphilotes in the Warner Mountains is needed.

As with most Euphilotes, the adult flight time of the single brood of E. heracleoides usually

coincides with the period just before the peak blooming time of its Eriogonum food plant, but adults can

sometimes be found for some time after the peak blooming period, although generally in worn condition.

Records of adults in Montana extend from 1 June through 9 July at 2900-3600’ elevation, but Oregon

flights can be earlier; late April to mid-July at elevations from 1500-7500’ (Warren, 2005), as indicated

by material examined. The timing of adult flights of any population may be variable on an annual basis, as

a result of local weather conditions.

Types: Holotype male: MONTANA: Sanders County: W. of Niarada, 3 June 2004, S. Kohler Coll.

(47.81588, -114.62969). Allotype female: MONTANA: Sanders County: 1.5 mi. N. of Niarada, 13

June 1996, S. Kohler Coll. (47.82988, -114.61261). Paratypes: MONTANA: Sanders County: 2 mi. W.

of Niarada, 3300’, 14 June 1996, S. Kohler, 453 522; 1.5 mi. W. of Niarada, 9 June 1997, S. Kohler, 62

52; same locality, 1 June 2000, S. Kohler, 19d 122; same locality, 2 June 2000, S. Kohler, 133 142; 5

mi. SW of Niarada, 8 June 2000, S. Kohler, 23; SW of Niarada, 15 June 2001, S. Kohler, 100 42; SW of

Niarada, 19 June 2001, S Kohler, 193 492; West of Niarada, 3 June 2004, S. Kohler, 323 8 (all

collected in association with Eriogonum heracleoides var. heracleoides).

Deposition of Types: The holotype male, allotype female and four male and four female paratypes will

be deposited in the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural

History, University of Florida, Gainesville, Florida. The remaining paratypes are in the Kohler

Collection.

Type Locality: USA: MONTANA: Sanders County: vicinity of Niarada.

Additional Specimens Examined: (All collected in association with Eriogonum heracleoides var. heracleoides). USA:

IDAHO: Boise County: FR 380, from summit of Mores Pass, 7119’, 43 56.796’, 115 41.029’, 14 July 2005, T. W.

Ortenburger, 6 ¢ (ADW); MONTANA: Flathead County: 1.5 mi. N. of Niarada, 13 June 1996, S. Kohler, 233 82 (SK);

Lake County: 3 mi. E. Hot Springs, 3600’, 20 June 1996, S. Kohler, 93 62 (SK); Lincoln County: Dancing Prairie, NW

Eureka, 3350’, 1 July 1996, S Kohler, 63 62 (SK); Dancing Prairie, 5 mi. NW Eureka, 25 June 1997, S. Kohler, 3d 42 (SK);

Burma Road, NE Eureka, 9 July 1999, S Kohler, 12 (SK); same locality, 6 July 2005, S. Kohler, 1d¢ 42 (SK); Dancing

Prairie, NW Eureka, 28 June 2011, S. Kohler, 463 262 (SK); same locality, 6 July 2011, S. Kohler, 16 22 (SK); lower

Burma Road, state section, NE of Eureka, 6 July 2011, 1d 12 (SK); S. Burma Road, 0.8 mi. SE Hwy. 93, 2840’, 5 June 2018,

S. Kohler, 463 112 (SK); Sanders County: 13 mi. N. of Perma, 13 June 1996, S. Kohler, 83 32 (SK); 4.5 mi. N. Camas

prairie, 9 June 1998, S. Kohler, 383 222 (SK); 1 mi. west of Perma, 11 June 2013, S. Kohler, 293 282 (SK); 1 mi. west of

Perma, south of Hwy. 200, 2675’, 19 June 2017, S. Kohler, 1036 52 (SK); OREGON: Baker County: Blue Mts., Elkhorn

Range, Rock Ck. Cyn., 1-2 mi. SW Nat. For. boundary, 21 June 2002, A. D. Warren, 1¢ 12 (ADW); Grant County: Aldrich

13

Mts., Fields Creek Cyn., vic. jct. Rd. 21 and NFD 2140, 4000-4200’, 28 May 2001, A. D. Warren, 23 (ADW); same locality,

16 June 2001, A. D. Warren, 43a (ADW); Aldrich Mts., Fields Creek Rd., T14S R28E, 23 June 1995, D. Ross, 13 (DR);

Wickup Ck., 0-1 mi. S. jct. Canyon Ck., S. of C.G., ca. 4600’, 20 June 2002, A. D. Warren, 1 ¢ (ADW); Rd. 16, 2-3 mi. W.

Malheur Co. line, ca. 5000’, 20 June 2002, A. D. Warren, 12 (ADW): Harney County: Blue Mts., King Mtn. Lookout, ca.

6500’, 16 June 1996, V. Covlin, 23 22 (ADW); same locality, 25 June 1997, V. Covlin, 1a 22 (ADW); Malheur County:

Bendire Mtn. — Hunter Creek area, ca. 6 mi. N. Beulah Res. on Bendire Rd., 4100-4500’, 17 May 2002, A. D. Warren, 13

(ADW); Malheur/Baker counties: N. end Sheephead Mtn. to E. Camp Creek Rd., 2-4 mi. W. Hwy. 26, 4700-4900’, 20 June

2002, A. D. Warren, 103 22 (ADW); Morrow County: Blue Mts., Rd. 53, 4 mi. W. Umatilla Co. line, ca. 5200’, 26 June

2002, A. D. Warren, 1a (ADW); Umatilla County: Blue Mts., Hwy. 244 at Lake Ck. C. G., 4200’, 25 June 2002, A. D.

Warren, 15 (ADW); Blue Mts., Hwy. 244, ca. 2 mi. SE Lane Ck. C. G., ca. 4300’, 25 June 2002, A. D. Warren, 33 22

(ADW); Blue Mts., Rd. 020 at Frazier Ck. C. G. 4400’, 25 June 2002, A. D. Warren, 33 22 (ADW); Union County: Marley

Ck. Lane (NFD 2442), ca. 2 mi. SW Starkey on Hwy. 244, 25 June 2002, A. D. Warren, 1 ¢ (ADW); Phillips Cyn., ca. 2 mi.

W. Elgin, ca. 3000’, 23 June 2002, A. D. Warren, 12 (ADW); Wallowa Mts., North Fork Catherine Creek, NFD 7785, ca. 3

mi. E. Medical Springs Hwy. (203), 22 June 2002, A. D. Warren, 23 (ADW); Walllowa County: Blue Mts., Powwatka Ridge,

ca. 11 mi. N. of Wallowa on Troy Rd., 4100’, 23 June 2002, A. D. Warren, 83 112 (ADW); Minam, along Wallowa River,

2900’, 23 June 2002, A. D. Warren, 93 (ADW); Wallowa Mts., Gumboot Ck., vic. jct. Imnaha River on Rd. 39, 3900’, 24 June

2002, A. D. Warren, 23 (ADW); Wallowa Mts., Salt Creek Summit Rec. Site, NFD 3915, 6000’, 24 June 2002, A. D. Warren,

13 (ADW).

Additional Records: The following specimens are listed here because they occur in association with a variety of Eriogonum

heracleoides other than Er. heracleoides var. heracleoides, because there may be doubt about food plant associations, and/or

because they were considered to represent another taxon in publications on the genus by Shields (e.g., 1975). Many museum

specimens labeled without specific food plant association information fall into the latter two categories. CANADA: BRITISH

COLUMBIA: Oliver, 3 June 1933, J. McDunnough, 12 (MGCL); same locality, 15 June 1978, 13 (MGCL); Penticon, 4

May 1978, 23 (MGCL); same locality, May 1978, K. Thorne, 23 (MGCL); same locality, May 1979, K. Thorne, 12

(MGCL); same locality, June 1978, K. Thorne, 1 2 (MGCL); Princeton, 26 June [??], B. Weber, 330 3 2 (MGCL); Shingle Ck.

Rd., Keremos, 19 June 1933, A. N. Gartrell, 1 ¢ (MGCL); Summerland, May 1978, 1 2 (MGCL); same locality, May 1979, 22

12 (MGCL); same locality, 6 June 1986, 13 (MGCL); 3 km N. of Grand Forks, assoc. Er. heracleoides, 3 June 2003, N.

Kondla, 193 102 (ADW); USA: IDAHO: Boise County: Arrowrock Dam, 15 May [??], 1a (MGCL); Banks NF

Campground, off SR-55, nr. Payette River, 2800’, 26 June 1990, D. L. Eiler, 1a 12 (MGCL); Cottonwood Canyon, 30 May

[??], J. H. Manning, 23° (MGCL); Deep Park, 4200’, N. Fork Boise River, [no date], J. H. Manning, 1 ¢ (MGCL); Grimes Ck.

Cyn., 2 mi. upstream ID Hwy. 21, 12 June 1989, L. D. Beutler, 23 (ADW); nr. Boise, 2700’, [no date], J. H. Manning, 3¢

(MGCL); road by Arrow Rock Reservoir just above Arrow Rock Dam, 12 June 1988, D. L. Eiler, 1 2 (MGCL); Shafer Butte

Picnic Area, near Bogus Basin Winter Sports Area, ca. 7000’, 28 July 1981, D. L. Eiler, 23 39 (MGCL); Camas County:

Sawtooth Nat. For., Couch Summit, 7000’, 15 July 1964, J. Baker, 1 ¢ (MGCL); Elmore County: Arrowrock Dam, 4000’, 30

May [??], J. H. Manning, 1¢ (MGCL); Sawtooth Nat. For., nr. Featherville, 4900’, 13 July 1964, J. Baker, 1a (MGCL);

Latah County: [no locality], 1 June 1986, 12 (MGCL); Lewis County: Clearwater River, 10 km west of Kamiah, 500m, 20

May 1982, J. Reichel, 1a (MGCL); same locality, 28 May 1981, J. Reichel, 1a (MGCL); same locality, 1 June 1984, J.

Reichel, 1a (MGCL); same locality, 3 June 1984, J. Reichel, 23 (MGCL); 10 mi. W. Kamiah, 29 May 1981, J. Reichel, 53

(SK); MONTANA: Lincoln County: Dancing Prairie, NW Eureka, 28 June 2011, assoc. Er. flavum, S. Kohler, 73 52 (SK);

N. Sophie Lake, 2600’, NW Eureka, 6 June 2018, assoc. Er. flavum, S. Kohler, 243 442 (SK); OREGON: Baker County:

along rd. between Joseph & Hwy. 86, 13.2 rd. mi. N. jct. Hwy. 86, Duck Creek turnoff to Fish Lake, 14 July 1970, S.K.

Dvorak & O. Shields, 23 (MGCL); Anthony Lake, 10 July 1947, J. Baker, 1 ¢ (OSAC); 17 July 1970, J. Baker, 23 (OSAC); 7

July 1973, J. Baker, 53 (OSAC); Anthony Lakes, Whitman Nat. For., 25 July 1946, 136 (MGCL); Baker, 4 July 1941, M.

Plomley, 33 (MGCL); Big Lookout Mtn., 13 July 1974, J. H. Baker, 12 (VC); Blue Mts., Pine Creek, 6 July 1967, J. M.

Plomley, 23 (MGCL); Burnt Riv. Cyn., Lookout Mtn. Rd. vic. jct. I-84 exit 338, 2700’, 11 June 1999, taken while feeding at

flower of Eriogonum compositum, no other Eriogonum observed in area, A. D. Warren, 1 do (ADW); Durkee, 8 June 1940, L.

W. Morley, 13 (OSAC); same locality, 2640’, 11 June 1942, J. H. Baker, 15 (OSAC); Eagle Creek nr. Richland, 11 July

1960, 23 (OSAC); FR 11 to Bald Mtn., 19 June 2001, V. Covlin, 13 12 (VC); Halfway, N. Fork Pine Creek, 20 June 1972, S.

Jewett, 13 (MGCL); Hwy. 86 at Snake River, near Homstead, 20 June 1971, 1a (ADW); N. Powder Lake, 23 July 1959, J.

Baker, 12 (OSAC); North Fork Pine Creek, near Halfway, 20 June 1972, L. M. Scott, 3a (SK); North Pine Ck. nr. Halfway,

20 June 1959, S. Jewett, 183 (OSAC); same locality, 4100’, 2 July 1942, J. Baker, 1a (OSAC); 4 June 1961, R. Woodley, 1¢

(OSAC); Pine Creek, 7 July 1954, B. Weber, 33 22 (MGCL); same locality, 6 July 1969, 23 (MGCL); Pine Creek Rd.,

4000’, 6 July 1968, 1a (ADW); Spring Creek, 21 June 1956, J. Baker, 23 (OSAC); Whitman Nat. For., Rock Creek, 18 July

1943, J. Baker, 1 (MGCL); Crook County: Ochoco Mts., creek N. of Hwy. 26, 2.7 mi. SW jet. Little Hay Creek Rd. (NFD

2610), assoc. Eriogonum heracleoides var. angustifolium, 4500’, 8 Jun 2001, A. D. Warren, 33 42 (ADW); same locality and

plant association, 10 June 2003, A. D. Warren, 53 52 (ADW); Ochoco Mts., Hwy. 26 at Viewpoint Rd., 4700’, 2 July 1968,

E. Dornfeld, 1a (OSAC); same locality, 15 June 2001, D. Ross, 1d¢ (DR); Ochoco Mts., McKay Ck. at Nat. For. boundary,

14

assoc. Er. heracleoides var. angustifolium, 19 June 2002, A. D. Warren, 1a (ADW); Deschutes County: vic. summit Pine

Mtn., ca. 6400’, S. of Millican (s. of Hwy. 20), assoc. Er. heracleoides var. angustifolium and Er. umbellatum, 29 June 2003,

A. D. Warren, 73 62 (ADW); same locality and plant associations, 2 July 2003, A. D. Warren, 73 62 (ADW); Gilliam

County: Lonerock, 12 June 1965, C. W. Nelson, 23 52 (OSAC); SR 206 by Rock Creek tributary, 3.6 mi. W. confluence,

ravine in grass rangeland, 22 June 1985, D. L. Eiler, 1a 12 (MGCL); 4,5 mi. S. Condon, 2500’, 2 May 1965, E. M. & S. F.

Perkins, 1a (MGCL); same locality, 14 June 1964, E. M. & S. F. Perkins, 33 72 (MGCL); Grant County: Beech Creek

Summit, 4 July 1962, B. Weber, 33 22 (MGCL); same locality, 9 July 1962, B. Weber, 1a 12 (MGCL); Fall Mtn., SE of

John Day, 23 June 1990, P. Savage, 43 72 (MGCL); Hwy. 395, 14 mi. S. John Day, 23 June 1990, P. Savage, 1 2 (MGCL);

Sugarloaf Gulch nr. Canyon Ck., 1 August 1975, 1¢ (ADW); Summit Prairie at jct. Rd. 162 & Rd. 1427, T16S, R34E,

Malheur Nat. For., SE of Prairie City, 13 July 1970, S. K. Dvorak & O. Shields, 1¢ (MGCL); 5 rd. mi. WSW jet. Rd. 162 &

Rd. 1427, along Rd. 162, T16S R33.5E, E. of Seneca, 13 July 1970, S. K. Dvorak & O. Shields, 1¢ (MGCL); Harney

County: Blue Mts., Divine Cyn., 5036’, 29 June 1964, R. Crowe, 53 (OSAC); same locality, 18 July 1969, E. Dornfeld, 1 3

(OSAC); 1 August 1965, Hinchliff, 1¢ (OSAC); Devine Cyn., 12 mi. N. Burns, 5036’, Hwy. 395, pine forest on Rd. to Boy

Scout Camp, Thimer Ck., 16 July 1964, C. R. Crowe, 23 (MGCL); Hwy. 395, Devine Cyn. N. of Burns, 23 June 1990, P.

Savage, 1d 12 (MGCL); King Mtn. Lookout, T21S R32E 827, 8, 16 June 1996, V. Covlin, 235 22 (ADW); same locality, 25

June 1997, V. Covlin, 1d¢ 22 (ADW); Stinkingwater Mountains, 24 June 1961, J. H. Baker, 1 2 (MGCL); Jefferson County:

Warm Springs, 4 June 1967, Hinchliff, 1a (OSAC); same locality, 6 July 1969, 13 (MGCL); Lake County: Warner Mts.,

Drake Peak, 19 July 1969, 12 (MGCL); Warner Mts., Drake Peak, 8000’, 3 August 1963, Newcomer, 12 (OSAC); Warner

Mts., Drake Peak Lookout, 8222’, 27 July 1995, V. Covlin, 23 22 (ADW); Warner Mts., vic. Drake Peak Lookout on Light

Peak, 8230’, 16 July 2003, assoc. Er. heracleoides, A.D. Warren, 53 62 (ADW); Morrow County: Rd. 21, ca. 0.5 mi. N. of

rd. to Lake Penland, at mud, 16 June 2001, D. Ross, 13 (DR); Willow Ck. Rd., T45 R28E S27, 31 May 1992, V. Covlin, 12

12 (VC); Sherman County: Deschutes Cyn. vic. Jones Cyn., Er. heracleoides var. angustifolium, 1100-1200’, A. D. Warren,

23 (ADW); Umatilla County: Blue Mts., Spring Creek Cyn., 18 July 1991, A. L. Alderman, 1 ¢ (OSAC); Pearson Ck. Rd. at

NF boundary, 3100’, 8 June 1996, V. Covlin, 43 (VC); same locality, 8 June 1997, V. Covlin, 1a (VC); Pole Spr., Skyline

Rd., 4500’, 2 July 1967, 53 (OSAC); S. Fork Umatilla Riv., 13 June 1972, A. L. Alderman, 13 (OSAC); Wallowa County:

along rd. between Joseph & Hwy. 86, 17.8 rd. mi. N. jct. Hwy. 86, 1.2 rd. mi. SW Wallowa Mtn. summit, 14 July 1970, S. K.

Dvorak & O. Shields, 13 12 (MGCL); Spring along FR 66, T5S R47E S832, 5100’, 20 June 2001, V. Covlin, 3d (VC);

Wallowa Mts., Wallowa Lake, 12 July 1948, B. Weber, 1a (MGCL); same locality, 9 July 1954, B. Weber, 23 (MGCL);

Wasco County: Beaver Creek, 13 May 1972, J. Wells, 1 ¢ (MGCL); Beaver Creek, Simnasho, 2400’, 10 June 1973, Hinchliff,

43 (OSAC); same locality, 8 June 1973, Hinchliff, 23 (OSAC); gulch at top of Butler Canyon, Hwy. 197, ca. 4 air mi. N.

Tygh Valley, assoc. Er. heracleoides var. angustifolium, 4 June 2003, A. D. Warren, 83 92 (ADW)); Juniper Flat, 10 mi. W.

Maupin, 1890’, 26 May 1964, E. M. & S. F. Perkins, 4a 72 (MGCL); same locality, 13 June 1964, E. M. & S. F. Perkins, 83

82 (MGCL); roadside, Rte. 216, 1 mi. W. to 6.5 mi. E. Wapinita, 2 June 1994, B. O’Hara, 1 ¢ 22 (ADW); Shitike Ck., 4 June

1967, Hinchliff, 1a 12 (OSAC); Simnasho Rd., 15 June 1968, Hinchliff, 2a (OSAC); Warm Springs Riv. Rd., W. of

Simnasho, 2500’, 30 May 1968, Hinchliff, 1 ¢ (OSAC); 2 mi. NNW Mill Cr. Bridge, Warm Spgs. Indian Res., 2500’, 18 June

1966, E. M. & S. F. Perkins, 1a 12 (MGCL); 2 mi. SE Tygh Valley, Hwy. 197, 1400’, 30 May 1965, E. M. & S. F. Perkins,

103 162 (MGCL); same locality, 13 June 1964, E. M. & S. F. Perkins, 33 112 (MGCL); same locality, assoc. Er.

heracleoides var. angustifolium, 6 June 2000, A. D. Warren, 1a (ADW); 11.2 mi. SW The Dalles, 1750’, 17 May 1964, E. M.

& S. F. Perkins, 63 82 (MGCL); WASHINGTON: Chelan County: Swakane Cyn., 6 June 1986, R. F. Hirzel #025, 16

(MGCL); Swakane Canyon Rd., 8 July 2010, S. Kohler, 63 (SK); Swakane Spring, Swakane Canyon Rd., 8 July 2010, S.

Kohler, 293 (SK); Columbia County: Blue Mts., Tucannon River, above Camp Wooten, 8 June 1963, R. Woodley, 33

(OSAC); Kittitas County: Colockum Pass Rd., at Sheep x Trail Creeks, 2595-2640’, assoc. Er. douglasii, 47.093N,

120.351 W, 21 May 1983, M. A. Peterson, 1 ¢ (ADW); same locality and plant association, 11 May 1990, M. A. Peterson, 1 ?

(ADW); Klickitat County: 2 mi. S. Bickleton, 5 June 1986, R. F. Hirzel #021, 3 9 (MGCL); Okanogan County: Black Cyn.,

28 May 1961, R. Woodley, 1 ¢ (OSAC); Brewster, 18 May 1940, J. C. Hopfinger, 1 2 (MGCL); same locality, 11 May 1946, J.

C. Hopfinger, 12 (MGCL); same locality, 18 May 1956, J. C. Hopfinger, 23 (MGCL); 5 May 1948, J. C. Hopfinger, 1 ?

(MGCL); 20 May 1956, J. C. Hopfinger, 19 (MGCL); 6 June 1959, J. C. Hopfinger, 23 12 (OSAC); 14 June 1948, J. C.

Hopfinger, 1a (MGCL); 23 June 1959, J. C. Hopfinger, 1a 12 (OSAC); Buttermilk Butte, 5471’, assoc. Er. heracleoides,

48.30N, 120.30W, 23 July 1983, M. A. Peterson, 1 2? (ADW); Carlton, 12 June 1986, R. F. Hirzel #031, 23 (MGCL); Hart’s

Pass, 6000’, 16 July 1971, 16 (ADW); Poorman Ck., 3200’, assoc. Er. heracleoides, 48.32N, 120.25W, 18 July 1983, S. J.

Collman, 1 2? (ADW); summit, Chelan Butte, 30 May 1965, 1 2 (OSAC); Tunk Mtn., 1845m, 25 June 1982, 1¢ 12 (OSAC).

Additional Washington Records: Washington records provided by Jonathan P. Pelham, some with host plant associations.

We have not examined these specimens, but consider the identifications reliable. USA: WASHINGTON: Adams County: 5

mi. SW Sprague, June, P. A. Opler, 1a (Shields, 1977); Sprague Lake, S. shore, 1890’, 27 May 1992, R. M. Pyle; Asotin

County: Lick Creek at Dry Fork Lick Creek, 12 July 1978, J. P. Pelham, 1¢ (UWBM); Chelan County: Chelan, 1089-1140’,

B. V. Leighton, 1946; Olalla Canyon, 1220’, 25 May 1941, D. R. Orcutt, 2a (UWBM) (recorded as Philotes battoides var.

oregonensis in Leighton, 1946 [Dryden]); Cashmere, 795’, May, D. L. Bauer, 3c 32 (Shields, 1977); Chelan Butte Lookout,

15

3820-3835’, 30 May 1965, R. E. Woodley, 13; “First Crator’, 1a (MCZ) (Shields, 1977); Tumwater Canyon, Drury,

footbridge, 1310-1325’, June, D. L. Bauer, 23 (Shields, 1977); same locality, May, D. L. Bauer, 14; same locality, 18 May

1968, J. P. Pelham, 63 (UWBM); 31 May 1969, J. P. Pelham, 1a (UWBM); 1 June 1975, J. P. Pelham, 4a (UWBM); 16 June

1975, J. P. Pelham, J. Rozdilsky, A. Mesford, 53 (UWBM); 18 June 1978, J. P. Pelham, 1a (UWBM); Tumwater Canyon,

The Alps, 1500-1510’, 31 May 1970, J. P. Pelham, 3a (UWBM); Tumwater Canyon, Mica Canyon, 1600-1680’, 31 May

1970, J. P. Pelham, 33 (UWBM); Leavenworth, 1150-1180’, May, D. L. Bauer, 13 (Shields, 1977); Mills Canyon, 1150-

1220’, 15 April 1971, J. P. Pelham, 1a (UWBM); Mills Canyon, 2010-2050’, 19 May 1972, J. P. Pelham, 33 12 (UWBM);

Mills Canyon Road, 3000’, 13 July 1985, R. M. Pyle, T. L. Pyle; Swakane Canyon, 800-1200’, 28 June 1976, D. R. Pauson,

63 (1 genitalia prep.) (UWBM); Swakane Canyon, 1180-1185’, 28 May 1984, R. M. Pyle, T. L. Pyle, 1¢ (UWBM); same

locality, 1480-1600’, 8 July 1984, R. M. Pyle, T. L. Pyle, 2¢ (UWBM); Swakane Spring, 5060-5070’, 7 July 1985, J. P.

