Volume 8 Number 8 30 December 2020

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

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

Review of the Speyeria egleis complex in Montana,

with the description of two new subspecies

(Lepidoptera: Nymphalidae: Heliconiinae)

Steve Kohler

2617 Glen Drive

Missoula, MT 59804

ABSTRACT: Four phenotypically and geographically distinct sets of populations of Speyeria egleis are identified in

Montana. Two new subspecies are named from northwestern and northcentral Montana. Two existing names are recognized

(S. e. macdunnoughi from southwestern Montana, southeastern Idaho and northwestern Wyoming, and S. e. albrighti from

central Montana. S. e. near macdunnoughi from Oregon and Washington is discussed.

Additional key words: Island ranges, Sweet Grass Hills, Morrell Mountain, Highwood Mountains.

ZooBank registration: urn:lsid:zoobank.org:pub:94D912CC-72BA-42EA-A 14E-E0OF51E1CD498

INTRODUCTION

At the present time, thirteen described subspecies of Speyeria egleis (Behr, 1862) are recognized

in North America (Pelham 2020). Of these, two occur in Montana, S. e. macdunnoughi (Gunder, 1932)

and S. e albrighti (Gunder, 1932). Historically, all Montana egleis populations with a brownish colored

under hindwing disc have been assigned to macdunnoughi and all populations with an olive-green colored

under hindwing disc have been assigned to albrighti (Ferris and Brown 1981; Kohler 1980). During the

course of my study of Montana butterflies, I have identified at least four distinct sets of populations which

can be phenotypically and geographically segregated. The purpose of this paper is to define the

characters of these four distinctive populations and to describe two new subspecies.

DISCUSSION

In this study, the two current subspecies of S. egleis are recognized. Two additional subspecies,

one from northwestern Montana and another from island mountain ranges in northcentral Montana are

designated below. Each of the subspecies is discussed and defined, and material examined for this study

is listed. Forewing length measurements (from the junction with the thorax to the wing apex) are given in

millimeters. The specimen figures are 0.8 life size and to scale.

Speyeria egleis macdunnoughi (Gunder, 1932)

(Figs.1-20)

Argynnis macdunnoughi Gunder, 1932, Can. Entomol. 64(12): 280-281.

Argynnis macdunnoughi Gund.; McDunnough, 1938, Mem. S. Calif. Acad. Sci. 1: 15.

Speyeria (Speyeria) egleis macdunnoughi (Gunder); dos Passos and Grey, 1947, Amer. Mus.

Novit. (1370): 5, 18; dos Passos, 1964, Mem. Lepid. Soc. (1): 94.

Speyeria egleis macdunnoughi (Gunder); Miller and Brown, 1981, Mem. Lepid. Soc. (2): 142;

Pelham, 2008, J. Res. Lepid. 40: 318; Pelham, 2020, www. butterfliesofamerica.com/US-Can-Cat.htm.

The type locality of macdunnoughi, as stated by Gunder is “Elkhorn Ranch Resort, Gallatin

County, Mont.” He also added a note, which read, “For the convenience of future students who may wish

to know the definite type locality of macdunnoughi, I quote from a letter received from Dr. McDunnough,

‘If you go up what they call (at the Resort) Spring Creek for about half a mile, the path branches to the

right about where the water ceases to flow and rises sharply over a ridge, ending in an open meadow. If

you cross this meadow to the top of the slope, you will find Argynnis plentiful in the open woods to the

right. Slightly earlier than my date might be a better collecting time.’” The additional data, “7000 feet,

Upper Gallatin Canyon” was added by dos Passos and Grey (1947).

Gunder named macdunnoughi for Dr. James H. McDunnough, who collected the holotype male,

allotype female and two male paratypes, and placed the specimens in the Canadian National Collection at

Ottawa. Dr. McDunnough is among some of the best-known names in Canadian butterfly study. He was

a tireless collector and systematist in the first half of the 19" century who built up the Canadian National

Collection considerably (Gunder, 1932; Layberry, Hall and Lafontaine, 1998). The material examined

by Gunder (1932) and upon which he based the description of macdunnoughi consisted of the four

Canadian National Collection specimens (holotype, allotype and two male paratypes) all collected by Dr.