Pelham, M. A. Peterson, 23; Swakne Canyon at creek ford, 2375’, 23 May 2019, M. Weiss, V. & M. Elledge, C. Bellin, C. C.

Weiss Folder, 14; same locality, 20 June 1993, 31 May 1998, 28 May 2005, D. & J. Nunnalee slides; 23 May 1998, D. & J.

Nunnallee slides; Swakane Canyon near reservoir, 1670’, 23 May 1998, 18 June 1995, D. & J. Nunnalllee; Swakane Canyon,

2080’, 25 June 1993, 18 June, 25 June 1995, D. & J. Nunnallee; Swakane Canyon, 1670’, 25 June 1995, D. & J. Nunnallee;

Swakane Canyon, 860’, 20 June 1993, D. & J. Nunnallee slides; Swakane Canyon, 3160-3240’, 17 June 2000’, J. P. Pelham, Z.

A. Pelham, K. Hiruma, D. & J. Nunnallee, 33 WBA; Swakane Springs, 5070’, 15 August 1999, D. Nunnallee, larvae; same

locality, 31 July 1993, 22 July 2006; Swakane Springs, 5060-5070", 6 July 1991, J. P. Pelham, K. Hiruma, 213 5? (UWBM);

Slide Ridge Road, 4510’, 15 July 1997, D. & J. Nunnallee; Hatchery Creek Road, 18 June 1978, J. P. Pelham, 3 3 (UWBM);

Sand Creek, 1850-1950’, 22 May 1998, J. P. Pelham, M. A. Peterson, J. P. Pearson, 16; Sand Creek, 1640-1680’, 13 May

1989, J. P. Pelham, K. Hiruma, 23; Chumstick Mountain, summit, 5780-5829’, 12 July 1982, J. P. Pearson, 43 82; same

locality, 7 July 1985, J. P. Pelham, M. A. Peterson, 1 2; same locality, 6 July 1991, J. P. Pelham, K. Hiruma, 2 ?; same locality,

28 June 1997, K. Hiruma, 56; same locality, 3 August 2002, K. Hiruma, 1 2 worn; same locality, 13 June 2005, C. C. LaBar,

13 D&V (figured in Butterflies of Sinlahekin Wildlife Area: 162); same locality, 27 July 2011, D. & J. Nunnallee, on Er.

heracleoides at parking area, several mating pairs on the flowers and numbers of individuals, mostly females. Collected a good

sample of adults, pinned; Chumstick Mountains Road, 4560’, 12 July 1982, J. P. Pearson, 53 22; Chumstick Mountain below

summit, 5480-5580’, 12 July 1982, J. P. Pearson, 134; Buck Spring, 5525-5650’, 12 July 1982, J. P. Pearson 53 3 2; Entiat

Ridge, SE Chumstick Mountain, 5400-5450’, 13 July 1985, R. M. Pyle, T. L. Pyle; Entiat Ridge, 4900-5221’, 12 July 1984, J.

P. Pearson, 13; same locality, 26 July 1984, J. P. Pearson, 1 3; Entiat Summit Road, below Chumstick Mountain, 5160’, 7 July

1985, J. P. Pelham, M. A. Peterson, 13 12; Rattlesnake Spring, 4300-4450’ 13 July 1985, R. M. Pyle, T. L. Pyle; Mission

Creek, 1440-1520’, 27 May 1984, J. P. Pearson, M. A. Peterson, 16; East Fork Mission Creek, 2136’, 27 May 1991, K.

Hiruma, 13; Antoine Creek, 1950-1970’, 14 June 1975, J. P. Pelham, J. Rozdilsky, A. Mesford, 23; Colockum Pass Road,

1534’, 10 June 1984, R. M. Pyle, T. L. Pyle, 1a (UWBM); Derby Canyon, 2230’, 28 May 1984, D. & J. Nunnallee; Eagle

Creek Road, near end of pavement, 2190-2210’, 9 July 1988, K. Hiruma, S. Tojo, 13; Dick Mesa 3250-3320’, 3 June 1995, J.

P. Pelham, Z. A. Pelham, K. Hiruma, B. Hardbarger, 103 92, saw many; Columbia County: Stockade Springs, 4530-4540’,

26 June 1941, B. V. Leighton, 1¢ 32 (UWBM); same locality, 12 July 1964, R. E. Miller, 13; Lewis Creek, 2720-2725’, 7

July 1957, R. E. Miller; same locality, 9-10 July 1978, J. P. Pelham, 23 (UWBM); Jasper Mountain road at Wilson Creek,

1700-1800’, 9 July 1978, J. P. Pelham, T. S. Pelham; North Touchet Road, 4000’, 12 July 1956, R. E. Miller; Dayton, 1556-

1600’, 9 July 1956, R. E. Miller; Camp Wooten State Park, 2700’, 8 June 1963, R. E. Woodley, 2; same locality, 20 June

1964, R. E. Woodley, 14; Douglas County: Bridgeport, 800-860’, 22 May 1955, G. F. Tiedt, 23 (UWBM); same locality, 6

June 1955, G. F. Tiedt, 4a (UWBM); same locality, 9 June 1955, G. F. Tiedt, 12 (UWBM); McGinnis Canyon, Badger

Mountain, 3510-3600’, 17 June 1978, J. P. Pelham, 103 102 (UWBM); Ruud Canyon, Badger Mountain, 3200-3500’, 17 June

1978, J. P. Pelham, 243 22 (UWBM); Ferry County: Lambert Creek, 3480-3500’, 27 June 1977, J. P. Pelham, 1¢ (UWBM);

Herron Creek Road, 3000’, 27 June 1977, J. P. Pelham, 113 42 (UWBM); Mires Creek Road, 3300-3450’, 28 June 1983, J. P.

Pelham, M. A. Peterson, J. P. Pearson, J. A. Austin, 43; Lake Roosevelt, 1290-1320’, 29 May 2007, F. Bentler photo;

Deadman Creek Road, 4320’, 24 July 2006, C. Loggers, Rick Moore (ID’d as E. columbiae); Garfield County: Pataha Creek,

3700-3950’, 13 July 1978, J. P. Pelham, T. S. Pelham, 3; Pataha Creek, 3780’, 19 June 1991, R. L. Crawford, J. P. Pelham,

133d 12; NF 40, 4270’ 29 July 2010, D. & J. Nunnallee, on Er. heracleoides, pinned; Kittitas County: Easton, 2150’, 3 2

(CAS) (Shields, 1977); Wymer, R.R. siding, 1300’, June, D. L. Bauer, 2 3 (Shields, 1977); Robinson Canyon, 2200-2400’, L.

Montgomery, 1a (CWSC); Colockum Pass Road, 2620-3000’, 18 May 1971, J. P. Pelham, M. D. Van Buskirk, S. Hills;

Colockum Pass Road, 4900’, 17 July 1971, J. P. Pelham, F. Van Buskirk, 23 (UWBM); Colockum Pass Road, 5000’, 17 July

1971, J. P. Pelham, F. Van Buskirk, 2? (UWBM); Colockum Pass Road, 4690’, 26 June 2011, D. & J. Nunnallee, assoc. Ev.

douglasii, pinned; Colockum Pass Road, 4695-4705’, 25 June 2012, D. & J. Nunnallee, assoc. Er. douglasii, 14 collected, no

other Eriogonum present, photographed mating pair on the douglasii; same locality, 21 June 2012, D. & J. Nunnallee, 13 12;

Colockum Pass Road, 3150-3550’, 9 June 1979, J. P. Pelham, S. F. Jones, R. Williams, 1 ¢ (UWBM); Colockum Pass Road,

4240-4320’, 9 June 1979, J. P. Pelham, S. F. Jones, R. Williams, 1a (UWBM); Colockum Pass Road, unnamed creek, 3200-

3280’, 14 May 1972, J. P. Pelham, L. R. Lew, R. Seto, 23 22 (UWBM); same locality, 21 May 1983, J. P. Pelham, M. A.

Peterson, J. P. Pearson, 73; Colockum Pass Road at unnamed creek, 3880-4000’, 26 May 2001, J. P. Pelham, D. Nunnallee, B.

16

Hardwick, 43 42; same locality, 7 June 2003, J. P. Pelham, M. W. Robinson, P. Charvet, 23 12; Colockum Pass Road, ridge

W., 3120-3240’, 2 June 1984, R. E. Kirk; same locality, 21 May 1983, assoc. Eriogonum douglasii, J. P. Pelham, M. A.

Peterson, J. P. Pearson, 23 22; Colockum Pass Road at Sheep Creek x Trail Creek, 2595-2640’, 2 June 1984, R. E. Kirk; same

locality, 2 June 1979, J. P. Pelham, S. F. Jones, R. Williams, 32 (UWBM); Umtanum Creek, mouth, 1350-1450’, 14 June

1964, J. H. Shepard, 13; same locality, 17 June 1984, J. P. Pelham, M. A. Peterson, J. P. Pearson, B. Ostenson, R. L.

Crawford, 33; same locality, 23 June 1984, J. P. Pelham, M. A. Peterson, J. P. Pearson, 5 3; same locality, 2 May 2015, D. G.

James, T. James, J. James, R. James, B. James, M. Weiss, WBA; same locality, 24 June 1990, K. Hiruma, 23 32; Umtanum

Creek at Old Durr Road, 1925-1950’, 24 June 1984, J. P. Pelham, M. A. Peterson, J. P. Pearson, 1c; same locality, 16 May

1987, J. P. Pelham, K. Burgess, G. Knesal, 23: (UWBM); same locality, 12 June 1989, K. Hiruma, 13; same locality, 23 May

1992, J. P. Pelham, K. Hiruma, 13; Reecer Creek Road, 4800’, 5 July 1982, M. A. Peterson, J. P. Pearson; same locality, 24

May 2018, M. Weiss WBA; Reecer Creek Road, 4640’ 4 July 1981, J. P. Pelham, M. A. Peterson, S. Fujikawa, 13; Reecer

Creek Road, 3850’, 26 May 1991, C. Albright; Reecer Creek Road at Garrison Spring, hill SE, 4925-5035’, 22 June 1985, R.

E. Kirk; same locality, 4 June 2005, C. C. LeBar, 3; Reecer Creek Road at water tank road, 4200’, 1 May 2016, D. & J.

Nunnallee; same locality, 24 May 1979, J. P. Pelham, S. F. Jones, T. S. Pelham, 1 ¢ (UWBM); same locality, 4 July 1987, K.

Hiruma, 1 2; Reecer Creek Road, rock quarry to Garrison, 4840’, D. & J. Nunnallee, digital; Reecer Creek Road, Azure corner,

3520-3560’, 28 June 2009, D. & J. Nunnallee; same locality, 8 June 2018, M. Weiss; same locality, 2 June 2017, C. Clark, M.

Weiss, 6; same locality, 28 May 2017, M. Weiss, 6; same locality, 30 May 2016, C. Clark, M. Weiss; same locality, 8 June

2008, C. C. LaBar, 12 (D&V, figures in Butterflies of Sinlahekin Wildlife Area: 162); Reecer Creek Road at Garrison Spring,

4925-5053’, 26 June 2011, C. LaBar, J. LaBar, R. LaBar, 53 12; Reecer Creek Road at Green Canyon Road, powerline, 2160-

2240’, 24 May 1979, J. P. Pelham, S. F. Jones, T. S. Pelham, 2? (UWBM); Reecer Creek Road at Johnson Creek, 2815-2820’,

24 May 1979, J. P. Pelham, S. F. Jones, T. S. Pelham, 63 (UWBM); Reecer Creek Road, 3400-3500’, 24 May 1979, J. P.

Pelham, S. F. Jones, T. S. Pelham, 73; Wilson Creek Road at Bar 14 Road, 2050-2060’, 3 July 1981, J. P. Pelham, M. A.

Peterson, S. Fujikawa, 13; McPherson Canyon, 1450-1600’, 23 June 1984, J. P. Pelham, M. A. Peterson, J. P. Pearson, 1 3;

Red Top Lookout, summit, 5040-5361’, 30 June 1985, R. E. Kirk; Elk Heights Road, 2255-2265’, 5 June 2009, D. & J.

Nunnallee, mating pair, ovipositing female on Er. heracleoides, photos; same locality, 6 June 2009, D. & J. Nunnallee, photos;

same locality, 21 June 2011, D. & J. Nunnallee, 1 mating pair on Er. heracleoides, however all other activity was on the Er.

douglasii, as it was the best nectar available, 3 specimens collected on Er. douglasii, pinned; same locality, 26-27 May 2012,

D. & J. Nunnallee, single males, females and mating pairs common on Er. douglasii in fresh bloom. Er. heracleoides and Er.

compositum both with tight green buds, at least two weeks from flower, but nothing on them, photos; same locality, 10 June

2012, D. & J. Nunnallee, on Er. heracleoides, 2 weeks after collecting Euphilotes on Er. douglasii at same site, 8 specimens

collected, mostly 3, strictly associated with Er. heracleoides, no Euphilotes seen on Er. douglasii. The Euphilotes on Er.

heracleoides were reasonably fresh and mostly males, so it appears these belong to a new hatch separate from the earlier May

27 batch. Er. compositum was also in bloom, and Euphilotes (probably columbiae) were seen on it; Elk Heights, 2270’, 22

June 2010, D. & J. Nunnallee, digital; same locality, 14 June 2012, D. J. Nunnallee, pinned; NF 9702 to Red Top Lookout, side

road, 3600-3620’, 6 June 1987, A. Swengel; Ingals Lake trail, 5000’, 22 June 2010, D. & J. Nunnallee, digital; Horse Canyon,

Bettas Road, 2440’, 25 May 1995, D. A. Rolfs, 1; Red Top Lookout, summit, 4920-5361’, 25 June 1994, J. P. Pelham, C. & J.

Albright, B. Byers, 13; South Fork Manastash Creek, 2800’, 12 June 1988, K. Hiruma, L. Crabo, 23; Old Durr Road at

Umtanum Ck., 1930’, 23 May 1992, D. & J. Nunnallee; Umptanum Road, “The Well’, 1980’, 21 June 1992, D. & J.

Nunnallee, slides; Klickitat County: East Prong Little Klickkitat River, 3520-3590’, 15 June 2002, J. P. Pelham, K. Hiruma,

43; State Highway 142, 1630-1640’, 1 June 2013, J. P. Pelham, 3c 29; Klickitat/Yakima County: Kusshi Creek at US

Highway 97, 1945-2000’, 20 May 1959, D. Carney, 1 ¢ (UWBM); same locality, 8 May 1960, R. Cheyne, 1 ¢ (UWBM); same

locality, 15 May 1964, C. W. Nelson; Lincoln County: Hawk Creek Campground, 1290-1300’, 19 May 1979, J. H. Shepard;

Avista Park, below Long Lake Dam, Spokane River, 1394’, 14 June 2019, J. Dammarell, photos; Highline Road N., 2080’ 15

June 2019, J. Dammarell; same locality, 2 July 2019, J. Dammarell, photo; Sylvan Lake, N. of, 1700-1750’, 2 June 1991, J. P.

Pelham, R. L. Crawford, 23 22; Okanogan County: Brewster, 790-830’, 6 May 1934, A. Anderson, 1a (UWBM); same

locality, 22 June 1955, J. C. Hopfinger (USNM); same locality, 2 May 1935, J. C. Hopfinger (USNM); same locality, 11-18

May 1946, J. C. Hopfinger (AMNH); same locality, 2 May 1936, T. W. Dunkle (AMNH); same locality, May, June, P. A.

Opler, 203 112 (PMNH, AMNH, NHMLA) (Shields, 1977); Pateros, 780-790’, May, 23 12 (MCZC, MCAS) (Shields,

1977); 4 mi. E. Brewster, May, assoc. Er. heracleoides var. angustifolium, 103 192 (AMNH) (Shields, 1977); Similkameen

River, June, 13 12 (EMEC) (Shields, 1977); Black Canyon, 1280-2000’, 26 May 1946, J. C. Hopfinger (USNM); same

locality, 18 June 1959, D. Carney, 13 (UWBM); same locality, 28 May 1961, R. E. Woodley, 12; same locality, 17 May

1973, J. P. Pelham, 33 (UWBM); Winthrop, 1750-1800’, 11 June 1961, J. H. Shepard, 73; same locality, 23 June 1961, J. H.

Shepard; Strawberry Mountain, 4400-4742’, 21 June 1961, J. H. Shepard, 12; Beaver Creek, 2600’, 22 June 1966, J. H. & S.

Shepard, 13; Chopaka Creek, 2400’ 6 June 1968, R. E. Miller; Chopaka Creek Road, 3600’, 16 July 1972, R. Littlefield, 2 3

72 (UWBM); Toats Coulee Creek, 1500-2000’, W. Rogers; Chesaw, 2900’, W. Rogers; Wannacut Lake, N. end, 1860-1880’,

9 June 1968, R. Littlefield, 23 (UWBM); same locality, 10 June 1968, R. Littlefield, 1a (UWBM); same locality, 16 June

1968, R. Littlefield, 3 (UWBM); same locality, 15 June 1972, R. Littlefield, 23 (1 genitalia prep.) (UWBM); Hicks Canyon,

2300-2500’, R. Littlefield; same locality, 26 May 1973, J. P. Pelham; Thirtymile Campground, 3435-3455’, 20 June 1970, J. P.

17

Pelham, D. Carney, M. D. Van Buskirk, M. Kimura; Alta Lake (State Park), 1160’, June, 1 ¢ (AMNH) (Shields, 1977); same

locality, May, D. M. McCorkle, 1c 22 (Shields, 1977); same locality, 17 June 1959, D. Carney, 1 ¢ (UWBM); same locality,

8 June 1963, D. Carney, 1¢ 22 (UWBM); same locality, 21 May 1971, D. Carney, 43 42 (UWBM); same locality, 17 May

1973, J. P. Pelham, 53 22 (UWBM); same locality, 25 May 1975, J. P. Pelham, 33 12 (UWBM); same locality, 8 May 1977,

J. P. Pelham, 33 12 (UWBM); Boulder Creek, 3050’, 27 June 1976, J. P. Pelham, 23 12 (1 genitalia prep.) (UWBM);

Bernhardt Creek, 5200-5240’, 27-28 June 1976, J. P. Pelham, T. S. Pelham, B. Ostenson, 1 ¢ (UWBM); Mount Hull, Summit

Lake Road at Forest Service Road 350, 3225-3260’, 2 July 1976, J. P. Pelham, B. Ostenson, 1d (UWBM); Tonasket Creek,

2990’, 2 July 1976, J. P. Pelham, B. Ostenson, 1a (UWBM); Rusty Creek at Salmon Meadows Road, 3000-3120’, 28 June

1976, J. P. Pelham, T. S. Pelham, B. Ostenson, 16 (UWBM); McKay Creek, 5770-5780’, J. P. Pelham, T. S. Pelham, B.

Ostenson, 1 do (UWBM); Salmon Meadows, 4400-5780’, 28-29 June 1976, J. P. Pelham, T. S. Pelham, B. Ostenson, 1 2; State

Highway 20, 2 mi. W. Winthrop, 1810-1820’, 27 June 1963, 43 92; Salmon Creek Road, below Happy Hill, 2080-2100’, 14

June 1981, J. P. Pelham, M. A. Peterson, 93 2? (66 12 UWBM); Salmon Creek Road at Salmon Creek, 1830-1860’, 14 June

1981, J. P. Pelham, M. A. Peterson, 93 12 (UWBM); Benson Canyon, lower, 1720-1800’, 15 June 1981, J. P. Pelham, M. A.

Peterson, 223 112 (11a 92 UWBM); Libby Creek, 1550-1650’, 15 June 1981, J. P. Pelham, M. A. Peterson, 83 112 (72

UWBM); Boulder Creek 3050’, 25 June 1983, J. P. Pelham, M. A. Peterson, J. P. Pearson, J. A. Austin, 1 2; Fish Lake, W.

end, Sinlahekin Wildlife Area, 1800-1815’, 30 June 1976, J. P. Pelham, 73 12; same locality, 25 May 1987, R. E. Kirk; same

locality, 25 June 1983, J. P. Pelham, M. A. Peterson, J. P. Pearson, J. A. Austin, worn 3 2; Loomis-Oroville Road, 6 mi. W.

Oroville, Similkameen River, N. bank, 1060’, 28 May 1961, D. Carney, 22? (UWBM); Paradise Hill Road [ca. 3 mi. N.

Brewster], 1360-1370’, 29 May 1971, D. Carney, 1 2; Mission Creek, 3200’, 18 July 1983, M. A. Peterson, J. P. Pearson, S. J.

Collman, 23 12; Mission Creek, 3640’, 18 July 1983, M. A. Peterson, J. P. Pearson, S. J. Collman, 183; Mission Creek,

4165-4175’, [NF nr. Blackpine Lake, 4080], 18 July 1983, M. A. Peterson, J. P. Pearson, S. J. Collman, 9 3; same locality, 23

July 1983, M. A. Peterson, J. P. Pearson, 1; Poorman Creek Road, 3200’, 18 July 1983, M. A. Peterson, J. P. Pearson, S. J.

Collman, 13 12; Mission Creek, 3640’, 23 July 1983, M. A. Peterson, J. P. Pearson, 83; Buttermilk Butte, 5200-5471’, 23

July 1983, M. A. Peterson, J. P. Pearson, 33 62; Toats Coulee Road, 3000’, 17 July 1971, J. Hinchliff; Mary Ann Creek,

3800’, 13 July 1976, J. Hinchliff; Finley Canyon, 2550-2650’, 13 & 15 June 1984, J. Austin, 3a 12 (UWBM); Mary Ann

Creek, 3500’, 28 June 2003, D. & J. Nunnallee, adults abundant on Er. heracleoides, collected 7°, slides; Lyman Lake to

Moses Meadow Road, pass, 4080’, T34N R29E S34, 24 May 1987, R. E. Kirk; Omak Mountain, 5640-5747’, 24 May 1987, R.

E. Kirk; Aeneas Mountain, south, 3900-4589’, R. E. Kirk; Pearrygin Lake State Park, N. end, entrance road to Methow

Wildlife Recreation Area, 2160-2170’, 14 June 1987, A. Swengel; Methow Wildlife Recreation Area, road at S. end Pearrygin

Lake State Park, N. of Y intersection, 1950-1960’, 14 June 1987, A. Swengel; Goat Peak Road at Goat Peak, 2400-2450’, 9

July 1991, R. E. Kirk, J. P. Pelham, 23; NF 37, 2855-2860’, 3 July 2013, R. Bjorklund, BAMONA photo; Long Swamp road

loop, 6130’, 29 June 2015, D. & J. Nunnallee, WBA; Goat Peak trail, 5770’, 24 July 2005, C. C. LaBar, 130; Blue Lake,

Sinlahekin Wildlife Area, 1705-1715’, 28 Junwe 2015, J. P. Pelham, D. & J. Nunnallee, WBA conference; Forde Lake,

campground S., Sinlahakin Wildlife Area, 1600’, 15 June 2005, C. C. LaBar 23; Forde Lake, deer lick along rd. E. of,

Sinlahekin Wildlife Area, 1570’, 17 June 2008, C. C. LaBar 23 (D&V, figured in Butterflies of Sinlahekin Wildlife Area:

162); Sinlahekin Road, southern pot hole slough, Sinlahekin Wildlife Area, 1782’, 5 June 2003, N. G. Kondla, 1030 19,

Sinlahekin Road, middle pot hole slough, Sinlahekin Wildlife Area, 1766’ 5 June 2003, N. G. Kondla; Blue Lake Campground,

Sinlahekin Wildlife Area, 1640’, N. G. Kondla 1c; Forde Lake, 1585’, N. G. Kondla, 73; Cecil Road, forest opening,

Sinlahekin Wildlife Area, 2461’, 5 June 2003, N. G. Kondla, 1&; Conner Lake, dam area, Sinlahekin Wildlife Area, 1555’, 4

June 2003, N. G. Kondla, 63; Fish Lake Road, wet spot, Sinlahekin Wildlife Area, 1781’, 28 June 2012, C. C. LaBar; Fish

Lake Road, wet spot, puddling, Sinlahekin Wildlife Area, 1755’, 28 June 2012, C. C. LaBar, 3 (figured in Butterflies of

Sinlahekin Wildlife Area: 163); Sinlahekin Creek Road, small creek north side of road, Sinlahekin Wildlife Area, 2574’, 28

June 2012, C. C. LaBar; ridge E. Sinlahekin Creek Road, Sinlahekin Wildlife Area, 1666’, 29 June 2012, C. C. LaBar;

Sinlahekin Creek Road, wet area at small tributary, Sinlahekin Wildlife Area, 2761’, 29 June 2012, C. C. LaBar, mating pair

(figured in Butterflies of Sinlahekin Wildlife Area: 162); Hunters Camp, Sinlahekin Road, Sinlehekin Wildlife Area, 1852’ 30

June 2012, C. C. LaBar, 1 & (figured in Butterflies of Sinlahekin Wildlife Area: 163); Hunters Camp, Sinlahekin Wildlife

Area, 1775-1780’, 6 June 2013, C. C. LaBar, puddling 3 (figured in Butterflies of Sinlahekin Wildlife Area: 163); Sinlahekin

Road, 1755’, meadow north of Fish Lake, 30 June 2012, C. C. LaBar; Forde Lake deer lick along rd. E. of, Sinlahekin Wildlife

Area, 1584’, 30 June 2012, C. C. LaBar; Sinlahekin Creek Road, 2761’, 1 July 2012, C. C. LaBar; Lemanasky S., Sinlahekin

Wildlife Area, 4185’, 10 July 2006, T. Stuart, 5; Zachman, Sinlahekin Wildlife Area, 3350’, 3 July 2006, T. Stuart, 3&2

(D&V figured in Butterflies of Sinlahekin Wildlife Area: 162); Eden Valley Guest Ranch, 3290-3310’, 22-24 June 2012, D. &

J. Nunnallee, on Er. heracleoides, 12, WBA conference; ca. 12 mi. N. Brewster, D. Carney, 3¢ 32 (UWBM); Pend Oreille

County: State Highway 20, Pend Oreille River, 2050’, 11 July 1961, J. H. Shepard, 13 12; same locality, 14 July 1976, J. H.

Shepard; Monumental Mountain, 5550-5700’, 24 June 1992, J. P. Pelham 13; Spokane County: Kepple Peninsula Overlook

Trail, near beginning, Turnbull National Wildlife Refuge, 2280-2290’, 5 June 1987, A. Swengel; Kepple Overlook Trail, .2 mi.

E., Turnbull National Wildlife Refuge, 2300’, 5 June 1987, A. Swengel, several; Auto Loop Road, between Blackhorse Lake

and 30 Acre Lake, Turnbull National Wildlife Refuge, 2260-2270’, 5 June 1987, A. Swengel; Dishman Hills Natural Area,

18

2010, 21 July 1994, D. & J. Nunnallee, Video Notes; Palisades Park, W. Spokane, 2200’, 23 July 1994, D. & J. Nunnallee,

Video Notes; Hog Lake, 1960-2000’, 2 July 2011, J. Baumann; same locality, 13 May 2019, J. Dammarell, Photo; Isaacson

Lake, west of, Turnbull Wildlife Refuge, 2270-2280’, 14 June 2009, J. Baumann; Knothead Loop, Little Spokane Natural

Area, 1650’, 6 June 2010, J. Baumann, 3; Smith Road, entrance sign to Headquarters, Turnbull National Wildlife Refuge,

2250’, 12 July 1990, T. L. Pyle, 1¢ (UWBM, ID’d as E. enoptes); Pine Loop Trail, pond edge, Turnbull National Wildlife

Refuge, 2180’, 5 June 1987, A. Swengel, 4; Spokane, slope above Hangman Creek, 2250-2290’, 8 June 2008, T. Stuart, mating

pair apparently associated with Er. compositum (?), photo; Kepple Peninsula Overlook Trail, near beginning, Turnbull National

Wildlife Refuge 2280-2290’, 24 June 2010, J. Baumann; Turnbull National Wildlife Refuge, SW corner trail, 2310-2320’, 5

July 2010, J. Baumann, many; Blackhorse Lake area, Turnbull National Wildlife Refuge, 2280-2300’, 22 July 2010, J.

Baumann; vic. Stubblefield Lake, Turnbull National Wildlife Refuge, 2330-2350’, 13 June 2010, J. Buamann, 20; same

locality, 23 June 2011, J. Baumann, abundant; Stubblefield Lake, mounds north of lake, 2335-2345’, 23 June 2011, J.

Baumann, 10; W. Burnett Road, Forest-Steppe south, 2350-2360’, 8 June 2013, J. Baumann, 5; Salnave Road, 2355’, 7 July

2012, D. & J. Nunnallee, pinned; Stevens County: Northport, 1325-1350’, 1946, Leighton (WSU); Walla Walla County:

South Fork Dry Creek, 2380-2400’, 18 June 1991, R. L. Crawford, J. P. Pelham, 23; Whitman County: Wawawai, 650-700’,

30 May 1964, J. H. Shepard; Snake River, below Wawawai, 600’, 26 May 1965, J. H. Shepard, 14; Rosalia, Pine Creek,

2200’, 25 June 1964, R. E. Miller, 12; Kamiak Butte, 3200-3650’, 27 June 2002, J. H. & S. Shepard; Yakima County:

Toppenish Creek, 1300’, [3 mi. SE Fort Simcoe], 6 May 1959, D. Carney, 23 (UWBM); Oak Creek, 1800-2500’, 23 June

1962, D. Carney, 1a (UWBM); Naches, 1450-1470’, 19 June, R. B. Nagle; Ahtanum Creek, 1800-2000’, 19 May 1958, E. J.

Newcomer; same locality, 26 May 1962, E. J. Newcomer, 1d 12, ex. J. H. Shepard; Bear Canyon, 2000-2500’, 25 June 1964,

S. J. Newcomer; same locality, 6 July 1968, J. P. Pelham, M. D. Van Buskirk, P. K. Wiley, 1a 12 (UWBM); same locality, 18

May 1970, J. P. Pelham, R. M. Pyle, M. D. Van Buskirk, 1 ¢ (UWBM); same locality, 10 July 1983, R. E. Kirk, 3a 12; Mount

Adams Highway, near gravel pit, 1280-1300’, 6 May 1959, E. J. Newcomer; same locality, 10 May 1958, E. J. Newcomer;

Cottonwood Canyon, 1850-1900’, 25 June 19674, E. J. Newcomer; same locality, 16 June 1961, D. V. McCorkle, 13; Priest

Rapids at Priest Rapids Dam, 480-500’, 2 June 1959, E. E. Newcomer; same locality, 26 May 1962, E. J. Newcomer, | 3, ex. J.

H. Shepard; same locality, June, 2d (NHMLA) (Shields, 1977); Short and Dirty Ridge, 5400-5650’, 14 July 1959, J. H.

Shepard, 1a; Satus Creek at U.S. Highway 97, 1050’, 22 April 1978, J. H. Shepard, 1 3; Satus Creek, Highway 97, 1440’, 22

May 1971, J. Hinchliff; Satus Creek at U.S. Highway 97, [15 mi. SW Toppenish], 960-1200’, May, D. V. McCorkle, 23

(Shields, 1977); Blue Slide Lookout Tower, 6785’, T12N R13E S4, 14 July 1959, J. H. Shepard, 33 12; lower Cowiche

Canyon, 1450-1460’, 24 May 2011, D. & J. Nunnallee, assoc. Er. heracleoides; Cowiche Creek Canyon, 1430’, 10 May 1998,

D. & J. Nunnallee, slides; Wenas Creek,. 1800’, 16 May 1958, E. J. Newcomer; N. Wenas Road at Umtanum Creek Trailhead,

2530’, 21 May 1959, D. V. McCorkle, 33 (OSAC) (Shields, 1977); same locality, 9 June 1991, S. Masta, 13; N. Wenas Road,

2680’, 17 June 2018, C. Clark, M. Weiss, photo; same locality, 23 May 1992, D. & J. Nunnallee, assoc. Er. heracleoides,

slides; N. Wenas Road at Umtanum Creek Trailhead, 2515-2530’, 22 June 2013, D. & J. Nunnallee; Audubon Road, [Wenas

Creek], 2550-2600’, June, P. A. Opler (Shields, 1977).

Euphilotes baueri baueri (Shields, 1975)

(Figs. 61-64, 261-262, 332)

Shijimiaeoides battoides baueri Shields, 1975, Bull. Allyn Mus. (28); 15-16.

Euphilotes battoides baueri (Shields); Miller & Brown, 1981, Mem. Lepid. Soc. (2): 118; Mattoni,

1989, J. Res. Lepid. 27(3/4): 175.

Euphilotes baueri (Shields, 1975); Pratt & J. Emmel, 1998, Syst. W. N. Am. Butts. (16): 220.

Euphilotes baueri baueri (Shields, 1975); Pelham, 2008, J. Res. Lepid. 40: 249; Pelham, 2021,

http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: California: Inyo County; W. side Gilbert Pass, 6200’.

Pratt & Emmel (1998) considered FE. baueri a species distinct from E. battoides. Male genitalia of

E. baueri are typical of the battoides complex, with bifurcate valvae (Figs. 261-262). The female is

typical for the complex as well, with a ring-shaped lodix, but the antrum is much shorter than in other

members of the complex (Fig. 332). Euphilotes b. baueri flies mainly from mid-May to early June, and

the larval food plant is Eriogonum ovalifolium var. ovalifolium. The most prominent distinguishing

characteristic of E. b. baueri is the large amount of blue on the dorsal wing surfaces of females (Fig. 63).

The distribution of E. 6. baueri includes northeastern to east-central California and west-central Nevada

(Fig. 385).

19

Euphilotes baueri vernalis Pratt & J. Emmel, 1998

(Figs. 65-68)

Euphilotes battoides vernalis Pratt & J. Emmel, 1998, Syst. N. Am. Butts. (16): 221.

Euphilotes baueri vernalis Pratt & J. Emmel; Davenport, 2004a, Tax. Rept. Int. Lepid. Surv. 4(7): 10.

Euphilotes battoides vernalis Pratt & J. Emmel, 1998; Pelham, 2008, J. Res. Lepid. 40: 247.

Euphilotes baueri vernalis Pratt & J. Emmel, 1998; Pelham, 2021, Cat. Butterflies U.S. Can.

http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: California: San Bernardino County; Coxey Meadow, 1600m.

Euphilotes b. vernalis was described as a subspecies of E. battoides (Pratt & Emmel, 1998), but

Davenport (2004a) aligned it with E. baueri, citing comments by Gordon Pratt, as reason for the change.

Pratt & Emmel (1998) cited Eriogonum kennedyi var. kennedyi, a close relative of Er. ovalifolium, as the

larval food plant. Females have significant blue dorsally, and adults usually fly in April. Euphilotes b.

vernalis 1s apparently endemic to the northwestern slopes of the San Bernardino Mountains, in San

Bernardino County, California (Fig. 385).

Euphilotes baueri orientis Austin, 1998

(Figs. 69-72, 263, 333)

Euphilotes baueri orientis Austin, 1998, Syst. W. N. Am. Butts. (45): 550-551; Pelham, 2008, J. Res.

Lepid. 40: 249; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Nevada: Lincoln County; Bristol Mountains, U. S. Highway 93, 12.5 road miles north of

Pioche, 1826m, T3N R66E (unsectioned) on USGS Bristol Range SE, Nev., 7.5’ quadrangle.

Austin (1998) discussed the differences between E. 6b. baueri and the more eastern populations

which he named E. b. orientis, as follows. Euphilotes b. orientis males are slightly more purplish than

those of EF. b. baueri, have broader dark margins on the forewing, and have more marginal black on the

hindwing. Females of E. 5. orientis have much less blue dorsally and have a narrower aurora on average.

The ventral wings of both sexes of E. b. orientis have larger macules and narrower aurorae than do E. b.

baueri. The larval food plant 1s Eriogonum ovalifolium. The distribution, given in the original

description, is central and southeastern Nevada, southwestern Utah, and northwestern Arizona (Fig. 385).

Euphilotes baueri (Shields, 1975) (far N. Great Basin northward segregate)

(Figs. 73-78, 264-266, 334-335)

Warren et al., 2016, Illustrated lists of American butterflies, Revised 21 Nov. 2017, Accessed 19 Feb.,

2021, http://www.butterfliesofamerica.com.

Warren (2005) considered all populations of Euphilotes in Oregon that use Eriogonum ovalifolium

var. ovalifolium as the larval food plant conspecific with E. baueri, mainly because of their food plant

association. He discussed populations in two regions of Oregon, the Alvord Desert of Harney County and

the Crook-Deschutes County line area east of Redmond. In Crook and Deschutes counties, adults of E.

baueri closely resemble those of parapatric populations of E. g. glaucon associated with Er. umbellatum,

but males average somewhat paler blue. In southeastern Oregon, most individuals of E. baueri generally

resemble E. b. orientis, and further study may suggest that the name E. b. orientis applies to them.

Populations of the E. battoides complex occurring in the central Columbia Basin that fly in early

May (vic. Richland, Benton Co., Washington; vic. Hermiston, Morrow Co., Oregon), might represent £.

baueri, but food plant associations have not been verified, and these have not been mapped herein (Fig.

385). Additional unassociated Euphilotes populations in Washington are known to feed on Eriogonum

sphaerocephalum Douglas ex Benth., as detailed by James & Nunnallee (2011). Whether or not these

20

populations should be aligned with EF. baueri, or another member of the E. battoides complex in the

region, remains unclear. For now, none of these populations have been included in our distribution map

for E. baueri (Fig. 385).

Specimens Examined: USA: NEVADA: Humboldt County: Bloody Run Hills, SW facing slopes along Sand Pass Rd.,

5100’, 0.1-1.4 mi. W. of Sand Pass Summit, 4.7 mi. W. jct. Hwy. 95, N. of Winnemucca, 18 May 2016, assoc. Er. ovalifolium,

R. L. Romeyn, 23 (SK); OREGON: Crook County: Hwy. 126, 0.5-1.5 mi. E. Deschutes Co. line, 3050’, 8 June 2003, on Er.

ovalifolium, A.D. Warren, 53 8? (ADW); Deschutes County: Hwy. 126, 2 mi. E. Redmond at E, end Roberts AFB, 3050’, 8

June 2003, on Er. ovalifolium, A.D. Warren, 33¢ 5? (ADW); Harney County: Alvord Desert Rd., 4 mi. NE Andrews, 4300’,

8 May 1990, 3a 32 (OSAC); ca. 1 mi. E. Folly Farm Rd., 32 mi. S. jct. Hwy. 78, 4200’, 16 May 2002, on Er. ovalifolium,

A.D. Warren, 83 72 (ADW); Carlson Creek, T36S R33E S8, 17 May 1997, V. Covlin, 1a (ADW); Home Creek, 22 May

1950, S. Jewett, 1a (OSAC); Mann Lake, 25 May 1950, S. Jewett, 1a (OSAC); nr. Andrews, 26 May 1979, Neyhart, 3d 2?

(OSAC); nr. end of Whitehorse Rd., T35S R39E S828, 4287’, 25 May 1999, V. Covlin, 1 2 (ADW); Rd. W. of Alvord Desert,

T35S R34E 820, 5353’ 17 May 1997, V. Covlin, 23 52 (VC); same locality, 23 May 1998, V. Covlin, 1a (ADW), 2c (VC);

27 May 1999, V. Covlin, 1d 12 (VC); W. Alvord Desert, 2 mi. NE Andrews, 4200’, Hinchliff, 1 (OSAC); Wildhorse Ck.,

Alvord Basin, 28 May 1960, J. Baker, 1a 12 (OSAC); same locality and date, S. Jewett, 235 22 (OSAC); same locality, 28

May 1961, S. Jewett, 11a 62 (OSAC); 29 May 1965, S. Jewett, 2a 42 (OSAC); Alvord Desert, 3 mi. NE Andrews, ca.

4000’, 8 May 1990, T. Stoddard, 12 (SK); Malheur County: Trout Creek, 23 May 1950, S. Jewett, 1a (OSAC); Trout Ck.

Mts., 5500’, 15 mi. WNW McDermitt, 9 June 1980, M. Smith, 1 t (OSAC)

Additional Probable Records: USA: NEVADA: Washoe County: Sheldon NWR, vic. Badger Creek cabin, 28 June 1999,

D. Ross, 23 (DR); OREGON: Harney County: Burns, 13 June 1964, C. W. Nelson, 12 (OSAC); same locality, 30 May

1964, R. Crowe, 1a (OSAC).

Euphilotes baueri borealis Kohler — new subspecies

(Figs. 79-94, 267-269, 336-338)

ZooBank registration: urn:lsid:zoobank.org:act:}6D9C6062-C226-46B 1-B391-6E71AA8FC7B6

As early as 2000, Kohler found what appeared to be two different species of Euphilotes in the E.

battoides complex occurring at the same locality in Sanders County, Montana. The area has an extensive

stand of Eriogonum ovalifolium, but also a considerable amount of Er. heracleoides. The first Euphilotes

species, E. baueri borealis, begins to fly when the Er. ovalifolium is in the full pre-bloom stage, which

typically occurs in early May. The Er. heracleoides in the same area blooms about one month later, in

early to mid-June, and the second E. battoides complex species, E. heracleoides, (described above) is

associated with it.

Definition: Like E. b. orientis, males of E. b. borealis are a darker shade of blue dorsally than those of E.

b. baueri, but the black wing borders of E. b. borealis males average somewhat wider than those of E. 5.

orientis. Dorsally, females of E. b. borealis are dark brown with dark orange aurorae, and have no or at

most a hint of blue coloration. Ventrally, both sexes of EF. b. borealis are similar to the other described

subspecies of E. baueri, having very large macules on the forewings, which appear smudged along the

posterior margin and in the tornus. There is also considerable dark suffusion in the basal areas of both

forewings and hindwings of E. b. borealis. The male genitalia of E. b. borealis are similar to other

described subspecies of EF. baueri and other members of the E. battoides complex, in having bifurcate

valvae (Figs. 267-269). The female genitalia are typical of other described subspecies of FE. baueri in

having a lodix with a short antrum and surrounding lamella (Figs. 336-338).

Etymology: The name of this subspecies, borealis means “northern” or pertaining to the north wind”, and

denotes the northern most known population of Euphilotes baueri.

Distribution and Phenology: To date, E. baueri borealis is known from only three localities in Montana

(Fig. 385). A large population exists in the area of the type locality, the main portion of which 1s a full

section of State of Montana-owned land leased for grazing purposes, between the town of Camas Prairie

and Markle Pass, 2 miles to the north. An extensive stand of Eriogonum ovalifolium Nutt. var. pansum

Reveal, the presumed larval food plant, covers a large portion of the section and is especially prevalent in

the shallow rocky soil along the tops of several low ridges that cross the section (Fig. 376). The single

brood of the adult butterflies begins to fly when the buckwheat is in the full pre-bloom stage, which

21

usually occurs the first part of May. Also occurring in the same area is Eriogonum heracleoides var.

heracleoides, which blooms about one month later, in early to mid-June. Euphilotes heracleoides is

associated with it. Even though E. b. borealis and E. heracleoides are present in the same area, E. b.

borealis is basically finished flying before E. heracleoides start to emerge. Recently, a second population

of E. b. borealis was discovered a few miles to the northeast of the type locality, but few adults have been

obtained. In 2016, an additional population was discovered in the area of Twin Bridges, Madison County,

east of the Continental Divide, about 180 miles southeast of the type locality. This site is higher in

elevation, 5375’, compared to 2960’ at the type locality. At the Madison County locality, the adult flight

occurred around mid-June in 2016 and 2017. The presumed food plant at this locality 1s Eriogonum

ovalifolium var. ovalifolium, and was the only species of Eriogonum present. Despite being widely

distributed in much of the western two-thirds of Montana, searches of Er. ovalifolium have not resulted in

the discovery of any additional E. baueri populations. At one locality in Lewis and Clark County where

stands of Er. ovalifolium occur at 7000’ elevation, the Euphilotes that were associated with it turned out to

be E. ancilla (discussed below).

Types: Holotype male: MONTANA: Saunders County: 4.5mi N of Camas Prairie, 5 May 2004, S.

Kohler Collector (47.53086, -114.60782). Allotype female: MONTANA: Sanders County: 4.5 mi N of

Camas Prairie, 6 May 2004, S. Kohler Collector. Paratypes: MONTANA: Sanders County: 5 mi. N.

Camas Prairie, 20 May 1997, S. Kohler, 20d 332 (SK); 4.5 mi. N. Camas Prairie, 20 May 1997, S.

Kohler, 403 232 (SK); same locality, 5 May 1998, S. Kohler, 24d 152 (SK); same locality, 6 May

1998, S. Kohler, 773 642 (SK); same locality, 7 May 1998, S. Kohler, 173 72 (SK); same locality, 15

May 1998, S. Kohler, 13 1? (SK); same locality, 5 May 2004, S. Kohler, 656 462 (SK); same locality,

6 May 2004, S. Kohler, 1093 422 (SK); same locality, 17 May 2006, S. Kohler, 1° (SK); 4.5 mi. north

Camas Prairie, west Hwy. 382, 2945’, 4 May 2016, S. Kohler, 36c0 182 (SK); same locality, 2950’, 13

May 2016, S. Kohler, 3a 52 (SK); same locality, 15 May 2018, S. Kohler, 136 5¢ (SK); 1 mi. S. of

Markle Pass, 24 May 1999, S. Kohler, 3a 42 (SK). All associated with Eriogonum ovalifolium var.

pansum.

Deposition of Types: The holotype male, allotype female and four male and four female paratypes will

be deposited in the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural

History, University of Florida, Gainesville, Florida. The remaining paratypes are in the Kohler

Collection.

Type Locality: USA: Montana: Sanders County: west of Highway 382, 4.5 miles north of Camas Prairie,

2950’ (Fig. 376, upper).

Additional Specimens Examined: USA: MONTANA: Madison County: Melrose Bench Rd., 5375’, 6 mi. W. Twin

Bridges, 22 June 2016, assoc. Eriogonum ovalifolium var. ovalifolium, S. Kohler, 53 4? (SK); Melrose Bench Rd., 5400’, 15

mi. SE of Melrose, 6 June 2017, assoc. Er. ovalifolium var. ovalifolium, S. Kohler, 213 1892 (SK); Sanders County: Wilks

Gulch Road, 2790’, SE of Hot Springs, 13 May 2016, assoc. Er. ovalifolium var. pansum, S. Kohler, 13 (SK).

Euphilotes baueri shoshone Kohler — new subspecies

(Figs. 95-106, 270-271, 339-340)

ZooBank registration: urn:lsid:zoobank.org:act:7E804961-6B1A-4B14-8066-5F3E7A833FD9

Prior to 2017, Euphilotes baueri had not yet been confirmed to fly in Idaho, despite appropriate

environmental factors and the presence of Eriogonum ovalifolium. In 2017, we began a study with the

goal of locating Idaho populations of FE. baueri and assessing their characteristics. Richard L. Romeyn

facilitated obtaining the required permit, coordinating with Todd C. Stefanic, Wildlife Biologist with

Craters of the Moon National Monument. On May 30-June 2, 2017, Kohler was able to obtain a

sufficient study series of Euphilotes from the area. It represents a population isolated from adjacent

populations to the north, west and south.

Definition: Euphilotes b. shoshone males are the darkest of any of the described subspecies of E. baueri,

being a more purplish-blue than the others, with wide black wing margins. The orange of the dorsal

22

hindwing aurora is variable, being absent, or at most, consisting of two or three lunules in the tornus area.

Ventrally, the wing surfaces of both sexes are typical of other described subspecies of E. baueri, with the

large forewing macules and the smudged appearance along the posterior margin and the tornus somewhat

more noticeable. Male genitalia (Figs. 270-271) are typical for E. baueri, with bifurcate valvae. The

female genitalia are also consistent with E. baueri, having a lodix with a short antrum and surrounding

lamella (Figs. 339-340).

Etymology: Members of the Shoshone and Bannock tribes and their ancestors had the most contact

among native inhabitants with the lava fields of Craters of the Moon. Extensive archaeological evidence

suggests the Shoshone and Bannock frequently visited Craters of the Moon in the vicinity of today’s loop

drive and Highways 20, 26 and 93. The Shoshone were a branch of the Northern Shoshone that inhabited

the upper Columbia River Basin, while the Bannock were a branch of the Northern Paiute. These two

groups both occupied the Snake River Plain, intermingled, traveled and hunted together, and otherwise

coexisted while speaking slightly different languages. The Shoshone and Bannock did not live in large,

highly-structured tribes with identified chiefs. Instead they tended to spread out into small, semi-nomadic

groups or bands of two or three families in summer, searching for food throughout the plain and into the

mountain valleys (National Park Service, 2020).

Distribution and Phenology: This subspecies has been found only at Craters of the Moon National

Monument, Butte County, Idaho (Fig. 385). The larval food plant, Eriogonum ovalifolium var. focarium

Reveal & Mansfield (Craters-of-the-Moon wild buckwheat), is endemic to the Monument (Fig. 377),

flowering from May-August in black volcanic cinder flats and slopes in sagebrush communities and

conifer woodlands (Reveal & Mansfield, 2014). In our 2017 sampling, adults were flying from late May

through early June.