McDunnough at the type locality Aug. 1, 1928, and an additional two males and one female paratypes in

Gunder’s collection, “together with a series of males and females taken by Mr. Hutchins, Mr. Wind and

Dr. Albright of Great Falls, Montana, in the same general region, including the Teton Range South in

Wyoming” (Gunder, 1932).

The original description was brief, and not much definitive information can be extracted from it.

Gunder said, “in size and superficial appearance near to oweni Edw. from the Mt. Shasta, Calif. but on the

upper sides obviously related to the chitone Edw. group, though dwarfed in size in comparison. Oweni

has an almost uniform dark ground color above, while macdunnoughi has a lighter aspect through the

limbal areas. Also in macdunnoughi the yellow band on the under side secondaries is more pronounced

or definite. Macdunnoughi also is of a drab greyish appearance in some specimens on the under side.

(One paratype in Canadian coll. is unsilvered .. .)”. The taxon chitone is now a subspecies of hesperis.

This subspecies is characterized by the usually medium-brown color of the discal area of the

under hindwing. The lighter aspect of the dorsal ground color in the limbal (post-median) wing areas is

present in both sexes, but is more extensive in females. It becomes more obvious in flight-worn

individuals. The under hindwing discal spots of macdunnoughi may be silvered or un-silvered (Figs. 13-

18) (Table 1). In the material examined for this study, approximately 25% of specimens were un-

silvered.

The distribution of macdunnoughi in Montana is shown in Fig. 127. It extends south into

northwestern Wyoming and has been recorded in the following counties: Big Horn, Fremont, Hot Springs,

Johnson, Park, Sheridan, Teton and Washakie. The distribution also includes parts of southern Idaho.

Probable records for that region include the following counties: Ada, Adams, Bannock, Bear Lake, Boise,

Bonneville, Caribou, Clearwater, Elmore, Franklin, Fremont, Idaho, Jefferson, Latah, Lemhi, Madison,

Power, Teton, Valley, and Washington (Pelham, pers. com). Both Dornfeld (1980) and Hinchliff (1994)

reported macdunnoughi from northeastern Oregon. However, Warren (2005) said, “until geographic

variation of S.egleis in eastern Oregon and Idaho can be studied in more detail, populations throughout the

Ochoco, Aldrich, Blue and Wallowa mountains are herein called Speyeria egleis nr. macdunnoughi.” He

listed its occurrence in the following counties: Baker, Crook, Grant, Harney, Umatilla and Union. A

Grant Co. specimen is shown in Figs. 21-22. Pelham (2020, pers. com.) also cited nr. macdunnoughi

records for Columbia and Garfield counties in southeastern Washington. Throughout its range,

macdunnoughi_ flies in a single brood from late June through August, depending on elevation and

progression of the season. Peak flight is normally from early to mid-July.

James and Nunalee (2011) gave the life history details for S$. egleis nr. macdunnoughi. They

reared it twice from gravid females collected near Mt. Misery, Garfield Co., Washington. They listed the

recorded host plants of egleis as Viola nuttallii (Nuttall’s Violet), V. praemorsa (Prairie Violet), V.

adunca (Blue Violet), V. purpurea (Goosefoot Violet) and V. walteri (Prostrate Blue Violet). Of these,

nuttallii, praemorsa and adunca are common and widespread in Montana and could be the presumed host

plants for our egleis. No life history work has been done for any of the Montana populations. Typical

macdunnoughi adults are shown in Figs. 1-20. For this study 52 males and 12 females were examined.

Average forewing length of males was 26.7 mm, with a range of 25.0 to 29.0 mm. Average forewing

length of females was 28.7 mm, with a range of 27.5 to 30.0 mm. Average forewing length of the series

of 8 males of near macdunnoughi from Grant Co. Oregon was 27.8 mm, with a range of 27.0 to 30.5 mm.

Material Studied: MONTANA: Beaverhead Co.: Baily Hill, Jackson, 3 July 1942, 16, H. A. Howland Coll.; Polaris, 16

July 1943, 1d, 22 July 1943, 14, 16 July 1944, 93, 18 July 1944, 66, H. A. Howland Coll.; 7 mi. NW Bannock Pass, 22 July

1986, 13, S. Kohler Coll.; Broadwater Co.: Avalanche Gulch, 6.5 mi. up, 8 July 2005, 13, S Kohler Coll.; Carbon Co.: vic.