Types: Holotype male: IDAHO: Butte County: NE jct. Craters Loop Rd. & Tree Molds Rd., 5875’,

Craters of the Moon Nat. Mon., 1 June 2017, S Kohler Collector (43.44107, -113.54249). Allotype

female: same data as holotype. Paratypes: same data as holotype, 233 102; IDAHO: Butte County:

Crater Loop Drive, 2.5 mi. S. Visitor Center, 5975’, Craters of the Moon Nat. Mon., 30-31 May 2017, S.

Kohler, 223 112; NE jct. Craters Loop Rd. & Tree Molds Rd., 5900’, Craters of the Moon Nat. Mon., 2

June 2017, S. Kohler, 11¢ 178.

Deposition of Types: The holotype male, allotype female and the majority of the paratypes will be

deposited in the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History,

University of Florida, Gainesville, Florida. The remaining paratypes are in the Kohler Collection.

Type Locality: USA: IDAHO: Butte County: vicinity of Crater Loop Road, 5875-5975’, Craters of the

Moon National Monument (Fig. 377, upper and lower).

THE EUPHILOTES ENOPTES COMPLEX

Pelham (2021) included the following species in the Euphilotes enoptes complex: E. ancilla (W.

Barnes & McDunnough) (Figs. 292-302), E. stanfordorum Opler & A. Warren, E. mojave (F. Watson &

W. Comstock), E. enoptes (Boisduval) (Figs. 303-307), E. dammersi (J. A. Comstock & Henne), and E.

columbiae (Mattoni) (Figs. 308-309). In this species complex, the male genitalia have valvae that are,

when viewed laterally, broad, simple and flat, curving upward distally, the broad distal margin being

furnished with a row of small teeth ending ventrally in a single strong spine below which is a rather deep

excavation (Fig. 303-306). In the female genitalia, the lodix is large and exposed, with the lamellae

surrounding the ostium bursae and antrum cylindrical in shape, with the lamella postvaginalis extending

well posterior of the ostium bursae. The lodix reaches its greatest development in E. enoptes (Figs. 366-

367) and E. columbiae (Figs. 368-369).

23

Euphilotes oakleyi Kohler — new species

(Figs. 107-140, 273-291, 341-351)

Euphilotes oakleyi oakleyi Kohler

(Figs. 107-124, 273-282, 341-343)

ZooBank registration: urn:lsid:zoobank.org:act:7884CBB3-3789-449 1 -8880-785488B6BF17

Shields (1975) stated, “a hybrid specimen of battoides glaucon X enoptes ancilla has come to my

attention, a male from Pattee Canyon, 3500’, Missoula Co., Montana, VI-23-62 (J. Scott). Face on, the

left valve is like battoides and the right valve (though somewhat shorter) is like enoptes! Also, two males

from 8 mi. N. of Jarbidge, Elko Co., Nevada, VI-28-67 (J. Lane) appear to have somewhat intermediate

genitalia between battoides glaucon and enoptes ancilla (i.e., both have bifurcate valvae but not as deeply

as 1s usual in battoides, and one has prominent teeth in a row, as in enoptes, along the concave surface).

Thus it would appear likely that these three specimens are hybrids between b. glaucon and e. ancilla, two

subspecies that are nearly identical in adult facies and which utilize the same hosts, Eriogonum

umbellatum and Er. heracleoides.” Shields (1977) also stated, “Ssp. ancilla approaches the range of

battoides glaucon in western Nevada, at Pyrmid Lake and Silver City. In Elko Co., Nevada, the two

nearly overlap in range, g/aucon occurring north of Jarbidge on the Nevada-Idaho border and ancilla

flying in the Jarbidge Mts. at Pine Creek near Jarbidge, Jarbidge, 3 mi. S. Jarbidge and Bonanza Gulch.

Based on genitalia, two specimens from 8 mi. N. of Jarbidge, appear to be hybrids between the two.” He

also provided SEM photos of the valva of two of the three specimens, one from Montana and one from

Nevada, as Figs. 32-35, pp. 62-65 (Shields, 1977).

In 2001, John Christenson, Megan Hansen and Will Kerling collected a male Euphilotes in the

Rattlesnake Recreation Area (Spring Gulch) north of Missoula, Missoula Co., Montana. John Christensen

performed a genitalic dissection and made a microscope slide of the valva, subsequently showing it to

Kohler, and commenting that it could possibly be a new species, since it matched neither the E. enoptes

nor E. battoides complex. Kohler recognized the similarity of the configuration of the valva to the figures

of the supposed hybrids in Shields (1977), and the following year (2002), obtained a sufficient study

series of males from the area. After performing multiple genitalia dissections, each resembling the

Shields photos, it became obvious that we were indeed dealing with a distinct new species. Kerling was

also able to document the food plant association further up the drainage (Stuart Peak Trail) in 2002 by

collecting adults of both sexes on Eriogonum umbellatum. In 2003, Kohler collected a series of males

and females on Er. umbellatum at the same locality. Since then, populations of this species have been

found at several more localities, as detailed below.

Definition: Overall aspects of male and female genitalia seem to place this species in the Euphilotes

enoptes complex, being nearer in appearance to that group, but clearly intermediate towards the E.

battoides complex, with the extended upper prong of the valva, but not to the extent of the bifurcate valva

of members of the E. battoides complex. The lodix of the female genitalia is moderately large and

exposed. Adults of this species are much like western Montana populations of Euphilotes ancilla

(discussed below) in appearance and phenology, and because both use Er. umbellatum as a food plant, the

only reliable method of identification is by examination of the genitalia. In male E. ancilla, the terminal

margin of the cucullus of the valva forms a rather straight line with many fine teeth (Figs. 292-297). In

male E. oakleyi, the dorsal portion of the cucullus of the valva is elongated into a curved prong extending

caudally, and when teeth are present along the terminal margin, they are fewer in number and larger (Figs.

273-282). In female FE. ancilla, the shape of the lodix is typical of the E. enoptes complex, with the

lamella postvaginalis being large and fully developed and extending caudally well beyond the ostium

bursae (Figs. 352-354). In female E. oakleyi, the central portion of the lamella postvaginalis still extends

caudally, but the peripheral portions on either side are somewhat reduced (Figs. 341-343).

Etymology: This species is named for Oakley Shields, a pioneer in the modern study of the genus

Euphilotes, and the first to illustrate the distinctive valva of the male genitalia of FE. oakleyi, though he

24

considered that the specimens he figured were hybrids, and did not recognize E. oakleyi as a distinct

species.

Distribution and Phenology: To date, records indicate a rather limited distribution for E. oakleyi,

primarily in western Montana (Fig. 386). Recent field work has expanded our knowledge of its

distribution and underscores the importance of genitalic examination of all specimens encountered. The

record for Elko Co., Nevada, leaves open the possibility that this species is much more widespread over a

large geographic expanse between Montana and Nevada, but has thus far gone undetected in this

intervening area. Most of the specimens from Montana were taken in association with Er. umbellatum

(Fig. 378, upper right and lower). The flight period for the single brood is June and July, depending on

seasonal development and elevation.

Types: Holotype male: MONTANA: Missoula County: Stuart Peak Trail, Rattlesnake Mtns., 18 June

2003, S. Kohler Collector (46.98790, -113.94785). Allotype female: same data as holotype. Paratypes:

MONTANA: Missoula County: Stuart Peak Trail, Rattlesnake Mts., 18 June 2003, S. Kohler, 136 92;

same locality, 4 July 2006, S. Kohler, 63 42; same locality, 16 July 2002, W. Kerling, 23 62; Spring

Gulch, Rattlesnake Mountains, 25 June 2002, W. Kerling, 14; same locality, 6 July 2002, S. Kohler,

353; same locality, 9 July 2002, S. Kohler, 23; same locality, 16 July 2002, S. Kohler, 23; same locality,

28 June 2007, A.D. Warren, 123; same locality, 28 June 2007, S. Kohler, 3d 1°; same locality, 5360’,

28 June 2004, S. Kohler, 1c.

Deposition of Types: The holotype male, allotype female and three male and three female paratypes will

be deposited in the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural

History, University of Florida, Gainesville, Florida. The remaining paratypes are in the Kohler Collection

and in ADW research material.

Type Locality: USA: MONTANA: Missoula County: vicinity of Spring Gulch and Stuart Peak Trail, 4.5

miles up, Rattlesnake Mountains, north of Missoula (Fig. 378, lower).

Additional Specimens Examined: USA: MONTANA: Deer Lodge County: Dry Cottonwood Creek, 6 mi. E. of Galen, 19

June 2007, S. Kohler, 43 (SK); Jefferson County: Warm Springs Creek, east of Alhambra, 23 June 2005, S. Kohler, 43 (SK);

Missoula County: Miller Cr., 1 July 1975, S. Kohler, 1¢ (SK); 6 mi. up Rattlesnake Cr., 12 July 1999, W. Kerling, 1 3 (SK);

Rattlesnake Rec. Area, 7 July 2001, M. Hanson, 1 3 (SK); Mount Jumbo, 14 June 2003, W. Kerling, 23 (SK); N. end Mount

Jumbo, NE of Missoula, 16 June 2004, W. Kerling, 14 (SK); same locality, 20 June 2004, W. Kerling, 13 (SK); Mt. Baldy

#361, MPG Ranch, 10 June 2015, J. S. Pippen, 23 12 (SK); Mount Baldy, MPG Ranch, 2 July 2014, assoc. Er. umbellatum, J.

S. Pippen, 12 (SK); Whaley Pond, MPG Ranch, 22 May 2015, J. S. Pippen, 1a (SK); Whaley Draw, MPG Ranch, 4 June

2014, J. S. Pippen, 13 (SK); same locality, 15 June 2015, J. S. Pippen, 13 (SK);); Ravalli County: Bass Cr., Florence, 3 July

1976, S. Kohler, 1¢ (SK); same locality, 17 June 1980, S. Kohler, 1 ¢ (SK); same locality, 21 July 1980, S. Kohler, 33 (SK);

same locality, 13 June 1982, S. Kohler, 1¢ (SK); same locality, 8 July 1985, S. Kohler, 1 ¢ (SK); mouth 3-Mile Cr., 4000’, SE

of Florence, 4 June 2018, 73 (SK); 3-Mile Cr. Rd., 5 mi. up, 4850’, SE of Florence, 4 June 2018, S. Kohler, 23 (SK); 8-Mile

Creek, E. of Florence, 26 June 1982, S. Kohler, 13 (SK); Silver Bow County: Rocky Ridge Trailhead, Divide Cr., 3 mi. W.

Feely, 18 June 2001, S. Kohler, 22 (SK); same locality, 22 June 2001, S. Kohler, 103 82 (SK); same locality, 30 June 2007,

S. Kohler, 1 2 (SK); Rocky Ridge Trailhead, W. of Feely, 30 June 2005, S. Kohler, 43 (SK); Rocky Ridge Trailhead, 2.5 mi.

west Feely, Divide Cr., 16 June 2003, S. Kohler, 5a 12 (SK); same locality, 19 June 2003, S. Kohler, 263 122 (SK); Rocky

Ridge Trailhead, 5900’, Divide Cr., 2.5 mi. west Feely exit I-15, 21 June 2007, S. Kohler, 183 22 (SK); same locality, 14

June 2012, S. Kohler, 13a (SK).

Additional Records: The following records were given by Shields (1977) and treated as possible hybrids between

Shijimiaeoides enoptes ancilla and S. battoides glaucon. USA: MONTANA: Missoula County: Pattee Canyon, 3500’, 23

June 1962, J. A. Scott, 1a; NEVADA: Elko County: 8 mi. N. of Jarbidge, 28 June 1967, J. Lane, 23. SEM photos of the

male genitalia of two of these specimens were given by Shields (1977, Figs. 32-35). Though Shields considered these were

possible hybrids, the figures clearly show the configuration of the valve as being consistent with what we now recognize as

characteristic of Euphilotes oakleyi.

25

Euphilotes oakleyi madisonensis Kohler — new subspecies

(Figs. 125-140, 283-291, 344-351)

ZooBank registration: urn:|sid:zoobank.org:act:5C9B 1 8DF-9B4C-453F-ABOC-FBE38 1F0E3EC

A population of Euphilotes discovered in 2008, displaying the same general configuration of male

and female genitalia as E. oakleyi, but isolated geographically from it, and utilizing a larval food plant

different than Eriogonum umbellatum, warrants subspecific status.

Definition: The appearance of the wing surfaces of both sexes of EF. oakleyi madisonensis is similar to

those of E. o. oakleyi, except that males of madisonensis are a lighter shade of blue dorsally. However,

the two subspecies differ considerably in details of the genitalia of both sexes. The curved prong of the

dorsal cucullus of the valva of male E. 0. madisonensis is more pronounced and elongated (Figs. 283-291)

than in E. o. oakleyi (Figs. 273-282, in many individuals approaching in superficial appearance the shape

of valvae of the E. battoides complex, except that the spine-like process on the inner surface of the valva

at the junction of the upper prong with the lower portion of the valva of the E. battoides complex is not

present on E. o. madisonensis. The conformation of the lodix of E. 0. madisonensis also differs

substantially from that of E. o. oakleyi. The lamella postvaginalis is much reduced on E. 0. madisonensis,

leaving only shoulder-like lateral projections with the central portion being narrow and much reduced

(Figs. 344-351). These aspects of the genitalia of E. 0. madisonensis place it even more intermediate

between the E. enoptes and E. battoides complexes than E. o. oakleyi, at least in superficial appearance.

Etymology: This subspecies is named for the river valley and county in which the population was

discovered, Madison River Valley, Madison County, Montana.

Distribution and Phenology: This subspecies has thus far been found at only a few localities in

southwestern Montana (Fig 386). At the type locality, the associated food plant, Eriogonum ovalifolium

var. ovalifolium grows in scattered patches over a large grassland area on the bench east of the Madison

River (Fig. 379, upper left). Flight period of adults of the single brood is from mid-June through early

July.

Types: Holotype male: MONTANA: Madison County: Varney Road, 4 mi NW of Cameron, 27 June

2013, 5345’, S. Kohler Collector (45.22658, -111.72227). Allotype female: same data as holotype.

Paratypes: MONTANA: Madison County: Varney Road, 4 mi. NW Cameron, 7 July 2008, S. Kohler,

1d 22; same locality, 30 June 2009, S. Kohler, 1d 92; same locality, 23 June 2010, S. Kohler, 202

122; same locality, 24 June 2010, S. Kohler, 153 72; Varney Road, 5315’ 4 mi. NW Cameron, 1 July

2011, S. Kohler, 223 449; Varney Road, 5345’, 4 mi. NW Cameron, 21 June 2012, S. Kohler, 1d 132;

same locality, 27 June 2013, S. Kohler, 255 562; same locality, 28 June 2013, S. Kohler, 473 612;

Varney Road, 5300’, 4 mi. NW Cameron, 17 June 2016, S. Kohler, 23 3 282.

Deposition of Types: The holotype male, allotype female and four male and four female paratypes will

be deposited in the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural

History, University of Florida, Gainesville, Florida. The remaining paratypes are in the Kohler

Collection.

Type Locality: USA: MONTANA: Madison County: state section, Varney Road, 5300-5345’, 4 miles

NW Cameron.

Additional Specimens Examined: Madison County: S. of Camp Creek Rd., 2 mi. E. of Melrose, 30 June 2007, S. Kohler,

13 (SK); Gravelly Range Rd., 7300’, N. Nat. For. boundary, 30 June 2015, S. Kohler, 1¢ (SK); Tobacco Root Mountains, S.

Willow Ck., ca. 8.0 rd. mi. SW jct. Norwegian Creek Rd., 6600’, 30 June 2007, A.D. Warren (at mud but ID’d by genitalia),

33 (ADW); Silver Bow County: Soap Gulch, 6980’, 7 mi. up, NE Melrose, 15 June 2015, S. Kohler, 33 (SK); Soap Gulch,

6850-7150’, 7-8 mi. up, NE Melrose,17 June 2015, S. Kohler, 303 (SK); Soap Gulch, 6900’, 6 mi. up, NE Melrose, 23 June

2016, S. Kohler, 33 (SK); Soap Gulch, 6875-7300’, 6-7 mi. up, NE Melrose, 24 June 2015, S. Kohler, 203 (SK); Soap Gulch,

6850-7300’, 6-7 mi. up, NE Melrose, 25 June 2015, S. Kohler, 83 (SK); Soap Gulch, 7300’, 7 mi. up, NE Melrose, 3 July

2020, S. Kohler, 63 (SK); Soap Gulch, NE of Melrose, 15 July 2011, S. Kohler, 93 (SK); Soap Gulch, 3-6 mi. up, NE of

Melrose, 22 July 2010, S. Kohler, 303 (SK); Soap Gulch, 6-8 mi. up, NE of Melrose, 20 July 2010, S. Kohler, 103 22 (SK);

Soap Gulch, 6 mi. up, NE of Melrose, 24 July 2010, 93 12 (SK); Soap Gulch, 3-6 mi. up, NE of Melrose, 3 August 2010, S.

Kohler, 23 (SK).

26

Euphilotes ancilla ancilla (W. Barnes & McDunnough, 1918)

(Figs. 141-146, 292, 352)

[Philotes enoptes] ancilla W. Barnes & McDunnough, 1918, Contrib. Nat. Hist. Lepid. N. Am.

4(2);79.

Philotes enoptes ancilla B. & McD; McDunnough, 1938, Mem. S. Calif. Acad. Sci. 1: 28.

Philotes enoptes ancilla Barnes & McDunnough, 1918; dos Passos, 1964, Mem. Lepid. Soc. (1): 67;

Langston, 1969, J. Lepid. Soc. 23(1): 51.

Shijimaeoides enoptes ancilla (Barnes & McDunnough, 1918); Shields, 1977, J. Res. Lepid.

16(1):11.

Euphilotes enoptes ancilla (Barnes and McDunnough); Miller & Brown, 1981, Mem. Lepid. Soc. (2):

118.

Euphilotes ancilla ancilla (Barnes & McDunnough, 1918); Pratt & Emmel, 1998, Syst. W. N. Am.

Butts. (16): 213; Pelham, 2008, J. Res. Lepid. 40: 250; Pelham, 2021, Cat. Butterflies U.S. Can.

http://www. butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Utah: [Juab County]; Eureka.

Pratt & Emmel (1998) and Warren (2005) treated Euphilotes ancilla as a species level taxon.

Pratt & Emmel (1998) considered Euphilotes a. ancilla to occupy an extremely wide range (Fig. 387),

although populations along Colorado’s Front Range have since been described as a separate subspecies

(see below). E. a. ancilla resembles E. g. glaucon in the appearance of the adults, differing mainly in the

conformation of the genitalia. Throughout its range, E. a. ancilla is mainly associated with Eriogonum

umbellatum, although it uses Er. heracleoides in parts of northern Nevada.

Further study of £. a. ancilla populations is needed to determine if additional subspecies-level

taxa are represented within its broad distribution. For now, all material from western Colorado, as well as

populations in southern Wyoming which utilize Er. umbellatum var. majus (Hook.) (= Er. subalpinum

Greene) as a larval food plant are included within the concept of E. a. ancilla, although these populations

may be somewhat intermediate between FE. a. ancilla and E. a. barnesi. These southern Rocky Mountain

populations will be treated in more detail by the authors in a subsequent publication, where data from

western Colorado and southern Wyoming will be provided.

Specimens Examined: USA: IDAHO: Caribou County: Wood Canyon, 6533’, 25 June 1995, V. Covlin, 1 ¢ (ADW); Lemhi

County: (all ADW coll., all taken at mud); W. side Bitterroot Range, Cow Ck., 5520’, off Lemhi Pass Rd., E. of Tendoy, 16

July 2005, T.W. Ortenburger, 43; same locality, 16 July 2006, T.W. Ortenburger, 13; same locality, 18 July 2006, T.W.

Ortenburger, 13 12; same locality, 22 July 2006, T.W. Ortenburger, 23 12; W. side Bitterroot Range, upper Cow Creek Rd.

fork, off Lemhi Pass Rd., E. of Tendoy, 6205’, 18 July 2006, T.W. Ortenburger, 1 3; W. side Bitterroot Range, Lake Yearian

Ck. Drainage ENE of Lemhi, 6327’, 18 July 2006, T.W. Ortenburger, 113; same locality, 22 July 2006, T.W. Ortenburger,

23; W. side Bitterroot Range, Pallee Ck. Rd.,NE of Tendoy, sandbar below 3-way-junction, 5141’, 16 July 2006, T.W.

Ortenburger, 23; NEVADA: Elko County: Spruce Mountain Road, 10.6 mi. E. Hwy. 93, 8600’, 29 June 2014, R. L. Romeyn,

103 12 (SK); UTAH: Box Elder County: Willard Peak, 17 July 1970, S. Kohler, 13 (SK); Cache County: Logan Canyon,

3 July 1980, S. Kohler, 283 (SK); Green Canyon, 1 July 1964, S. Kohler, 8a (SK); same locality, 11 July 1965, S. Kohler, 1?

(SK); same locality, 2 July 1979, S. Kohler, 143 (SK); same locality, 14 July 1993, S. Kohler, 53 (SK); Green Canyon Rd., 3

mi. up, 1 July 2010, S. Kohler, 116 (SK); Green Canyon Rd., 4-5 mi. up, 2 July 2010, S. Kohler, 133 (SK); Hyde Park

Canyon, 21 June 1965, S. Kohler, 1 2 (SK); WYOMING: Fremont County: vic. Dickinson Park Guard Station, 9300’, Wind

River Range, 26 July 2017, S. Kohler, 1d (SK); Lincoln County: Swift Creek, Afton, 2 July 1978, L. P. Grey, 23 (SK);

Swift Cr., 2.5 mi. E. Afton, 7 July 1982, F. E. Holley, 1 ¢ (SK); Park County: Clarks Fork Overlook, Hwy. 212, 14 July 1982,

S. Kohler, 14 (SK); Teton County: .5 mi. S. Togwotee Pass, 9660’, 18 July 2005, S. Kohler, 1d (SK).

Za

Euphilotes ancilla barnesi Opler & M. Fisher, 2009

(Figs. 147-150, 293-294)

Euphilotes ancilla barnesi Opler & M. Fisher, 2009, In Fisher, M. S. 2009, Butterflies Colo. 4: 126-

128; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Colorado: Jefferson County; Indian Gulch, 1 mile west of Golden.

Euphilotes a. barnesi differs from E. a. ancilla by its darker grayish-white appearance below

(Opler & Fisher, 2009). We consider EF. a. barnesi to be endemic to Colorado’s Front Range, where it is

associated with Eriogonum umbellatum var. umbellatum. Records exist (from north to south) from

Larimer, Boulder, Gilpin, Clear Creek, Jefferson, Douglas, Elbert and Park counties (Stanford & Opler,

1993; ADW pers. obs. 1990-2020). For now, populations further north, along the Colorado-Wyoming

border in larimer County, and on Colorado’s west slope, which utilize Eriogonum umbellatum var. majus

(= Er. subalpinum), are considered to represent E. a. ancilla (Fig. 387), although as noted above, these

may be somewhat intermediate towards E. a. barnesi. Populations of FE. ancilla to the northeast of E. a

barnesi, in prairie habitats utilizing Er. flavum as a larval food plant, are described as a separate

subspecies, below.

Euphilotes ancilla montosa Kohler — new subspecies

(Figs. 151-170, 296, 353-361)

ZooBank registration: urn:|sid:zoobank.org:act:0945 169D-7192-487B-8A4A-1 18EA9FB68E2

Definition: FE. a montosa differs from E. a. ancilla in the blue color of the male dorsal wings, which are a

darker shade of blue in E. a. montosa, and the greater frequency of at least a partial dorsal hindwing

orange aurora in the males of E. a. montosa. The underside wing surfaces of both sexes of E. a. montosa

also have larger black macules, particularly on the hindwing, and more basal suffusion of dark scales on

both the forewing and hindwing. The macules of the underside wing surfaces of E. a. ancilla are smaller

and there is less dark basal suffusion. The usual larval food plant of E. a. montosa is Eriogonum

umbellatum, although some populations at the higher elevations utilize Eriogonum ovalifolium var.

depressum Blank. (Fig. 379, upper right and lower). Eriogonum flavum var. piperi (Greene) M. E. Jones

is also utilized at some sites (Fig. 380, upper and lower), as detailed below.

Etymology: The Latin word “montosa” means mountainous, which is descriptive of the type of habitat

where this subspecies is found.

Distribution and Phenology: This subspecies occurs in mountainous regions of western Montana,

mainly west of the Continental Divide (Fig. 387). The adult flight period of the single brood extends

through June and July, depending on elevation and seasonal development.

Types: Holotype male: MONTANA: Missoula County: Ninemile Prairie, Greenough, 14 June 1979, S.

Kohler Coll. (46.95443, -113.54590). Allotype female: MONTANA: Missoula County: Ninemile

Prairie, 16 June 2001, S. Kohler Collector. Paratypes: MONTANA: Missoula County: Ninemile Prairie,

nr. Greenough, 18 June 1977, S. Kohler, 1¢ 42; same locality, 7 June 1979, S. Kohler, 23 32; 9-Mile

Prairie, 29 June 1978, S. Kohler, 153 102; Ninemile Prairie, Greenough, 9 June 1979, S. Kohler, 162

222; same locality, 11 June 1979, S. Kohler, 183 142; same locality, 12 June 1979, S. Kohler, 213142;

Ninemile Prairie, 25 May 1992, S. Kohler, 200132; same locality, 16 June 2001, S. Kohler, 1d 19;

same locality, 26 June 2004, S. Kohler, 12; same locality, 20 June 2005, S. Kohler, 22; N. Ninemile

Prairie, 3900’, 8 June 2001, S. Kohler, 256 219°.

Deposition of Types: The holotype male, allotype female and four male and four female paratypes will

be deposited in the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural

History, University of Florida, Gainesville, Florida. The remaining paratypes are in the Kohler

Collection.

Type Locality: USA: MONTANA: Missoula County: Ninemile Prairie, northwest of Greenough.

28

Additional Specimens Examined: USA: MONTANA: Beaverhead County: Lacy Creek area, FR 73, south of Wise River,

25 June 2015, assoc. Er. umbellatum, S. Kohler, 23 (SK); May Creek area, 6340’, Hwy. 43, west of Wisdom, 25 June 2015,

assoc. Er. umbellatum, S. Kohler, 13 (SK); Gold Creek, 6600’, FR 73, 15 mi. S. Wise River, 30 June 2017, assoc. Er.

umbellatum, S. Kohler, 23 1° (SK); Broadwater County: Avalanche Gulch, 16 July 2004, S. Kohler, 23 (SK); Avalanche

Gulch, 4-8 mi. up, 30 June 2004, S. Kohler, 93 (SK); Avalanche Gulch, 6.5 mi. up, 8 July 2005, S. Kohler, 33 (SK);

Avalanche Gulch, 6-8 mi. up, 12 June 2006, S. Kohler, 30d (SK); Avalanche Gulch, 8-9 mi. up, 20 June 2007, S. Kohler, 114

(SK); Avalanche Gulch, ca. mi. 4.0-4.8, 27 June 2007, A.D. Warren, 436 (at mud) (ADW); Avalanche Gulch, ca. mi. 7.8-8.1,

27 June 2007, A.D. Warren, 1 3 (at mud) (ADW); Avalanche Gulch, ca. mi. 9.0-10.0, 4 July 2007, A.D. Warren, 113 (at mud)

(ADW); Confederate Gulch, 30 June 2004, S. Kohler, 43 32 (SK); Confederate Gulch, 8.5 mi. up, 16 June 2004, S. Kohler,

23 (SK); Carbon County: nr. Red Lodge, 13 June 1978, S. Kohler, 13 (SK); East Rosebud Lake, 6200’, 2 July 1989, B.

Vogel, 13 12 (SK); East Rosebud Lake, 6100’, 30 June 1990, B. Vogel, 23 32 (SK); ridge W. Cub Creek, 5-7 mi. SE of

Belfry, 17 May 2007, S. Kohler 12 (SK); Hollenbeck Draw, 5 mi. SE of Belfry, 5 June 2007, S. Kohler, 32 (SK); Hell

Roaring Plateau Rd., 22 July 2008, S. Kohler, 1a (SK); 2-3 mi. SW Belfry, 25 June 2008, S. Kohler, 1a 12 (SK); same

locality, 19 June 2008, S. Kohler, 1 2 (SK); 2-3 mi. SW Belfry, 4125-4300’, 11 June 2009, S. Kohler, 14 12 (SK); 2-3 mi. SW

Belfry, 4100-4325’, 17 June 2009, S. Kohler, 63 (SK); 2-3 mi. SW Belfry, 4240-4350’, 19 June 2009, S. Kohler, 33 12 (SK);

vic. MT-WY state line, Main Fork Rock Cr., 8100’, SW of Red Lodge, 23 July 2019, S. Kohler, 73 (SK); same locality, 8 July

2020, S. Kohler, 1¢ (SK); Deer Lodge County: Forest Road 683, 2 mi. E. Silver Lake, 19 July 1993, S. Kohler, 43 (SK);

same locality, 19 July 2004, S. Kohler, 13 (SK); E. Georgetown Lake, 6440’, 5 July 2001, S. Kohler, 23 32 (SK); nr. Storm

Lake Pass, 9200’, Anaconda Range, 15 July 2003, S. Kohler, 1 2 (SK); Dry Cottonwood Cr., 6 mi. E. of Galen, 19 June 2007,

S. Kohler, 53 (SK); N. Fork Cottonwood Cr., 6150’, 9 mi. east of Galen, 29 June 2013, S. Kohler, 60 42 (SK); N. Fork

Cottonwood Cr., 6475’, 10 mi. E. of Galen, 29 June 2013, S. Kohler, 23 (SK); N. Fork Cottonwood Cr., 6525’, 11 mi. E.

Galen, 5 July 2013, S. Kohler, 13 12 (SK); Cable Cr. bog, 6325’, FR 683, 2 mi. E. Silver Lake, 18 June 2015, S. Kohler, 1¢

1¢ (SK); Cable Cr. bog, 6330’, FR 683, 2 mi. E. Silver Lake, 28 June 2016, assoc. Er. umbellatum, S. Kohler, 13 (SK); S.

slope Mount Tiny, 9150’, W. Storm Lake Pass, Anaconda Range, 16 July 2018, S. Kohler, 1 + (SK); Gallatin County: Bridger

Range, Bridger Ski Bowl Area, ca. 6100’, 1 July 2007, A.D. Warren, 23 (at mud) (ADW); Glacier County: Mount Henry,

near Two Medicine, 20 July 1990, S. Kohler, 13 (SK); same locality, 22 July 2005, S. Kohler, 83 62 (SK); Spot Mountain,

near Two Medicine, 3 July 2001, S. Kohler, 12 (SK); Apistoki Trail, nr. Two Medicine, 28 July 2004, S. Kohler, 1 2 (SK);

Apistoki Trail to scenic overlook, above Two Medicine, 10 July 2017, S. Kohler, 1 ¢ 22 (SK); scenic overlook trail, above

Two Medicine, 25 June 2018, S. Kohler, 13 (SK); Granite County: W. of Phillipsburg, 30 June 1981, S. Kohler, 53 492

(SK); Black Pine Road, 3 July 1989, S. Kohler, 1¢ (SK); Black Pine Road, 5675’, 6 mi. NW Phillipsburg, 20 June 2001, S.

Kohler, 31a 72 (SK); Jefferson County: 4 mi. W. Jefferson City, 23 June 2005, S. Kohler, 23 (SK); Warm Springs Creek,

west of Alhambra, 23 June 2005, S. Kohler, 1a 12 (SK); Lewis & Clark County: Skelly Gulch, 6500’, 16 mi. NW of

Helena, 21 June 1992, B. Vogel, 13 (SK); 5 mi. W. Rogers Pass, 10 July 2001, S. Kohler, 1d (SK); 0.5 mi. N. of Wolf Creek,

2 July 2002, S. Kohler, 1¢ (SK); Cottonwood Creek, Beartooth Game Range, 22 June 2005, S. Kohler, 53 (SK); mouth of

Cottonwood Ck., 26 June 2007, A.D. Warren, 13 (at mud) (ADW)); railroad tracks, S. of Wolf Creek, 21 June 2005, S.

Kohler, 23 (SK); Elkhorn Creek, Beartooth Game Range, 13 June 2006, S. Kohler, 43 (SK); same locality, 10 July 2009, S.

Kohler, 12 (SK); W. side Rogers Pass, 7200’, 25 July 2008, B. Weber, 1 ? (SK); Crown Saddle, 7300’, Scapegoat Wilderness,

17 July 2009, S. Kohler, 23 (SK); W. side Rogers Pass, 14 July 2010, S. Kohler, 43 (SK); Dead Man Coulee, north of Wolf

Creek, 7 June 2012, S. Kohler, 1 ¢ (SK); Red Mountain, 10 mi. N. Lincoln, 29 July 2010, assoc. Er. ovalifolium, S. Kohler, 1¢

12 (SK); Red Mountain, 7840’, NE of Lincoln, 16 July 2013, assoc. Er. ovalifolium, S. Kohler, 3d (SK); trail above Copper

Camp, 7800’, NE Lincoln, 7 July 2014, assoc. Er. ovalifolium, S. Kohler, 53 82 (SK); same locality, 26 July 2014, assoc. Er.

ovalifolium, S. Kohler, 1¢ 22 (SK); Red Mountain Trail, 7400’ NE Lincoln, 15 July 2015, assoc. Er. ovalifolium, S. Kohler,

12 (SK); Red Mountain trail above Copper Camp, 7500’, NE Lincoln, 1 July 2016, assoc. Er. ovalifolium, S. Kohler, 583

202 (SK); Red Mtn. trail above Copper Camp, 7000’, N. Lincoln, 4 July 2017, assoc. Er. ovalifolium, S. Kohler, 103 62

(SK); Missoula County: Miller Cr., 19 June 1971, S. Kohler, 1 ¢ (SK); same locality, 5 June 1972, S. Kohler, 1 ¢ (SK); Twin

Creek, 24 July 1974, S. Kohler, 1a 22 (SK); Mt. Sentinel, 4 July 1975, S. Kohler, 43 (SK); Randolph Property, N. of

Waterworks Hill, 1 July 2002, S. Kohler, 23 82 (SK); Mount Jumbo, 14 June 2003, W. Kerling, 1¢ 22 (SK); Shoo Fly

Meadows, 28 June 2003, W. Kerling, 33 12 (SK); same locality, 3 July 2004, W. Kerling, 23 (SK); N. end Mount Jumbo, 16

June 2004, W. Kerling, 12 (SK); Trail 42, vic. Holland Lookout, Swan Range, 21 July 2004, S. Kohler, 1¢ 2? (SK); Park

County: Eagle Creek Camp, near Jardine, 26 June 2004, W. Kerling, 43 (SK); L. Fork Bear Creek, near Jardine, 26 June

2004, W. Kerling, 23 (SK); Powell County: Boulder Mts., Priest Pass Rd., 1.5-1.8 mi. W. pass summit, ca. 5500’, 5 July

2007, A.D. Warren, 33 12 (ADW); Boulder Mts., Telegraph Ck. Rd., 4.3-5.3 mi. SSE Hwy. 12, ca. 4800’, 5 July 2007, A.D.

Warren, 23 (at mud) (ADW); near MacDonald Pass, 29 June 1981, S. Kohler, 23 (SK); same locality, 30 June 1981, S.

Kohler, 1d (SK); NW of Deer Lodge, 30 June 1982, S. Kohler, 14 (SK); nr. Morrell Lookout, 11 July 1988, S. Kohler, 13

(SK); same locality, 16 July 1997, S. Kohler, 13 (SK); below Morrell Lookout, E. of Seeley Lake, 11 July 2000, S. Kohler,

263 212 (SK); below Morrell Lookout, 7450’, east of Seeley Lake, 17 July 2001, S. Kohler, 23 22 (SK); near Browns Lake,

2 July 2002, S. Kohler, 13 (SK); vic. Morrell Lookout, 10 mi. E. Seeley Lake, 12 June 2004, assoc. Er. flavum, S. Kohler, 13 3

29

92 (SK); same locality, 6 July 2007, assoc. Er. flavum, A.D. Warren, 363 40? (ADW); same locality, 3 July 2008, assoc. Er.

flavum, S. Kohler, 343 142 (SK); same locality, 9 July 2008, assoc. Er. flavum, S. Kohler, 533 29? (SK); Swamp Creek, 4-5

mi. NE Seeley Lake, 16 July 2008, 13 (SK); vic. Morrell Lookout, 7300’, 7 July 2009, assoc. Er. flavum, S. Kohler, 163 3 2

(SK); vic. Morrell Lookout, 7600’, 10 mi. E. Seeley Lake, 7 July 2009, assoc. Er. flavum, S. Kohler, 393 52 (SK); same

locality, 9 July 2009, assoc. Er. flavum, S. Kohler, 453 182 (SK); vic. Morrell Lookout, 10 mi. E. Seeley Lake, 23 July 2011,

assoc. Er. flavum, S. Kohler, 393 22 (SK); Morrell Lookout Rd., 6985’, 10 mi. E. Seeley Lake, 29 June 2016, S. Kohler, 83

(SK); vic. Morrell Lookout, 7400-7700’, 10 mi. E. Seeley Lake, 29 June 2016, assoc. Er. flavum, S. Kohler, 433 72 (SK);

same locality, 3 July 2017, assoc. Er. flavum, S. Kohler, 13 72 (SK); Ravalli County: 8-Mile Creek, E. of Florence, 26 June

1982, S. Kohler, 1d (SK); 3-Mile Creek, SE of Florence, 29 June 1990, S. Kohler, 13 (SK); same locality, 1 June 1992, S.

Kohler, 1d (SK); same locality, 5 June 1992, S. Kohler, 13 (SK); Bass Creek, Florence, 14 June 1988, S. Kohler, 13 (SK);

same locality, 4 July 1997, S. Kohler, 13 (SK); Silver Bow County: near Butte, 28 June 1994, S. Kohler, 1 2 (SK); head of

Soap Gulch, 7530’, NE Melrose, 2 July 2020, assoc. Er. flavum, S. Kohler, 43 (SK); Soap Gulch, NE of Melrose, 15 July

2011, S. Kohler, 103 (SK); Soap Gulch, 7725’, NE Melrose, 6 July 2013, S. Kohler, 136 (SK); Soap Gulch, 3-6 mi. up, NE of

Melrose, 22 July 2010, S. Kohler, 113 (SK); Soap Gulch, 6-8 mi. up, NE of Melrose, 20 July 2010, S. Kohler, 343 12 (SK);

Soap Gulch, 6 mi. up, NE of Melrose, 24 July 2010, S. Kohler, 93 (SK); Stillwater County: Benbow Mine Road, 9000’, 17

mi. SW Fishtail, 18 July 1989, B. Vogel, 13 (SK); Sweet Grass County: Swamp Creek Rd., 12 July 1978, S. Kohler, 13 12

(SK); Picket Pin Rd., 8500’, W. Nye, 4 July 2013, S. Kohler, 23 (SK); Teton County: near jct. Mount Wright Trail & Trail

114, 5650’, 5 July 2017, S. Kohler, 13 (SK).

Additional Records: USA: MONTANA: Carbon County: Beartooth Plateau, just N. of Wyoming border, 26 July 1994, C.

E. Harp (Lep. Soc.Seas. Sum. as Euphilotes sp.); East Rosebud Lake, 6200’, 5 July 1991, B. Vogel, 83 (BV Collection);

Gallatin County: Swan Creek Trail, 23 July 1970, C. D. Ferris, 13 12 (CDF Collection); Three Forks, June, 23° (CM)

(Shields, 1977); Glacier County: 1km E. scenic point, 10km NW East Glacier, 2195m, 12 July 1978, A Peters, 1 a6 3? (GNP

Collection); Granite County: N. Hogback Cabin, Rock Cr. Rd., 25 mi. SE Clinton, 22 June 2012, G. Marangelo, 1¢

(BAMONA photo); Judith Basin County: Blacktail Hills, Dry Wolf Cr. Rd., S. of Stanford, 20 August 1994, C. E. Harp (Lep.

Soc. Seas Sum. as Euphilotes sp.); Lewis & Clark County: Skelly Gulch, 5000’, 10 mi. WNW Helena, 17 June 1990, B.

Vogel, 12 (BV Collection); 2.5 mi. W. of Austin, FR 1805, 1 July 2002, J. Christenson, 1¢ 12 (JC Collection); same locality,

2 July 2002, J. Christenson, 23 (JC Collection); same locality, 29 July 2002, J. Christenson, 1 3 1? (JC Collection); 4 mi. S. of

Rimni, 7000’, 8 July 1990, B. Vogel, 1a 12 (BV Collection); Flesher Pass, 13 July 2002, J. Christenson, 1 3 (JC Collection);

N. Wolf Creek along tracks, 13June 2003, J. Christenson, 1 3 (JC Collection); Priest Pass, 10 July 2002, T. Aneiros, 1¢ (TA

Collection); same locality, 10 July 2002, J. Christenson, 13 (JC Collection); Rogers Pass, 16 July 1965, J. Oberfoell, 136 (JO

Collection); Stemple Pass, 14 July 2004, J. Christenson, 3 3 (JC Collection); Madison County: Antelope Flats, Forest Road

556, 1 July 2003, J. Christenson, 14 (JC Collection); Rock Creek and Madison River, 1a 32 CM) (Shields, 1977); Teepee

Creek and Madison River, 23 22 (CM) (Shields, 1977); Missoula County: Missoula, 3 May 1934, D. Smith, 1¢ (MSU

Collection); Park County: Big Creek, 2-4 mi. W. Hwy. 89, 6000-6500’, 7 July 2009, M. S. Fisher; Powell County: Basin Cr.

Rd., 19 June 2016, N. & D. Ewer (Lep. Soc. Seas. Sum.); Silver Bow County: Pipestone Pass, 6418’, 22 July 1968, W. D.

Patterson (Lep. Soc. Seas. Sum.); Sweet Grass County: E. side Crazy Mountains, 27 June 1966, J. A. Scott (Lep. Soc. Seas.

Sum.); Half Moon Campground, 6500’, Big Timber Canyon, 1 « (AME) (Shields, 1977); Half Moon Park, 6500’ Big Timber

Canyon, Crazy Mts., 28 June 1966, D. C. Ferguson, 13 (YPM); Little Timber Creek, 5743’, Crazy Mountains, 29 June 1966,

D. C. Ferguson, 1a (YPM); near Big Timber Cr. 4400’, 28 June 1966, J. A. Scott, 1d (JAS Collection) (Shields, 1977);

Swamp Cr. Rd., 5500’, nr. Big Timber, 29 June 1966, J. A. Scott (JAS Collection); same locality, 2 July 1966, J. A. Scott,

(JAS Collection); Teton County: Pine Butte Ranch, 5100’, 23 June 1998, R. E. Stanford (Lep. Soc. Seas. Sum.); Teton

Canyon, W. of Choteau, 11 July 1989, J. Oberfoell, 1a (JO Collection).

Euphilotes ancilla campestris Kohler — new subspecies

(Figs. 171-196, 297-301, 362-365)

ZooBank registration: urn:lsid:zoobank.org:act:C7CAAA9D-87F9-4AF2-9A7A-65413CEA8754

Definition: Euphilotes a. campestris adult males are a lighter blue dorsally than those of E. a. montosa,

and very rarely have even a hint of orange aurora on the hindwing. They have narrow black borders

dorsally, especially on the hindwing, where often the border is reduced to a series of spots. The adult

female orange aurora of E. a. campestris is narrower than that of E. a. montosa. The black macules on the

underside wing surfaces of both sexes of E. a. campestris are smaller than those of E. a. montosa, and

there is less basal suffusion of dark scales on both the forewing and hindwing. The male and female

genitalia of E. a. campestris are typical for E. ancilla. The larval food plant is Eriogonum flavum var.

flavum (Fig. 381, lower).

30

Etymology: The word campestris means “level, even or flat ground” or “on open plain” or “Plains-

dwelling”, which is descriptive of the type of habitat where this subspecies is found.

Distribution and Phenology: Euphilotes a. campestris is widely distributed in the prairie regions of

eastern Montana (Fig. 387), and is found near dry prairie hilltops and on low ridges with shallow soils

that are thinly vegetated, such as the type locality (Fig. 381, upper). An evaluation of existing records for

Euphilotes ancilla from the prairie regions of eastern Wyoming (and possibly far northeastern Colorado,

e.g., Weld County, as indicated by Stanford & Opler (1993)), western North Dakota, South Dakota and

Nebraska, as well as the southern portions of the Canadian provinces of Alberta and Saskatchewan,

strongly suggests that those records all represent E. a campestris. Though not recorded for North Dakota

(Royer, 2003), a more recent record for that state exists; Square Butte, Golden Valley County, North

Dakota, 12 June 2004, B. Reynolds and R. Royer. Marrone (2002) reported records for Lawrence and

Fall River counties, South Dakota. His figures (p. 174) are consistent with the appearance of E. a.

campestris. The specimens figured by Ferris & Brown (1981) on page 216 as E. enoptes ancilla from E.

Laramie, Albany County, Wyoming, 28 June 1972, are consistent with the appearance of E. a. campestris.

Hooper (1973) reported one specimen of F. ancilla taken near Elkwater in the Cypress Hills of Alberta,

Canada, 18 June 1963, one specimen at Val Marie, Saskatchewan, Canada, 25 June 1968, and one from

Rosefield, Saskatchewan, Canada, 26 June 1968. Layberry et al. (1998) map the above Canadian records

and also additional localities for Alberta and Saskatchewan on page 155. They state that these records

represent E. a. ancilla, but their discussion of the probable food plant as Eriogonum flavum, and the

appearance of the adults (Plate 11, Figs. 42, 43) confirm that all of these Canadian records most probably

also represent Euphilotes a. campestris. The period of adult flight of the single brood is mainly May and

June.

Types: Holotype male: MONTANA: Powder River Co.: .8 mi. W. Pumpkin Cr. Rd. 24 mi. W. Broadus,

S. Hwy., 13 June 2007, S. Kohler Collector (45.54154, -105.83300). Allotype female: MONTANA:

Powder River Co.: .8 mi. W. Pumpkin Cr. Rd., 24 mi. W. Broadus, S. Hwy., 15 June 2007, S. Kohler

Collector. Paratypes: same locality, 13 July 2007, S. Kohler Coll., 36d 162 (SK); same locality, 15 June

2007, S. Kohler, 83 112 (SK); hills S. Hwy 212, 24 mi. W. Broadus, 26 June 2008, S. Kohler, 263 182

(SK); hills S. of Hwy. 212, 3650’, 22 mi. W. Broadus, 10 June 2015, S. Kohler, 7a 52 (SK). All assoc.

Er. flavum.

Deposition of Types: The holotype male, allotype female and three male and three female paratypes will

be deposited in the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural

History, University of Florida, Gainesville, Florida. The remaining paratypes are in the Kohler

Collection.

Type Locality: USA: MONTANA: Powder River County: hills south of Highway 212, 0.8 miles west of

Pumpkin Creek Road, 24 miles west of Broadus.

Additional Specimens Examined: USA: MONTANA: Cascade County: bluff 2.5 mi. N. Ft. Shaw, 3815’, 8 June 2010, S.

Kohler, 43 42 (SK); 1 mi. NW Lowry Bridge, 5.5 mi. W. of Simms, 22 June 2009, S. Kohler, 13 12 (SK); Carter County:

FR 4035, Long Pines area, 36 mi. SE of Ekalaka, 14 June 2007, S. Kohler, 1o (SK); Hill County: badlands NW of Havre,

2550’, 19 May 1998, G. Lawson, 1c (SK); same locality, 24 May 2001, G. Lawson, 23 (SK); same locality, 25 May 2001, G.

Lawson, 13 (SK); Bears Paw Mountains, S. of Havre, 3 June 1986, S. Kohler, 1¢ (SK); Pondera County: hills 5 mi. W.

Conrad, 3 July 2009, S. Kohler, 13 (SK); Stillwater County: Lone Tree Road, 1.5 mi. N. Columbus, 12 June 2007, S.

Kohler, 1346 42 (SK); 8.5 mi. N. Columbus, 4600’, 15 August 1989, B. Vogel, 1 ¢ 12 (BV Collection); Teton County: rim E.

of Hwy. 287, 4500’, 7 mi. S. of Choteau, 1 July 2009, S. Kohler, 22 (SK); W. Floweree Butte, 3815’, 5 mi. NW Simms, 7

June 2010, S. Kohler, 23 12 (SK); same locality, 8 June 2010, S. Kohler, 1¢ (SK); same locality, 26 June 2010, S. Kohler,

343 342 (SK); same locality, 28 June 2010, S. Kohler, 193 212 (SK); Toole County: N. of Cameron Lake, Middle Butte,

Sweet Grass Hills, 11 July 2002, assoc. Er. flavum, S. Kohler, 83 12 (SK); Pratt Canyon, 4600’, W. Butte, Sweet Grass Hills,

24 June 2009, S. Kohler, 23 (SK); 0.8 mi. N. Cameron Lake, 4150’, Middle Butte, Sweet Grass Hills, 25 June 2019, assoc. Ev.

flavum, S. Kohler, 13 (SK); vic. Cameron Lake, 4080’, Middle Butte, Sweet Grass Hills, 22 June 2020, assoc. Er. flavum, S.

Kohler, 13 (SK); same locality, 26 June 2020, assoc. Er. flavum, S. Kohler, 13 (SK); Yellowstone County: 2 mi. east of

Waco, 23 May 1994, S. Kohler, 13 (SK); WYOMING: Laramie County: | mi. W. I-25 on Terry Ranch Rd., 4 mi. N. CO

state line, 27 June 1993, prob. assoc. Er. flavum, A.D. Warren, 1? (ADW).

Additional Records: Although not examined by us, the following records most likely represent this subspecies: USA:

MONTANA: Fergus County: 10 mi. N. Big Snowy Mts., 22 June 1989, R. A. Royer (Lep. Soc. Seas. Sum.); Liberty

31

County: Corral Cr., East Butte, Sweet Grass Hills, 14 June 2005, 14; south rim of Marias River, 3100’, N. of Ft. Benton, 19

June 1994, C. E. Harp (CEH Collection); Toole County: 3 mi. W. of Sunburst, 23 May 1987, N. G. Kondla, 1d (NGK

Collection).

Euphilotes ancilla (W. Barnes & McDunnough, 1918) (SE Oregon segregate)

Warren et al., 2016, Illustrated lists of American butterflies, Revised 21 Nov. 2017, Accessed 19 Feb.,

2021. http://www.butterfliesofamerica.com.

Warren (2005) discussed populations of Euphilotes ancilla in northern Harney County, Oregon,

found in association with Eriogonum sphaerocephalum var. sphaerocephalum. He stated that the adults

are unique among members of the E. enoptes complex in Oregon, in that ventral wing spots are large and

bold, and females have prominent, very well-developed hind wing aurorae. Before 2003, this segregate of

E. ancilla was known from just a few specimens, all from Harney County. In 2003, Warren located large

populations of FE. ancilla in the Hines-Burns area. Hundreds of individuals were observed among dense

stands of Er. sphaerocephalum var. sphaerocephalum. Eriogonum strictum was also present at the site in

Burns, but was ignored by Euphilotes adults.

In addition to the population of this segregate in the Burns-Hines area, Richard Romeyn (pers.

comm., 2021) has recently documented the occurrence of both Er. sphaerocephalum var.

sphaerocephalum and the associated EF. ancilla at multiple sites along the Highway 395 corridor south

into Modoc County, California (not mapped on Fig. 387). He noted that adults at all sites sampled in the

Warner Mountains, east of Alturus, California, were indistinguishable in phenotype from those flying at

Burns-Hines, Oregon. Thus, the overall distribution of this segregate (Fig. 387) may well be far greater

than originally thought. The use of the species name Euphilotes ancilla for these populations is tentative,

however, as they may be more closely related to E. columbiae. Further details and a formal description of

these populations will be provided by the authors in a subsequent publication.

Euphilotes ancilla giulianii Pratt & J. Emmel, 1998

Euphilotes ancilla giulianii Pratt & J. Emmel, 1998, Syst. W. N. Am. Butterflies (16): 214-215;

Pelham, 2008, J. Res. Lepid. 40: 250; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-

Cat.htm.

Type Locality: Nevada: Nye County; Grapevine Mountains, ridge 0.8km southeast of Wahguyhe Peak,

2300-2400m.

Pratt & Emmel (1998) stated that this subspecies is the most distinct of the EF. ancilla group,

looking superficially much like EF. baueri. It is different from other Eriogonum umbellatum-feeding

populations in the increased amount of blue in females, the narrow width of the black border in males, and

its small size. The larval food plant 1s Eriogonum umbellatum var. versicolor S. Stokes. The range of E.

a. giulianii 1s limited to the Grapevine Mountains and Last Chance Range of southeastern California and

southwestern Nevada (Fig. 387).

Euphilotes ancilla gilvatunica Austin, 1998

(Fig. 302)

Euphilotes ancilla gilvatunica Austin, 1998, Syst. W. N. Am. Butterflies (45): 553-554; Pelham, 2008,

J. Res. Lepid. 40: 251; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Nevada:, Washoe County; Nevada State Route 341, 3.3 road miles east of US 395,

1525m, T18N R20E S5 on USGS Steamboat, Nev. 15’ quadrangle.

32

Austin (1998) stated that E. a. gilvatunica differs from other E. ancilla taxa in its relatively large

size, the black of the outer margins of males tending to be of individual spots in each cell (especially on

the hindwing), its dark brown females, and its yellowish tinged ventral surface with relatively large

macules. Males also have less submarginal orange on the dorsal hindwing than is usual among E. a.

ancilla. This subspecies occurs in western Nevada from about the Reno, Washoe County area south into

Mineral County and east into the mountains of central Nevada (Fig. 387). The single brood has been

recorded from late May through early August (mostly in June). No food plant association was provided

in the original description, but E. a. gilvatunica reportedly uses Eriogonum umbellatum var. umbellatum

(Shields, 1977) and Eriogonum lobbii Torr. & Gray (Austin, pers. comm., 2003).

Euphilotes ancilla shieldsi Austin, 1998

Euphilotes ancilla shieldsi Austin, 1998, Syst. W. N. Am. Butterflies (45): 553; Pelham, 2008, J. Res.

Lepid. 40: 251; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Nevada: Esmeralda County; White Mountains, Trail Canyon Road, 9.5 road miles west of

Nevada state route 264, 2286m, T1S R34E S6 on USGS Davis Mountain, Nev.-Calif. 15’ quadrangle.

Austin (1998) stated that E. a. shieldsi is most similar to E. a. purpura (see below), but is not as

deeply colored, has narrower black margins and white wing fringes. The gray flush of the ventral

forewing is not as extensive, and the macules of the submargin tend to be separate rather than fused, as on

E. a. purpura. Euphilotes a. shieldsi differs from EF. a. ancilla in its deeper purplish (less blue) color,

broader black margins and larger ventral macules. This subspecies is known only from the White

Mountains, Esmeralda County, Nevada, and the Quinn Canyon Range, Nye County, Nevada (Fig. 387).

The known flight period extends from mid-June to late July. The larval food plant is Eriogonum

umbellatum.

Euphilotes ancilla purpura Austin, 1998

(Figs. 197-198)

Euphilotes ancilla purpura Austin 1998, Syst. W. N. Am. Butterflies (45): 552; Pelham, 2008, J. Res.

Lepid. 40: 251; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Nevada: Clark County; Spring Mountains, Willow Creek, 1829m, T18S R55E S2 on

USGS Mt. Stirling quadrangle, 15’ series.

Austin (1998) stated that males of EF. a. purpura are readily recognizable from those of all other

Euphilotes by their deep purple-blue color, very broad black margins, the black forewing fringe, and the

strong gray flush on the ventral forewing. He noted the distribution of this subspecies as middle

elevations of the Spring Mountains of Clark County and the road to Mt. Stirling in southern Nye County,

Nevada (Fig. 387). Austin (1998) also stated that E. a. purpura is found locally near colonies of its

probable larval food plant, Eriogonum umbellatum var. subaridum S. Stokes, and that the single brood

flies from late May to mid-August. This information was discussed, clarified, and corrected in Austin et

al. (2008), where the authors documented the existence of two cohorts of E. ancilla feeding on two

different taxa of Eriogonum in the Spring Mountains. They determined that the early season cohort flying

in May and June and utilizing Eriogonum umbellatum var. juniporinum Reveal is E. a. purpura, and that

the late season cohort flying in July-August and utilizing Er. umbellatum var. subaridum 1s E. a. cryptica

(see below).

33

Euphilotes ancilla cryptica Austin & Boyd, 2008

(Figs. 199-200)

Euphilotes ancilla cryptica Austin & Boyd, 2008, In Austin et al., J. Lepid. Soc. 62(3): 157-158;

Pelham, 2021, http://www. butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Nevada: Clark County; Spring Mountains, Cold Creek, 1825m in elevation, towards the

northern end of these mountains.

As discussed above, EF. a. cryptica was described (Austin et al., 2008) as the late season cohort of

E. ancilla in the Spring Mountains of Nevada, with larvae feeding on Eriogonum umbellatum var.

subaridum. Austin et al. (2008) stated that E. a. cryptica is distinguished from E. a. purpura by several

biological traits, including larval food plants, flight season and diapause intensity. They also stated that

size, wing pattern and genital morphology of E. a. cryptica are apparently identical with EF. a. purpura.

This taxon is known from several sites on both slopes of the Spring Mountains in Nye and Clark counties,

Nevada (Fig. 387).

Euphilotes ancilla pseudointermedia Pratt & J. Emmel, 1998

Euphilotes ancilla pseudointermedia Pratt & J. Emmel, 1998, Syst. W. N. Am. Butterflies (16): 214;

Pelham, 2008, J. Res. Lepid. 40: 251; Pelham, 2021, http://www.butterfliesofamerica.com/US-Can-

Cat.htm.

Type Locality: California: Nevada County; Castle Peak near Donner Pass, 2700m.

Pratt & Emmel (1998) stated that this subspecies is probably closely related to E. a. ancilla.

Euphilotes a. pseudointermedia 1s associated with Eriogonum umbellatum and its resemblance to E.

glaucon intermedia is remarkable. Pratt & Emmel (1998) stated that the two cannot be distinguished by

wing morphology. Males of E. a. pseudointermedia have a broad black border above and both sexes have

a narrow orange aurora on the dorsal hindwing. The distribution of E. a. pseudointermedia includes the

alpine region of the Sierra Nevada Mountains in Nevada and Mono counties, California (Fig. 387).

THE EUPHILOTES RITA COMPLEX

Pelham (2021) included the following species in the Euphilotes rita complex: E. spaldingi (W.

Barnes & McDunnough) (Figs. 310, 370), E. rita (W. Barnes & McDunnough) (Figs. 311-314, 371-374)

and E. pallescens (Tilden & Downey) (Figs. 315, 375). In this species complex, the valvae of the male

genitalia have christae, and the valva is noticeably narrowed laterally in the mid-portion and widened

distally to become paddle-shaped, with long bristle-like terminal teeth (Fig. 311). The falces are very

short, compared to the E. battoides and E. enoptes species complexes. In the female genitalia, the lodix

(genital plate) is much reduced and is elongated, narrow, and tube-shaped in £. rita and E. pallescens

(Figs. 371, 375), but more shortened and somewhat broader in E. spaldingi (Fig. 370).

Euphilotes rita rita (W. Barnes & McDunnough, 1916)

(Figs. 201-202)

Lycaena rita W. Barnes & McDunnough, 1916, Can. Entomol. 48(7): 223-224.

Philotes rita B. & McD., 1916; W. Barnes & McDunnough, 1916, Contrib. Nat. Hist. Lepid. N. Am.

3(2): Plate XI, Fig. 3; McDunnough, 1938, Mem. S. Calif. Acad. Sci. 1: 28.

Philotes rita rita (Barnes & McDunnough), 1916; dos Passos, 1964, Mem. Lepid. Soc. (1): 67;

Langston, 1969, J. Lepid. Soc. 23(1): 52.

Shijimaeoides rita rita (Barnes & McDunnough), 1916); Shields, 1975, Bull. Allyn Mus. (28): 16;

Shields, 1977, J. Res. Lepid. 16(3): 164.

34

Euphilotes rita rita (Barnes and McDunnough); Miller & Brown, 1981, Mem. Lepid. Soc. (2): 119.

Euphilotes rita rita (W. Barnes & McDunnough, 1916); Pelham, 2008, J. Res. Lepid. 40: 251; Pelham,

2021, http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Arizona: Cochise County; Huachuca Mountains, near Ramsey Canyon.

Euphilotes r. rita occurs from extreme southern Utah (although these may be closer to the Four

Corners Area segregate discussed below), south through central Arizona, then east through southern New

Mexico into western Texas (Fig. 388) (Stanford & Opler, 1993). Females have bright, broad aurorae on

the dorsal hindwings, and both sexes have bold aurorae on the ventral hindwings. Males often have a

poorly-developed aurora on the dorsal hindwing, as well. The larval food plant for E. r. rita was cited as

“close to Eriogonum wrighti” by Mattoni [1966] and Langston (1969), and was confirmed as Er. wrightii

Torr. Ex Benth. by Bailowitz & Brock (1991). Shields (1977) and Bailowitz & Brock (1991) described

the flight period of E. r. rita as mid-August to late September.

Euphilotes rita coloradensis (Mattoni, [1966])

(Figs. 203-206, 311, 371)

Philotes rita coloradensis Mattoni, 1966, J. Res. Lepid. 4(2): 88-92; Langston, 1969, J. Lepid. Soc.

23(1): 52.

Shijimiaeoides rita coloradensis (Mattoni); Shields, 1977, J. Res. Lepid. 16(3): 164.

Euphilotes rita coloradensis (Mattoni); Miller & Brown, 1981, Mem. Lepid. Soc. (2): 119.

Euphilotes rita coloradensis (Mattoni, [1966]); Pelham, 2008, J. Res. Lepid. 40: 251; Pelham, 2021,

http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Type Locality: Colorado: Lincoln County; 7 miles south of Kendrick.

Mattoni [1966] noted that in general, E. r. coloradensis is similar to E. r. rita, but that E. r.

coloradensis may be distinguished by the male having a purplish-blue overlay and wider terminal line

(dark wing margins) dorsally. In both sexes, the darker greyish underside ground color and clear halos

are diagnostic of E. r. coloradensis. Otherwise, the females were thought to be indistinguishable. Mattoni

[1966] noted the larval food plant as Eriogonum effusum. Shields (1977) also reported Er. effusum var.

effusum as a food plant for E. rita coloradensis, and stated that it and E. r. rita are distinguishable, but

closely related subspecies that intergrade in north-central New Mexico. Cary (2021) recognized both E. r.

rita and E. r. coloradensis as occurring in New Mexico; the former utilizing Er. wrightii in the southern

part of the state, and the latter utilizing Er. effusum var. effusum and Er. rotundifolium Benth. in central

and northeastern New Mexico.

At most locales, the single generation of FE. r. coloradensis flies in mid-July through August. The

currently understood distribution of E. r. coloradensis includes south-central and southeastern Wyoming,

far southwestern Nebraska (e.g., Dundy Co., see Stanford & Opler, 1993), eastern Colorado (also far NW

Colorado in Moffat Co.), and northern and central New Mexico (Fig. 388). However, subtle patterns of

geographic variation have been detected within this region; e.g., females from the southern San Luis

Valley in Colorado (Costilla Co.) have reduced aurorae on the hindwings. A review of Colorado

populations of E. rita, in conjunction with study of populations discussed below, will be completed in a

subsequent publication by the authors.

35

Euphilotes rita (W. Barnes & McDunnough, 1916) (Four Corners Area segregate)

Warren et al., 2016, Illustrated lists of American butterflies, Revised 21 Nov. 2017, Accessed 19 Feb.,

2021, http://www.butterfliesofamerica.com.

Adults in populations of F. rita examined from far northwestern New Mexico, near Aztec, and

adjacent parts of northeastern Arizona and southern Utah (Fig. 388) differ from those of E. r. rita or E. r.

coloradensis in having generally narrower dorsal (on females) and ventral aurorae, as shown by Warren et

al. (2016), and females are paler above compared to those of E. r. rita and E. r. coloradensis, with a

generally faded appearance approaching that seen in females of E. pallescens emmeli (Shields). Adults

fly in August. These populations are associated with Eriogonum leptocladon Torr. & A. Gray. Further

study of these populations is required to determine if they are unique at the subspecies-level.

Euphilotes rita montanensis Kohler — new subspecies

(Figs. 207-234, 312-314, 372-374)

ZooBank registration: urn:lsid:zoobank.org:act: BF83A4D8-77AC-4C27-B438-08B5584B2CD2

Werner et al. (2004), in Amphibians and Reptiles of Montana, divided the state into five ecological

provinces, two in the prairie regions and three in the western mountains (Fig. 389). Ecological provinces

are based on geographical features, such as mountains and plains, and on large plant communities, such as

grasslands and forests. The Great Plains Dry Grassland Province is the largest of these five provinces. It

includes everything east of the Rocky Mountains except for several island mountains ranges and the small

portion of arid land comprising the Intermountain Semi-desert Province. This latter province is the small

northward extension of a large arid region of Wyoming, the Bighorn Basin, and lies in the rain shadow of

the Beartooth Mountains. This dry desert habitat near Belfry, Montana (Fig. 382, upper), has been

fascinating to study and explore, as it is home to several species of butterflies of more arid, southerly

distributions, including such species as Cercyonis meadiii (W. H. Edwards), C. sthenele (Boisduval),

Chlosyne acastus (W. H. Edwards), Euphydryas anicia wecoeut M. Fisher, Spomer & Scott, and the more

recent discovery of Hesperopsis alpheus (W.H. Edwards). It was here that populations of Euphilotes rita

were first discovered in 2007.

Definition: The dorsal wing color of male E. 7. montanensis 1s similar to that of E. r. coloradensis but not

quite as purplish, more intermediate in shade between E. r. rita and E. r. coloradensis. The black wing

borders of E. r. montanensis are also not quite as wide as in E. r. coloradensis, but wider than in E. r. rita.

The dorsal hindwing orange aurora of female E. r. montanensis 1s narrower than in E. r. coloradensis.

The ground color of the underside wing surfaces of both sexes of E. r. montanensis is not as dark of gray

as in E. r. coloradensis, but a shade darker than E. r. rita. The series of secondary submarginal macules

on the ventral forewing of E. r. coloradensis is usually heavy and the spots more rounded, while this

series of macules in FE. r. montanensis is faint and the spots not rounded but more crescent-shaped, like E.

r. rita. The shape of the valva of the male genitalia of EF. r. montanensis also differs from both E. r. rita

and E. r. coloradensis. In these two subspecies the anal angle of the ventral portion of the cucullus of the

valva forms a fairly sharp right angle with small indentations along the terminal margin (Fig. 311). In £.

r. montanensis, the anal angle has a more curved radius and the terminal margin 1s more entire, without

the small indentations (Figs. 312-314). The adults of E. r. montanensis are closely associated with

Eriogonum pauciflorum Pursh, the presumed larval food plant (Fig. 382, lower).

Etymology: The word montanensis means “from Montana”, and denotes the primary distribution of this

subspecies. It is consistent with the nearest named subspecies, E. r. coloradensis.

Distribution and Phenology: Thus far, E. r. montanensis has been found only in a few localities near

Belfry in Carbon County, Montana, and at an adjacent area in northern Park County, Wyoming (Fig. 388).

The single brood flies from late May through June. Euphilotes r. montanensis is remarkable for having a

36

May-June period of adult flight as opposed to August-September for EF. r. rita and July-August for E. r.

coloradensis. Also remarkable is the food plant association of Eriogonum pauciflorum.

Types: Holotype male: MONTANA: Carbon Co.: Hollenbeck Draw, 5 mi. SE of Belfry, 23 May 2007,

S. Kohler Collector (45.00569, -109.04955). Allotype female: MONTANA: Carbon Co.: Hollenbeck

Draw, 5 mi SE of Belfry, 5 June 2007, S. Kohler Collector. Paratypes: MONTANA: Carbon County:

Hollenbeck Draw, 5 mi. SE of Belfry, 5 June 2007, S. Kohler, 13 (SK); same locality, 23 May 2007, S.

Kohler, 13 (SK); same locality, 9 June 2010, S. Kohler, 1a (SK); same locality, 22 June 2011, S. Kohler,

543 232 (SK); same locality, 9 June 2015, S. Kohler, 43 12 (SK); Hollenbeck Draw, 4125-4200’, 5 mi.

SE Belfry, 11 June 2009, S. Kohler, 4a 12 (SK); Hollenbeck Draw, 4050-4250’, 5 mi. SE Belfry, 12

June 2009, S. Kohler, 1a 22 (SK); same locality, 16 June 2009, S. Kohler, 8a 22 (SK); Hollenbeck

Draw, 4000-4320’, 5 mi. SE Belfry, 19 June 2009, S. Kohler, 3a 42 (SK); same locality, 8 June 2015, S.

Kohler, 3¢ 2? (SK).

Deposition of Types: The holotype male, allotype female, and three male and three female paratypes will

be deposited in the McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural

History, University of Florida, Gainesville, Florida. The remaining paratypes are in the Kohler collection.

Type Locality: USA: Montana: Carbon County: Hollenbeck Draw, 4200’, 5 miles southeast of Belfry.

Additional Specimens Examined: USA: MONTANA: Carbon County: 2-3 mi. SW Belfry, 19 June 2008, S. Kohler, 1d 12

(SK); same locality, 25 June 2008, S. Kohler, 23 12 (SK); same locality, 21 June 2011, S. Kohler, 123 42 (SK); 2-3 mi. SW

Belfry, 4125-4300’, 11 June 2009, S. Kohler, 163 142 (SK); 2-3 mi. SW Belfry, 4100-4325’, 17 June 2009, S. Kohler, 446

372 (SK); 2-3 mi. SW Belfry, 4240-4350’, 19 June 2009, S. Kohler, 93 202 (SK); 2-4 mi. SW Belfry, 25 June 2014, S.

Kohler, 1¢ 22 (SK); draw E. Hwy. 72, 8.5 mi. S. Belfry, 21 June 2011, S. Kohler, 13 (SK); E. Wolf Cr. Rd., 2-4 mi. SW

Belfry, 4200-4400’, 8 June 2015, S. Kohler, 53 62 (SK); ridge W. Cub Cr., 5-7 mi. SE Belfry, 16 May 2007, S. Kohler, 13

(SK); WYOMING: Park County: draw E. Hwy. 72, 11 mi. S. Belfry, 10 June 2009, S. Kohler, 1 2 (SK).

37

Z| ZS

Figs. 1-24. Euphilotes glaucon. Fig.1. E. g. glaucon, Nevada: Douglas Co.: Clear Creek Canyon, 2 mi. NW US 395, 29 May

1980, G.T. Austin 3 (d). Fig. 2. Same (v). Fig. 3. Same ¢ (d). Fig. 4. Same (v). Fig. 5. EF. g. glaucon, Montana: Silver Bow

Co.: Soap Gulch, 6-8 mi. up, NE of Melrose, 20 July 2010, S. Kohler 3 (d). Fig. 6. Same (v). Fig. 7. Same, 6-7 mi. up, 6850-

7300’, 25 June 2015, assoc. Er. umbellatum, S. Kohler ? (d). Fig. 8. Same (v). Fig. 9. Same, 6850-7150’, 7-8 mi. up, NE

Melrose, 17 June 2015, S. Kohler 3 (d). Fig. 10. Same (v). Fig. 11. Same, 7300’, 7 mi. up, NE Melrose, 3 July 2020, S.

Kohler 3 (d). Fig. 12. Same (v). Fig. 13. Same, 6900’, 6 mi. up, NE Melrose, 23 June 2016, S. Kohler 2 (d). Fig. 14. Same

(v). Fig. 15. £. g. glaucon, Montana: Beaverhead Co.: near Lemhi Pass, 20 July 1982, S. Kohler 3 (d). Fig. 16. Same (v).

Fig. 17. Same, Gold Cr., 6000’, FR 73, 15 mi. S. Wise River, 30 June 2017, S. Kohler 3 (d). Fig. 18. Same (v). Fig. 19. £. g.

glaucon, Montana: Ravalli Co.: Railroad Creek off Skalkaho-Rye Road, 25 June 2001, assoc. Er. umbellatum, S. Kohler 3 (d).

Fig. 20. Same (v). Fig. 21. Same 2 (d). Fig. 22. Same (v). Fig. 23. E. g. oregonensis, Oregon: Klamath Co.: Hwy. 97 at mile

226.4, 27 June 2004, T. Stoddard 3 (d). Fig. 24. Same (v). Figs. 1.6X life size and to scale. Photos by S. Kohler.

38

45

Figs. 25-38. Euphilotes glaucon. Figs. 39-48. Euphilotes heracleoides. Fig. 25. E. g. oregonensis, Antelope Desert, Sand

Creek area, flats W. Hwy. 97 at mi. 226, 4557’, 29 June 2017, R.L. Romeyn, 2? (d). Fig. 26. Same (v). Fig. 27. FE. g.

australoglaucon, California: Inyo Co.: Sierra Nevada Mts.,flats S. FR 7S01, 4.5 mi. W. Hwy. 168 via Buttermilk Rd., 6840’,

23 May 2017, R.L. Romeyn, ¢ (d). Fig. 28. Same (v). Fig. 29. Same 2? (d). Fig. 30. Same (v). Fig. 31. £. g. intermedia,

California: Shasta Co.: Sacramento River bike path, between Rock Cr. and Gold Run Cr., 0.4-1.3 mi. E. Keswick trailhead, 10

May 2015, assoc. Er. umbellatum, R.L. Romeyn ¢ (d). Fig. 32. Same (v). Fig. 33. Same ? (d). Fig. 34. Same ¢ (v). Fig. 35.

E. glaucon (N. Oregon-Washington Cascades segregate), Oregon: Clackamas/Hood River Cos.: Mt. Hood, NW Timberline

Lodge, 5700-6000’, 20 July 2003, on Er. umbellatum var. haussknechtii, A.D. Warren 3 (d). Fig. 36. Same (v). Fig. 37. Same

? (d). Fig. 38. Same (v). Fig. 39. E. heracleoides, Montana: Sanders Co.: west of Niarada, 3 June 2004, assoc. Er.

heracleoides, S. Kohler, Holotype 3 (d). Fig. 40. Same (v). Fig. 41. E. heracleoides, Montana: Sanders Co.: 1.5 mi. N. of

Niarada, 13 June 1996, S. Kohler, Allotype ? (d). Fig. 42. Same (v). Fig. 43. Same, west of Niarada, 3 June 2004, S. Kohler,

Paratype 3 (d). Fig. 44. Same (v). Fig. 45. Same, Paratype 3 (d). Fig. 46. Same (v). Fig. 47. E. heracleoides, Montana:

Sanders Co.: 2 mi. W. of Niarada, 3300’, 14 June 1996, S. Kohler, Paratype ? (d). Fig. 48. Same (v). Figs. 1.6X life size and

to scale. Photos by S. Kohler.

39

68

VE

Figs. 49-60. Euphilotes heracleoides. Figs. 61-72. Euphilotes baueri. Fig. 49. E. heracleoides, Montana: Sanders Co.: 1.5 mi.

W. of Niarada, 1 June 2000, S. Kohler, Paratype 2? (d). Fig. 50. Same (v). Fig. 51. Same, 1 mi. W. of Perma, 11 June 2013, S.

Kohler ? (d). Fig. 52. Same (v). Fig. 53. E. heracleoides, Montana: Lincoln Co.: Dancing Prairie, NW Eureka, 28 June 2011,

S. Kohler 3 (d). Fig. 54. Same (v). Fig. 55. Same, 6 July 2011, S. Kohler 2 (d). Fig. 56. Same (v). Fig. 57. E. heracleoides,

Oregon: Wallowa Co.: Blue Mts., Powwatka Ridge, ca. 11 mi. N. of Wallowa on Troy Rd., 4100’, 23 June 2002, on Er.

heracleoides, A.D. Warren 3 (d). Fig. 58. Same (v). Fig. 59. E. heracleoides, Washington: Chelan Co.: Swakane Spring,

Swakane Canyon Rd., 8 July 2010, S. Kohler 3 (d). Fig. 60. Same (v). Fig. 61. E. b. baueri, California: Inyo Co.: W. side

Gilbert Pass, 6 mi. NE Deep Springs, 19 May 2005, assoc. Er. ovalifolium, S. Kohler 3 (d). Fig. 62. Same (v). Fig. 63. Same

? (d). Fig. 64. Same (v). Fig. 65. E. b. vernalis, California: San Bernardino Co.: Ord Mts., E. & W. slopes FR 3N14 Coxey

Truck Trail, 0.3 mi. S. FR 4N16, 5785’, 30 April 2017, assoc. Er. kennedyi, R.L. Romeyn ¢ (d). Fig. 66. Same (v). Fig. 67.

Same ? (d). Fig. 68. Same (v). Fig. 69. FE. b. orientis, Nevada: Lincoln Co.: Bristol Wells Road, 3.5 mi. W. Hwy. 93, 20 May

2005, assoc. Er. ovalifolium, S. Kohler 3 (d). Fig. 70. Same (v). Fig. 71. Same 2 (d). Fig. 72. Same (v). Figs. 1.6X life size

and to scale. Photos by S. Kohler.

40

93 95

Figs. 73-96. Euphilotes baueri. Fig. 73. E. baueri (far N. Great Basin northward segregate), Nevada: Humboldt Co.: Bloody

Run Hills, SW facing slopes along Sand Pass Rd., 5100’, 0.1-1.4 mi. W. of Sand Pass Summit, 4.7 mi. W. jct. Hwy. 95, N. of

Winnemucca, 18 May 2016, assoc. Er. ovalifolium, R.L. Romeyn 3 (d). Fig. 74. Same (v). Fig. 75. Same, Oregon: Harney

Co.: ca. 1 mi. E. Folly Farm Rd., 32 mi. S. jct. Hwy. 78, 4200’, 16 May 2002, on Er. ovalifolium, A.D. Warren 3 (d). Fig. 76.

Same (v). Fig. 77. Same, Alvord Desert, 3 mi. NE Andrews, ca. 4000’, 8 May 1990, T. Stoddard 3 (d). Fig. 78. Same (v).

Fig. 79. E. b. borealis, Montana: Sanders Co.: 4.5 mi. N. of Camas Prairie, 5 May 2004, S. Kohler, Holotype 3 (d). Fig. 80.

Same (v). Fig. 81. Same, 6 May 2004, S. Kohler, Allotype ¢ (d). Fig. 82. Same (v). Fig. 83. Same, west Hwy. 382, 2950’, 15

May 2018, S. Kohler, Paratype 3 (d). Fig. 84. Same (v). Fig. 85. Same, Paratype 3 (d). Fig. 86. Same (v). Fig. 87. Same,

2945’, 4 May 2016, S. Kohler, Paratype ? (d). Fig. 88. Same (v). Fig. 89. E. b. borealis, Montana: Sanders Co.: 4.5 mi. N.

Camas Prairie, 5 May 2004, S. Kohler, Paratype 2 (d). Fig. 90. Same (v). Fig. 91. £. b. borealis, Montana: Madison Co.:

Melrose Bench Rd., 5400’, 15 mi. SE of Melrose, 6 June 2017, assoc. Er. ovalifolium var. ovalifolium, S. Kohler 3 (d). Fig.

92. Same (v). Fig. 93. Same 2 (d). Fig. 94. Same (v). Fig. 95. E. b. shoshone, Idaho: Butte Co.: NE jct. Craters Loop Rd. &

Tree Molds Rd., 5875’, Craters of the Moon Nat. Mon., 1 June 2017, S. Kohler, Holotype o (d). Fig. 96. Same (v). Figs. 1.6X

life size and to scale. Photos by S. Kohler.

4]

105 ** 106 107 108

109

113

ie foe

Figs. 97-106. Euphilotes baueri. Figs. 107-120. Euphilotes oakleyi. Fig. 97. E. b. shoshone, Idaho: Butte Co.: NE jct. Craters

Loop Rd. & Tree Molds Rd., 5875’, Craters of the Moon Nat. Mon., 1 June 2017, S. Kohler, Allotype 2 (d). Fig. 98. Same

(v). Fig. 99. Same, Paratype 3 (d). Fig. 100. Same (v). Fig. 101. Same, Paratype ? (d). Fig. 102. Same (v). Fig. 103. Same,

Crater Loop Drive, 5975’, 2.5 mi. S. Visitor Center, Craters of the Moon Nat. Mon., 30-31 May 2017, S. Kohler, Paratype 3

(d). Fig. 104. Same (v). Fig. 105. Same, Paratype 2 (d). Fig. 106. Same (v). Fig. 107. E. 0. oakleyi, Montana: Missoula Co.:

Stuart Peak Trail, Rattlesnake Mts., 18 June 2003, S. Kohler, Holotype ¢ (d). Fig. 108. Same (v). Fig. 109. Same, Allotype ?

(d). Fig. 110. Same (v). Fig. 111. Same, Paratype ¢ (d). Fig. 112. Same (v). Fig. 113. Same, Paratype 3 (d). Fig. 114.

Same (v). Fig. 115. Same, Paratype ? (d). Fig. 116. Same (v). Fig. 117. £. 0. oakleyi, Montana: Missoula Co.: Stuart Peak

Trail, Rattlesnake Mountains, 4 July 2006, S. Kohler, Paratype ? (d). Fig. 118. Same (v). Fig. 119. E. 0. oakleyi, Montana:

Silver Bow Co.: Rocky Ridge Trailhead, 5900’, Divide Cr., 2.5 mi. west Feely exit I-15, 21 June 2017, S. Kohler 3 (d). Fig.

120. Same (v). Figs. 1.6X life size and to scale. Photos by S. Kohler.

42

135

137 138

143 144

Figs. 121-140. Euphilotes oakleyi. Figs. 141-144. Euphilotes ancilla. Fig. 121. E. 0. oakleyi, Montana: Silver Bow Co.:

Rocky Ridge Trailhead, 2.5 mi. west of Feely, Divide Cr., 19 June 2003, S. Kohler 2 (d). Fig. 122. Same (v). Fig. 123. Same,

Montana: Ravalli Co.: Bass Creek, Florence, 13 June 1982, S. Kohler 3 (d). Fig. 124. Same (v). Fig. 125. E. o. madisonensis,

Montana: Madison Co.: Varney Road, 5345’, 4 mi. NW of Cameron, 27 June 2013, S. Kohler, Holotype 3 (d). Fig. 126. Same

(v). Fig. 127. Same, Paratype 3 (d). Fig. 128. Same (v). Fig. 129. Same, Allotype ? (d). Fig. 130. Same (v). Fig. 131.

Same, Paratype 3 (d). Fig. 132. Same (v). Fig. 133. Same, 28 June 2013, S. Kohler, Paratype ¢ (d). Fig. 134. Same (v).

Fig. 135. Same, 27 June 2013, S. Kohler, Paratype ? (d). Fig. 136. Same (v). Fig. 137. Same, Paratype 2 (d). Fig. 138.

Same (v). Fig. 139. E. 0. madisonensis, Montana: Madison Co.: Gravelly Range Rd., N. Nat. Forest boundary, 7300’, 30 June

2015, S. Kohler 3 (d). Fig. 140. Same (v). Fig. 141. £. a. ancilla, Utah: Cache Co.: Green Canyon Rd., 3 mi. up, 1 July 2010,

S. Kohler 3 (d). Fig. 142. Same (v). Fig. 143. E. a. ancilla, Utah: Cache Co.: Logan Canyon, 3 July 1980, S. Kohler 3 (d).

Fig. 144. Same (v). Figs. 1.6X life size and to scale. Photos by S. Kohler.

43

145

157 ' ig =O” | 159 160

161

165 ) | 167

Figs. 145-168. Euphilotes ancilla. Fig. 145. E. a. ancilla, Utah: Cache Co.: Hyde Park Canyon, 21 June 1965, S. Kohler ¢ (d).

Fig. 146. Same (v). Fig. 147. E. a. barnesi, Colorado: Jefferson Co.: Ralston Creek, 7500’, GGCSP East Gate, 22 June 2005,

C. Slater 3 (d). Fig. 148. Same (v). Fig. 149. Same 3 (d). Fig. 150. Same (v). Fig. 151. E. a. montosa, Montana: Missoula

Co.: Ninemile Prairie, Greenough, 14 June 1979, S. Kohler, Holotype 3 (d). Fig. 152. Same (v). Fig. 153. Same, 16 June

2001, S. Kohler, Allotype ? (d). Fig. 154. Same (v). Fig. 155. Same, 25 May 1992, S. Kohler, Paratype ¢ (d). Fig. 156.

Same (v). Fig. 157. Same, Paratype 3 (d). Fig. 158. Same (v). Fig. 159. Same, 12 June 1979, S. Kohler, Paratype 2 (d). Fig.

160. Same (v). Fig. 161. Same, 9 June 1979, S. Kohler, Paratype 2 (d). Fig. 162. Same (v). Fig. 163. E. a. montosa,

Montana: Powell Co.: vic. Morrell Lookout, 10 mi. E. Seeley Lake, 12 June 2004, assoc. Er. flavum, S. Kohler 3 (d). Fig.

164. Same (v). Fig. 165. Same 2 (d). Fig. 166. Same (v). Fig. 167. E. a. montosa, Montana: Lewis & Clark Co.: Red Mtn.

trail above Copper Camp, 7500’, NE Lincoln, 1 July 2016, assoc. Er. ovalifolium, S. Kohler 3 (d). Fig. 168. Same (v). Figs.

1.6X life size and to scale. Photos by S. Kohler.

44

189 190 | 191 192

Figs. 169-192. Euphilotes ancilla. Fig. 169. E. a. montosa, Montana: Lewis & Clark Co.: Red Mtn. trail above Copper Camp,

7500’, NE Lincoln, 1 July 2016, assoc. Er. ovalifolium, S. Kohler ? (d). Fig. 170. Same (v). Fig. 171. E. a. campestris,

Montana: Powder River Co.: .8 mi. W. Pumpkin Cr. Rd., 24 mi. W. Broadus, S. Hwy., 13 June 2007, S. Kohler, Holotype 3

(d). Fig. 172. Same (v). Fig. 173. Same, 15 June 2007, S. Kohler, Allotype ? (d). Fig. 174. Same (v). Fig. 175. Same, 13

June 2007, S. Kohler, Paratype 3 (d). Fig. 176. Same (v). Fig. 177. Same, Paratype 3 (d). Fig. 178. Same (v). Fig. 179.

Same, Paratype 3 (d). Fig. 180. Same (v). Fig. 181. Same, 15 June 2007, Paratype ¢ (d). Fig. 182. Same (v). Fig. 183.

Same, 13 June 2007, S. Kohler, Paratype 2 (d). Fig. 184. Same (v). Fig. 185. E. a. campestris, Montana: Hill Co.: badlands

NW Havre, 2550’, 24 May 2001, G. Lawson & (d). Fig. 186. Same (v). Fig. 187. E. a. campestris, Montana: Stillwater Co.:

Lone Tree Road, 15 mi. N. Columbus, 12 June 2007, S. Kohler 3 (d). Fig. 188. Same (v). Fig. 189. Same ¢ (d). Fig. 190.

Same (v). Fig. 191. E. a. campestris, Montana: Teton Co.: W. Floweree Butte, 3815’, 5 mi. NW Simms, 26 June 2010, S.

Kohler 3 (d). Fig. 192. Same (v). Figs. 1.6X life size and to scale. Photos by S. Kohler.

45

197 . 198 ) 199 200

205

209

Ale)

Figs. 193-200. Euphilotes ancilla. Figs. 201-216. Euphilotes rita. Fig. 193. E. a. campestris, Montana: Teton Co.: W.

Floweree Butte, 3815’, 5 mi. NW Simms, 26 June 2010, S. Kohler 2 (d). Fig. 194. Same (v). Fig. 195. E. a. campestris,

Montana: Toole Co.: vic. Cameron Lake, 4080’, Middle Butte Sweet Grass Hills, 26 June 2020, S. Kohler 3 (d). Fig. 196.

Same (v). Fig. 197. E. a. purpura, Nevada: Clark Co.: Willow Creek, Spring Range, 5 June 1979, G.T. Austin 3 (d). Fig.

198. Same (v). Fig. 199. E. a. cryptica, Nevada: Clark Co.: Spring Mts., Willow Cr., 6000’, 20 July 2000, assoc. Er.

umbellatum var. subaridum, B. Boyd ¢ (d). Fig. 200. Same (v). Fig. 201. E. r. rita, Arizona: Cochise Co.: Ramsey Canyon,

Huachuca Mts., 3 September 1978 ¢& (d). Fig. 202. Same (v). Fig. 203. E. r. coloradensis, Colorado: El Paso Co.: Ramah Rd.,

Hwy. 94, 6100’, 6 August 1973, M. Fisher & (d). Fig. 204. Same (v). Fig. 205. Same ¢ (d). Fig. 206. Same (v). Fig. 207. £.

r. montanensis, Montana: Carbon Co.: Hollenbeck Draw, 5 mi. SE of Belfry, 23 May 2007, S. Kohler, Holotype 3 (d). Fig.

208. Same (v). Fig. 209. Same, Allotype 2 (d). Fig. 210. Same (v). Fig. 211. Same, 4200’, Paratype 3 (d). Fig 212. Same

(v). Fig. 213. Same, 16 June 2009, Paratype 3 (d). Fig. 214. Same (v). Fig. 215. E. r. montanensis, Montana: Carbon Co.:

Hollenbeck Draw, 5 mi. S. of Belfry, 22 June 2011, S. Kohler, Paratype 3 (d). Fig. 216. Same (v). Figs. 1.6X life size and to

scale. Photos by S. Kohler.

46

233

Figs. 217-234. Euphilotes rita. Fig. 217. E. r. montanensis, Montana: Carbon Co.: Hollenbeck Draw, 5 mi. S. of Belfry, 22

June 2011, S. Kohler, Paratype <3 (d). Fig. 218. Same (v). Fig. 219. FE. r. montanensis, Montana: Carbon Co.: Hollenbeck

Draw, 4200’, 5 mi. SE of Belfry, 11 June 2009, S. Kohler, Paratype ? (d). Fig. 220. Same (v). Fig. 221. Same, 19 June 2009,

S. Kohler, Paratype ? (d). Fig. 222. Same (v). Fig. 223. E. r. montanensis, Montana: Carbon Co.: Hollenbeck Draw, 5 mi. S.

of Belfry, 22 June 2011, S. Kohler, Paratype ? (d). Fig. 224. Same (v). Fig. 225. Same, Paratype ? (d). Fig. 226. Same (v).

Fig. 227. E. r. montanensis, Montana: Carbon Co.: 2-3 mi. SW Belfry, 21 June 2011, S, Kohler 3 (d). Fig. 228. Same (v).

Fig. 229. E. r. montanensis, Montana: Carbon Co.: 2-3 mi. SW Belfry, 4199-4325’, 17 June 2009, S. Kohler 3 (d). Fig. 230.

Same (v). Fig. 231. Same ? (d). Fig. 232. Same (v). Fig. 233. E. r. montaenesis, Montana: Carbon Co.: 2-3 mi. SW Belfry,

4240-4350’, 19 June 2009, S. Kohler 2 (d). Fig. 234. Same (v). Figs. 1.6X life size and to scale. Photos by S. Kohler.

47

236 aM

238 ; 239

241 242

Figs. 235-243. Male genitalia of Euphilotes showing shape of left valva in left lateral view. Fig. 235. E. battoides battoides,

California: Inyo Co.: Mono Pass Trail, Mosquito Flats to above Ruby Lake, 10300-12000’, 11 July 1989, J. & F. Preston. Fig.

236. E. bernardino bernardino, California: Los Angeles Co.: CR N-4, 1 mi. E. Bob’s Gap Rd., 27 June 1990, B. O’Hara. Fig.

237. E. b. martini, Nevada: Clark Co.: Newberry Mountains, Christmas Tree Pass, 22 April 1980, G.T. Austin. Fig. 238. £.

glaucon glaucon, Nevada: Douglas Co.: US 395, 2.1 mi. S. Carson City line, 30 May 1980, G.T. Austin. Fig. 239. EF. g.

glaucon, Oregon: Deschutes Co.: Rd. 16, 1-2 mi. S. Sisters, 7 June 2003, A.D. Warren. Fig. 240. £. g. glaucon, Oregon:

Klamath Co.: Hwy. 66, 1.5 mi. W. jct. Hwy. 97, SW edge Klamath Falls, 4300’, 18 June 2003, A.D. Warren. Fig. 241. E. g.

glaucon, Montana: Beaverhead Co.: near Lemhi Pass, 20 July 1982, S. Kohler. Fig. 242. E. g. glaucon, Montana: Ravalli Co.:

Railroad Cr., off Skalkaho-Rye Rd., 25 June 2001, S. Kohler. Fig. 243. FE. g. glaucon, Montana: Silver Bow Co.: Barrel Spring

trail, 4-5 mi. NE of Melrose, 5 June 2006, S. Kohler. Figs. approximately 50X life size and to scale. Photos by S. Kohler.

48

251 DEPP

250

Figs. 244-252. Male genitalia of Euphilotes showing shape of left valva in left lateral view. Fig. 244. £. glaucon glaucon,

Montana: Silver Bow Co.: Camp Creek Road, 4.5 mi. E. of Melrose, 4 June 2004, S. Kohler. Fig. 245. E. g. glaucon,

Montana: Silver Bow Co.: Camp Creek Road, 4-8 mi. E. of Melrose, 14 June 2004, S. Kohler. Fig. 246. E. g. oregonensis,

Oregon: Klamath Co.: Hwy. 97, vic. mile 224-226, ca. 4700’, 21 June 2000, A.D. Warren. Fig. 247. FE. glaucon (N. Oregon-

Washington Cascades segregate), Oregon: Clackamas/Hood River Cos.: Mt. Hood, NW of Timberline Lodge, 5700-6000’, 20

June 2003, A.D. Warren. Fig. 248. E. glaucon (N. Oregon-Washington Cascades segregate), Washington: Yakima Co.: Bethel

Ridge, Microwave Tower Rd., 18 July 2009, D. Nunnallee. Fig. 249. EF. glaucon near intermedia, Oregon: Josephine Co.:

NFD 4609, at ridge E. of Little Gray Jack Peak, 4500’, ca. 10 air miles E. Cave Junction, 11 July 2003, A.D. Warren. Fig. 250.

E. heracleoides, Holotype, Montana: Sanders Co.: W. of Niarada, 3 June 2004, S. Kohler. Fig. 251. E. heracleoides, Paratype,

same locality. Fig. 252. E. heracleoides, Paratype, Montana: Sanders Co.: 1.5 mi. W. of Niarada, 9 June 1997, S. Kohler.

Figs. approximately 50X life size and to scale. Photos by S. Kohler.

49

256 3 257 258

261

259

Figs. 253-261. Male genitalia of Euphilotes showing shape of left valva in left lateral view. Fig. 253. E. heracleoides,

Montana: Flathead Co.: 1.5 mi. N. of Niarada, 13 June 1996, S. Kohler. Fig. 254. FE. heracleoides, Montana: Lake Co.: 3 mi.

E. Hot Springs, 3600’, 20 June 1996, S. Kohler. Fig. 255. E. heracleoides, Montana: Lincoln Co.: Dancing Prairie, 5 mi. NW

Eureka, 25 June 1997, S. Kohler. Fig. 256. E. heracleoides, Idaho: Boise Co.: FR 380, from summit of Mores Pass, 7110’, 14

July 2005, T.W. Ortenburger. Fig. 257. FE. heracleoides, Oregon: Wallowa Co.: Blue Mts., Powwatka Ridge, ca. 11 mi. N. of

Wallowa on Troy Rd., 4100’, 23 June 2002, A.D. Warren. Fig. 258. FE. heracleoides, same locality. Fig. 259. E. heracleoides,

same locality. Fig. 260. E. heracleoides, Oregon: Crook Co.: Ochoco Mts., creek N. of Hwy. 26, 2.7 mi. SW ject. Little Hay Cr.

Rd. (NFD 2610), 4500’, 10 June 2003, A.D. Warren. Fig. 261. E. baueri baueri, California: Inyo Co.: W. side Gilbert Pass, 6

mi. NE Deep Springs, 19 May 2005, S. Kohler. Figs. approximately 50X life size and to scale. Photos by S. Kohler.

50

262 263

265

267

268 269 270

Figs. 262-270. Male genitalia of Euphilotes showing shape of left valva in left laterial view. Fig. 262. E. baueri baueri,

California: Inyo Co.: W. side Gilbert Pass, 6 mi. NE Deep Springs, 19 May 2005, S. Kohler. Fig. 263. EF. baueri orientis,

Nevada: Lincoln Co.: Bristol Wells Road, 3.5 mi. W. Hwy. 93, 20 May 2005, S. Kohler. Fig. 264. E. baueri (far N. Great

Basin northward segregate), Oregon: Crook Co.: Hwy. 126, 0.5-1.5 mi. E. Deschutes Co. line, 3050’, 8 June 2003, A.D.

Warren. Fig. 265. E. baueri (far N. Great Basin northward segregate), Oregon: Harney Co.: ca. 1 mi. E. Folly Farm Rd., 32

mi. S. ject. Hwy. 78, 4200’, 16 May 2002, A.D. Warren. Fig. 266. E. baueri (far N. Great Basin northward segregate), same

locality. Fig. 267. E. baueri borealis, Holotype, Montana: Sanders Co.: 4.5 mi. N. of Camas Prairie, 5 May 2004, S. Kohler.

Fig. 268. E. baueri borealis, Paratype, Montana: Sanders Co.: 5 mi. N. Camas Prairie, 20 May 1997, S. Kohler. Fig. 269. E.

baueri borealis, Montana: Madison Co.: Melrose Bench Road, 5400’, 15 mi. SE of Melrose, 6 June 2017, S. Kohler. _ Fig.

270. E. baueri shoshone, Holotype, Idaho: Butte Co.: NE jct. Craters Loop Rd. & Tree Molds Rd., 5875’, Craters of the Moon

Nat. Mon., 1 June 2017, S. Kohler. Figs. approximately 50X life size and to scale. Photos by S. Kohler.

51

271 272 273

275

ATG 278 2719

Figs. 271-279. Male genitalia of Euphilotes showing shape of left valva in left lateral view. Fig. 271. E. baueri shoshone,

Paratype, Idaho: Butte Co.: NE jct. Craters Loop Rd. & Tree Molds Rd., 5875’, Craters of the Moon Nat. Mon., 1 June 2017,

S. Kohler. Fig. 272. E. centralis centralis, Colorado: El Paso Co.: 2 mi. N. exit for Hwy. 16 on I-25, 30 July 1990, A.D.

Warren. Fig. 273. E. oakleyi oakleyi, Holotype, Montana: Missoula Co.: Spring Gulch, Rattlesnake Mountains, 18 June 2003,

S. Kohler. Fig. 274. E. 0. oakleyi, Paratype, Montana: Missoula Co.: Stuart Peak Trail, Rattlesnake Mountains, 18 June 2003,

S. Kohler. Fig. 275. E. 0. oakleyi, Paratype, Montana: Missoula Co.: Spring Gulch, Rattlesnake Mountains, 25 June 2002, W.

Kerling. Fig. 276. E. o. oakleyi, Montana: Missoula Co.: Miller Creek, 1 July 1975, S. Kohler. Fig. 277. E. o. oakleyi,

Montana: Ravalli Co.: Bass Cr., Florence, 13 June 1982, S. Kohler. Fig. 278. E. 0. oakleyi, same locality. Fig. 279. E. o.

oakleyi, Montana: Ravalli Co.: Bass Creek, Florence, 17 June 1980, S. Kohler. Figs. approximately 50X life size and to scale.

Photos by S. Kohler.

52

280 281 282

283 284 285

288

286 287

Figs. 280-288. Male genitalia of Euphilotes showing shape of left valva in left lateral view. Fig. 280. E. oakley oakleyi,

Montana: Ravalli Co.: Bass Creek, Florence, 3 July 1976, S. Kohler. Fig. 281. E. 0. oakleyi, Montana: Ravalli Co.: 8-Mile

Creek, E. of Florence, 26 June 1982, S. Kohler. Fig. 282. E. 0. oakleyi, Montana: Silver Bow Co.: Rocky Ridge Trailhead,

Divide Cr., 3 mi. W. Feely, 22 June 2001, S. Kohler. Fig. 283. E. oakleyi madisonensis, Holotype, Montana: Madison Co.:

Varney Road, 5345’, 4 mi. NW of Cameron, 27 June 2013, S. Kohler. Fig. 284. E. 0. madisonensis, Paratype, same locality.

Fig. 285. E. o. madisonensis, Paratype, Montana: Madison Co.: Varney Road, 4 mi. NW of Cameron, 7 July 2008, S. Kohler.

Fig. 286. E. o. madisonensis, Montana: Madison Co.: Gravelly Range Rd., N. Nat. Forest boundary, 7300’, 30 June 2015, S.

Kohler. Fig. 287. E. 0. madisonensis, Montana: Silver Bow Co.: Soap Gulch, 6980’, 7 mi. up, NE Melrose, 15 Jun 2015, S.

Kohler. Fig. 288. E. 0. madisonensis, Montana: Silver Bow Co.: Soap Gulch, 6850-7150’, 7-8 mi. up, NE Melrose, 17 June

2015, S. Kohler. Figs. approximately 50X life size and to scale. Photos by S. Kohler.

53

295 296

Figs. 289-297. Male genitalia of Euphilotes showing shape of left valva in left lateral view. Fig. 289. E. oakleyi madisoni,

Montana: Silver Bow Co.: Soap Gulch, 6875-7300’, 6-7 mi. up, NE Melrose, 24 June 2015, S. Kohler. Fig. 290. E. o.

madisonensis, Montana: Silver Bow Co.: Soap Gulch, 6850-7300’, 6-7 mi. up, NE Melrose, 25 June 2015, S. Kohler. Fig. 291.

E. o. madisonensis, same locality. Fig. 292. E. ancilla ancilla, Utah: Cache Co.: Green Canyon, 14 July 1993, S. Kohler. Fig.

293. E. ancilla barnesi, Colorado: Jefferson Co.: Ralston Creek, 7500’, GGCSP East Gate, 22 June 2005, C. Slater. Fig. 294.

E. a. barnesi, Colorado: Gilpin Co.: Hwy. 7 at Smith Gulch, 29 June 1996, B.B. Brinkman. Fig. 295. E. a. ancilla, Colorado:

Grand Co.: USFS Rd. 133 (Co. Rd. 50), 2-4 mi. E. jct. Hwy 40, 8 mi. W. Hot Sulphur Springs, 25 June 1989, A.D. Warren.

Fig. 296. E. ancilla montosa, Holotype, Montana: Missoula Co.: Ninemile Prairie, near Greenough, 14 June 1979, S. Kohler.

Fig. 297. E. ancilla campestris, Holotype, Montana: Powder River Co.: .8 mi. W. Pumpkin Cr. Rd., 24 mi. W. Broadus, S.

Hwy. 212, 15 June 2007, S. Kohler. Figs. approximately 50X life size and to scale. Photos by S. Kohler.

54

Figs. 298-306. Male genitalia of Euphilotes showing shape of left valva in left lateral view. Fig. 298. E. ancilla campestris,

Paratype, Montana: Powder River Co.: .8 mi. W. Pumpkin Cr. Rd., 24 mi. W. Broadus, S. Hwy. 212, 15 June 2007, S. Kohler.

Fig. 299. E. a. campestris, Montana: Hill Co.: Bears Paw Mountains, S. of Havre, 3 June 1986, S. Kohler. Fig. 300. F. a.

campestris, Montana: Carter Co.: Forest Road 4035, Long Pines, 36 mi. SE Ekalaka, 14 June 2007, S. Kohler. Fig. 301. £. a.

campestris, Montana: Stillwater Co.: Lone Tree Road, 15 mi. N. Columbus, 12 June 2007, S. Kohler. 302. E. ancilla

gilvatunica, Nevada: Lander Co.: US 50 at Austin Summit, 7500’, nr. Austin, 20 June 2002, J. & F. Preston. Fig. 303. E.

enoptes enoptes, Oregon: Lake Co.: Warner Mts., Bullard Cyn. at E. end Lakeview, ca. 5100’, 18 June 2003, A.D. Warren.

Fig. 304. E. enoptes enoptes, same locality. Fig. 305. E. enoptes aridorum, Nevada: Douglas Co.: Montreal Canyon, 31 July

1983, J.B. Vernon. Fig. 306. FE. enoptes bayensis, California: Marin Co.: Tiburon, 5 June 1983, J.B. Vernon. Figs.

approximately 50X life size and to scale. Photos by S. Kohler.

55

307

310 311 312

313 315

Figs. 307-315. Male genitalia of Euphilotes showing shape of left valva in left lateral view. Fig. 307. E. enoptes tildeni,

California: Los Angeles Co.: Sawmill Ridge, 4000’, 1 mi. W. Sawmill Camp, 9 September 1981, G.A. Gorelick. Fig. 308. E.

columbiae, Oregon, Sherman Co.: Jones Cyn. off Deschutes Canyon, ca. 1000’, 2 May 2003, A.D. Warren. Fig. 309. E.

columbiae, Oregon: Jefferson Co.: W. end Lake Billy Chinook, vic. mouth of Spring Creek, 1700’, 19 May 2003, A.D.

Warren. Fig. 310. E. spaldingi spaldingi, Utah: Millard Co.: Scipio Pass, 6700’, 1 mi. S. I-15, ex. ova, em. 1 June 2000, J.

Wolfe. Fig. 311. EF. rita coloradensis, Colorado: El Paso Co.: Ramah Rd., Hwy. 94, E. Colorado Springs, 6100’, 6 August

1973, M.S. Fisher. Fig. 312. E. rita montanensis, Holotype, Montana: Carbon Co.: Hollenbeck Draw, 5 mi. SE of Belfry, 23

May 2007, S. Kohler. Fig. 313. E. r. montanensis, Paratype, Montana: Carbon Co.: Hollenbeck Draw, 5 mi. SE of Belfry, 5

June 2007, S. Kohler. Fig. 314. E. r. montanensis, Montana: Carbon Co.: ridge W. Cub Creek, 5-7 mi. SE of Belfry, 16 May

2007, S. Kohler. Fig. 315. E. pallescens pallescens, Utah: Juab Co.: Jerico Dunes, 1 mi. W. Hwy. 50, mile 14, 8 August 1984,

J. Johnson. Figs. approximately 50X life size and to scale. Photos by S. Kohler.

56

319

5K)

| 304 ~ Sag , 327

Figs. 316-327. Female genitalia of Euphilotes, ventral view showing shape of lodix. Fig. 316. E. battoides battoides,

California: Inyo Co.: Mono Pass Trail, Mosquito Flats to above Ruby Lake, 10300-12000’, 11 July 1989, J. & F. Preston. Fig.

317. E. bernardino bernardino, California: Los Angeles Co.: Los Angeles, 1 June 1966, S. Kohler. Fig. 318. EF. bernardino

martini, Nevada: Clark Co.: Newberry Mountains, Bridge Canyon at Nev. 163, 22 April 1980, G.T. Austin. Fig. 319. E£.

glaucon glaucon, Nevada: Washoe Co.: 3 mi. E. US 395 on Hwy. to Virginia City, 19 June 1980, J.B. Vernon. Fig.320. E. g.

glaucon, Montana: Ravalli Co.: Railroad Creek, off Skalkaho-Rye Road, 25 June 2001, S. Kohler. Fig. 321. E. g. glaucon,

same locality. Fig. 322. EF. glaucon (N. Oregon-Washington Cascades segregate), Oregon: Clackamas/Hood River Cos.: Mt.

Hood, NW of Timberline Lodge, 5700-6000’, 20 July 2003, A.D. Warren. Fig. 323. EF. glaucon (N. Oregon-Washington

Cascades segregate), Washington: Yakima Co.: Bethel Ridge, Microwave Tower Rd., 18 July 2009, R.M. Pyle. Fig. 324. E.

heracleoides, Allotype, Montana: Sanders Co.: 1.5 mi. N. of Niarada, 13 June 1996, S. Kohler. Fig. 325. E. heracleoides,

Paratype, Montana: Sanders Co.: 1.5 mi. W. of Niarada, 2 June 2000, S. Kohler. Fig. 326. E. heracleoides, Montana: Sanders

Co.: 4.5 mi. N. Camas Prairie, 9 June 1998, S. Kohler. Fig. 327. E. heracleoides, Montana: Lake Co.: 3 mi. E. Hot Springs,

3600’, 20 June 1996, S. Kohler. Figs. approximately 50X life size and to scale. Photos by S. Kohler.

ST

Figs. 328-339. Female genitalia of Bupnilotes ventral view showing shape of lodix. Fig. 328. E. heracleoides, Montana:

Lincoln Co.: Dancing Prairie, 3350’, NW of Eureka, 1 July 1996, S. Kohler. Fig. 329. FE. heracleoides Oregon: Wallowa Co.:

Blue Mts., Powwatka Ridge, ca. 1 mi. N of Wallowa on Troy Rd., 4100’, 23 June 2002, A.D. Warren. Fig. 330. E

heracleoides, same locality. Fig. 331. E. heracleoides, Oregon: Union Co.: Phillips Cyn., ca. 2 mi. W. of Elgin, ca. 3000’, 23

June 2002, A.D. Warren. Fig. 332. EF. baueri baueri, California: Inyo Co.: W. side Gilbert Pass, 6 mi. NE Deep Springs, 19

May 2005, S. Kohler. Fig. 333. FE. baueri orientis, Nevada, Lincoln Co.: Bristol Wells Road, 3.5 mi. W. Hwy. 93, 20 May

2005, S. Kohler. Fig. 334. E. baueri (far N. Great Basin northward segregate), Oregon: Crook Co.: Hwy. 126, 0.5-1.5 mi. E.

Deschutes Co. line, 3050’, 8 June 2003, A.D. Warren. Fig. 335. FE. baueri (far N. Great Basin northward segregate), Oregon:

Harney Co.: Alvord Desert, 3 mi. NE Andrews, ca. 4000’, 8 May 1990, T. Stoddard. Fig. 336. E. baueri borealis, Alllotype,

Montana: Sanders Co.: 4.5 mi. N. Camas Prairie, 6 May 2004, S. Kohler. Fig. 337. E. b. borealis, Paratype, same locality.

Fig. 338. E. b. borealis, Montana: Madison Co.: Melrose Bench Rd., 5400’, 15 mi. SE of Melrose, 6 June 2017, assoc. Er.

ovalifolium var. ovalifolium, S. Kohler. Fig. 339. E. baueri shoshone, Allotype, Idaho: Butte Co.: NE jct. Craters Loop Rd. &

Tree Molds Rd., 5875’, Craters of the Moon Nat. Mon., 1 June 2017, S. Kohler. Figs. approximately 50X life size and to scale.

Photos by S. Kohler.

58

wf ; oa 351 \

Figs. 340-351. Female genitalia of Euphilotes, ventral view showing shape of lodix. Fig. 340. FE. baueri shoshone, Paratype,

Idaho: Butte Co.: NE jct. Craters Loop Rd. & Tree Molds Rd., 5875’, Craters of the Moon Nat. Mon., 1 June 2017, S. Kohler.

Fig. 341. EF. oakleyi oakleyi, Allotype, Montana: Missoula Co.: Stuart Peak Trail, Rattlesnake Mountains, 18 June 2003, S.

Kohler. Fig. 342. E. o. oakleyi, Paratype, Montana: Missoula Co.: Stuart Peak Trail, Rattlesnake Mountains, 4 July 2006, S.

Kohler. Fig. 343. E. 0. oakleyi, Montana: Silver Bow Co.: Rocky Ridge Trailhead, 2.5 mi. W. of Feely, Divide Cr., 19 June

2003, S. Kohler. Fig. 344. E. oakleyi madisonensis, Allotype, Montana: Madison Co.: Varney Road, 4 mi. NW Cameron,

5345’, 27 June 2009, S. Kohler. Fig. 345. E. 0. madisonennsis, Paratype, Montana: Madison Co.: Varney Road, 4 mi. NW of

Cameron, 7 July 2008, S. Kohler. Fig. 346. FE. 0. madisonensis, Paratype, same locality. Fig. 347. E. 0. madisonensis,

Paratype, Montana: Madison Co.: Varney Road, 4 mi. NW Cameron, 30 June 2009, S. Kohler. Fig. 348. £. 0. madisonensis,

Paratype, same locality. Fig. 349. EL. 0. madisonensis, Paratype, same locality. Fig. 350. E. o. madisonensis, Paratype, same

locality. Fig. 351. E. o. madisonensis, Paratype, same locality. Figs. approximately 50X life size and to scale. Photos by S.

Kohler.

59

e?

rgd p -

Figs. 352-363. Female genitalia of Euphilotes, ventral view showing shape of lodix. Fig. 352. E. ancilla ancilla, Utah: Cache

Co.: Green Canyon, 11 July 1965, S. Kohler. Fig. 353. E. ancilla montosa, Paratype, Montana: Missoula Co.: Ninemile

Prairie, nr. Greenough, 12 June 1979, S. Kohler. Fig. 354. E. a. montosa, Paratype, same locality. Fig. 355. E. a. montosa,

Montana: Carbon Co.: Hollenbeck Draw, 5 mi. SE of Belfry, 5 June 2007, S. Kohler. Fig. 356. E. a. montosa, Montana:

Carbon Co.: 2-3 mi. SW Belfry, 19 June 2008, S. Kohler. Fig. 357. E. a. montosa, Montana: Deer Lodge Co.: nr. Storm Lake

Pass, Anaconda Range, 9200’, 15 July 2003, S. Kohler. Fig. 358. E. a. montosa, Montana: Glacier Co.: Mount Henry, near

Two Medicine, 22 July 2005, S. Kohler. Fig. 359. E. a. montosa, Montana: Granite Co.: Black Pine Road, 6 mi. NW

Phillipsburg, 5675’, 20 June 2001, S. Kohler. Fig. 360. EF. a. montosa, Montana: Jefferson Co.: Warm Springs Creek, E. of

Alhambra, 23 June 2005, S. Kohler. Fig. 361. E. a. montosa, Montana: Powell Co.: below Morrell Lookout, E. of Seeley Lake,

11 July 2000, S. Kohler. Fig. 362. £. ancilla campestris, Allotype, Montana: Powder River Co.: .8 mi. W. Pumpkin Cr. Rd.,

24 mi. W. Broadus, S. Hwy 212, 15 June 2007, S. Kohler. Fig. 363. E. a. campestris, Paratype, same locality. Figs.

approximately 50X life size and to scale. Photos by S. Kohler.

60

ann 4, ; + 4 tothe

“ afk . ie ee ae |

Figs. 364-375. Female genitalia of Euphilotes, ventral view showing shape of lodix. Fig. 364. E. ancilla campestris, Montana:

Stillwater Co.: Lone Tree Road, 15 mi. N. Columbus, 12 June 2007, S. Kohler. Fig. 365. E. ancilla campestris, Montana:

Toole Co.: N. of Cameron Lake, Middle Butte Sweet Grass Hills, 11 July 2002, S. Kohler (lateral view). Fig. 366. E. enoptes

enoptes, Oregon: Lake Co.; Warner Mts., Bullard Cyn. at end of Lakeview, ca. 5100’, 19 June 2003, A.D. Warren. Fig. 367.

E. enoptes tildeni, California: Los Angeles Co.: Sawmill Ridge, 4000’, 1 mi. W. Sawmill Camp, 9 September 1981, G.A.

Gorelick. Fig. 368. E. columbiae, Washington: Skamania Co.: Hwy. 14 at SE base of Dog Mtn., 0.25 mi. W. Dog Creek Falls,

ca. 100’, 22 May 2003, A.D. Warren. Fig. 369. FE. columbiae, same locality. Fig. 370. E. spaldingi spaldingi, Utah: Salt Lake

Co.: E. Wasatch Blvd, SLC, 1 mi. N. Hwy. 190, ex. lar. 16 September 2006, em. 25 October 2006, J. Wolfe. Fig. 371. E. rita

coloradensis, Colorado: Saguache Co.: 4 mi. S. of Villa Grove, 22 July 1969, J. Scott. Fig. 372. E. rita montanensis, Allotype,

Montana: Carbon Co.: Hollenbeck Draw, 5 mi. SE of Belfry, 5 June 2007, S. Kohler. Fig. 373. E. r. montanensis, Montana:

Carbon Co.: 2-3 mi. SW Belfry, 19 June 2008, S. Kohler. Fig. 374. FE. r. montanensis, same locality. Fig. 375. E, pallescens

pallescens, Utah: Tooele Co.: nr. Willow Springs, 14 August 1971, K. Tidwell (lateral view). Figs. approximately 50X life

size and to scale. Photos by S. Kohler.

61

Figure 376. Upper. Habitat of Euphilotes baueri borealis at the type locality, 4.5 miles north of Camas Prairie, Sanders

County, Montana. Lower. Larval food plant of Euphilotes baueri borealis, Eriogonum ovalifolium var. pansum Reveal.

Photos by S. Kohler.

Se ee... . ih teed

Pee | sey ¥ % , om V ' fy Was rons, : : 1 weE ae

Figure. 377. Upper. Habitat of Euphilotes baueri shoshone at the type locality, vic. Crater Loop Road, Craters of the Moon

National Monument, Butte County, Idaho. Lower. Larval food plant of Euphilotes baueri shoshone, Eriogonum ovalifolium

var. focarium Reveal & Mansfield. Photos by S. Kohler.

Figure 378. Upper Left. Larval food plant of Euphilotes heracleoides, Eriogonum heracleoides var. heracleoides Nutt.,

Sanders County, Montana. Upper Right. Larval food plant of Euphilotes oakleyi oakleyi, Eriogonum umbellatum var. aureum

(Gand.) Reveal, at the type locality, Stuart Peak Trail, Rattlesnake Mountains, Missoula County, Montana. Lower. Habitat of

Euphilotes oakleyi oakleyi at the same locality. Photos by S. Kohler.

64

rl _

hall Aes

Le

sd

a : Z

a a

tig eee ;

Figure 379. Upper Left. Larval food plant of Euphilotes oakleyi madisonensis, Eriogonum ovalifolium var. ovalifolium

Nutt., Varney Road, 4 miles west of Cameron, Madison County, Montana. Upper Right. Local larval food plant of

Euphilotes ancilla montosa, Eriogonum ovalifolium var. depressum Blank, Red Mountain trail above Copper Camp, 7500’,

north of Lincoln, Lewis & Clark County, Montana. Lower. Habitat of Euphilotes ancilla montosa, at the same locality.

Photos by S. Kohler.

65

Figure. 380. Upper. Habitat of Euphilotes ancilla montosa, vicinity of Morrell Lookout, 10 miles east of Seeley Lake, Powell

County, Montana. Lower. Local larval food plant of Euphilotes ancilla montosa, Eriogonum flavum var. piperi (Greene) M.E.

Jones, from the same locality. Photos by S. Kohler

ot a : * a ' rf t . ma z c 3 i z . -

Figure. 381. Upper. Habitat of Euphilotes ancilla campestris at the type locality, 24 miles west of Broadus, Powder River

County, Montana. Lower. Larval food plant of Euphilotes ancilla campestris, Eriogonum flavum var. flavum Nutt., from the

same locality. Photos by S. Kohler.

67

Me

Figure. 382. Upper. Habitat of Buphilowes rita montanensis at the type foceihy ce Horlgabeck Dawe 5 emilee SE of Belfry,

Carbon County, Montana. Lower. Larval food plant of Euphilotes rita montanensis, Eriogonum pauciflorum Pursh., from the

same locality. Photos by S. Kohler.

68

Figure 383. Distribution of Euphilotes glaucon (W. H. Edwards, 1871).

Wai

1 Wk eos. rat 4 ; '

Euphilotes glaucon (W.H. Edwards)

@ E. glaucon glaucon

© FE. glaucon oregonensis

© E. glaucon (N. Oregon-Washington Cascades segregate)

© E. glaucon australoglaucon

© FE. glaucon comstocki

© FE. glaucon intermedia

69

Figure 384. Distribution of Euphilotes heracleoides Kohler & A. Warren, 2021

ere cnet,

Lie 9

*) 1

Euphilotes heracleoides Kohler & A. Warren

® FE. heracleoides

70

Figure 385. Distribution of Euphilotes baueri (Shields, 1975).

» weue

| i

3)

) “

ee = \

Euphilotes baueri (Shield

@ FE. baueri baueri

E. baueri orientis

E. baueri vernalis

E. baueri (far N. Great Basin northward segregate)

E. baueri borealis

E. baueri shoshone

0000 0

71

Figure 386. Distribution of Euphilotes oakleyi Kohler, 2021.

bil .

Euphilotes oakleyi Kohler

@ E. oakleyi oakleyi

® EF. oakleyi madisonensis

72

Figure 387. Distribution of Euphilotes ancilla (W. Barnes & McDunnough, 1918).

Euphilotes ancilla (W. Barnes & McDunnough)

@ EF. ancilla ancilla

ancilla barnesi

. ancilla montosa

. ancilla campestris

. ancilla (SE Oregon segregate)

ancilla giulianii

ancilla gilvatunica

. ancilla shieldsi

. ancilla purpura

. ancilla cryptica

. ancilla pseudointermedia

©®270® 000 OO

73

Figure 388. Distribution of Euphilotes rita (W. Barnes & McDunnough, 1916).

' 4 : ' on ‘A te i ” Sie ‘ , ‘a ,

' { y , a’ .

\ 7h : VA Oo -t = Li , Le ae

Pet ‘aay ‘dba SE A a i

1 - = $e oe 2 i

Euphilotes rita (W. Barnes & McDunnough)

® F. rita rita

® E. rita coloradensis

© FE. rita montanensis

@ E. rita (Four Corners Area segregate)

74

Figure 389. Ecological Provinces of Montana (after Werner et al., 2004).

ACKNOWLEDGMENTS

Appreciation is given to Jonathan P. Pelham and Richard L. Romeyn for their encouragement and

review of this paper and for providing specimens and distribution records. Peter Lesica has identified

many Eriogonum host plants over the years, and also reviewed this paper. Many people have provided

specimens, collection records, information and other assistance. They include, among others: T. Aneiros,

George T. Austin, B. Boyd, Barton B. Brinkman, John Christenson, Vernon Covlin, Chris Durden, John

F. Emmel, David Ewer, Nancy Ewer, Clifford D. Ferris, Michael S. Fisher, G. A. Gorlick, Lionel P. Grey,

Megan Hansen, Chuck E. Harp, Frank E. Holley, J. Johnson, Kim Kendall, Will Kerling, Norbert G.

Kondla, John Lane, Gary Lawson, Glen Marangelo, David Nunnallee, Jo Nunnallee, James Oberfoell, B.

A. O’Hara, Paul A. Opler, Tom W. Ortenburger, W. D. Patterson, Adam Peters, Jeffrey S. Pippen,

Gordon F. Pratt, Floyd Preston, June Preston, Robert M. Pyle, Dana Ross, Ronald A. Royer, James A.

Scott, L. M. Scott, Charles Slater, Ray E. Stanford, Terry Stoddard, Kenneth Tidwell, Dave Trochlell,

John B. Vernon, Bea Vogel, Byron Weber and Jacque Wolfe.

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