MT-WY state line, 8100’, Main Fork Rock Cr., SW Red Lodge, 23 July 2019, 13, S. Kohler Coll.; Gallatin Co.: Jackson Cr.

Trailhead, Bridger Mts., NE Bozeman, 23 June 2014, 14, S. Kohler Coll.; below Sacajawea Pass, 8550’, Bridger Mts., 25 July

2013, 13, S Kohler Collector; Sacajawea Peak, Bridger Mts., 19 July 2007, 13, S. Kohler Coll.; above Fairy Lake, 7600’,

Bridger Mts., 19 July 2007, 22, 24 July 2013, 23, S. Kohler Coll.; Jefferson Co.: Boulder River Rd., SW Basin, 24 August

1993, 13, S. Kohler Coll.; Lewis & Clark Co.: Deadman Coulee Rd., 4200’, N. Wolf Creek, 25 August 2011, 12, S. Kohler

Coll.; Madison Co.: Gravelly Range Rd., 7290’, at N. USFS boundary, 27 June 2013, 14 12, S. Kohler Coll.; S. slope Lone

Mtn., Madison Range, 2 August 2006, 1¢ 22, S. Kohler Coll.; Ravalli Co.: near Skalkaho Pass, 21 August 1981, 19, S.

Kohler Coll.; Sanders Co.: Baldy Mountain, 6800’, NE Plains, 24 July 1996, 136, S. Kohler Coll.; Stillwater Co.: Benbow

Mine Rd., 9000’, 17 mi. SW Fishtail, 18 July 1989, 13, B. Vogel Coll.; Benbow Mine Rd., 10000’, Stillwater Plateau, near

Dean, 8 August 2013, 1d, S. Kohler Coll.; Emerald Lake, West Rosebud Canyon, 10 July 1997, 13, S. Kohler Coll.; Mystic

Lake Rd., 15 August 1986, 22, S. Kohler Coll.; Sweet Grass Co.: Picket Pin Rd., 8500’, W. Nye, 4 July 2013, 1 2, S. Kohler

Coll.; Picket Pin Mtn., 10000’, W. of Nye, 17 July 2014, 14, S. Kohler Coll.; Picket Pin Rd., 9000-9500’, East Boulder

Plateau, 17-19 mi. W. of Nye, 25 July 2019, S. Kohler Coll.; vic. Picket Pin Mtn., 8950-9500’, W. Nye, 31 July 2013, 16, S.

Kohler Coll.; S. slope Picket Pin Mtn., 9000-9700’, W. of Nye, 31 July 2015, 12, S. Kohler Coll; WYOMING: Lincoln Co.:

Swift Cr., near Afton, 28 June 1972, 23, 28 June 1973, 44, 29 June 1973, 23, L. P. Grey Coll.; Swift Cr., Afton, 27 June

1972, 23, 1 August 1973, 23 29, D. Eff Coll.; OREGON: Grant Co.: Aldrich Mtn. Lookout Rd., 7.1-7.5 mi. E. jct. Fields

Creek Rd., Aldrich Mts., 16 July 2016, 83, R. L. Romeyn Coll.

Speyeria egleis albrighti (Gunder, 1932)

(Figs. 23-44)

Argynnis albrighti Gunder, 1932, Can. Entomol. 64(12): 281-282.

Argynnis albrighti Gund.; McDunnough, 1938, Mem. S. Calif. Acad. Sci. 1: 15.

Speyeria (Speyeria) egleis albrighti (Gunder); dos Passos and Grey, 1947, Amer. Mus. Novit.

(1370): 5, 18; dos Passos, 1964, Mem. Lepid. Soc. (1): 94.

Speyeria egleis albrighti (Gunder); Miller and Brown, 1981, Mem. Lepid. Soc. (2): 142; Pelham,

2008, J. Res. Lepid. 40: 318; Pelham, 2020, www. butterfliesofamerica.com/US-Can-Cat.htm.

The type locality of albrighti as stated by Gunder is, “Highwood Mts., Chouteau County,

Montana”. The type series in Gunder’s collection (now in LACM) consisted of only three specimens, the

holotype male and allotype female, both collected June 24, 1931 at the type locality, and one paratype

male collected July 26, 1931 at Monarch, Cascade County, Montana. Monarch is in the Little Belt Mts.

south of the Highwood Mts. All three specimens were collected by C. C. Albright, for whom Gunder

named this subspecies. Clifton Clarence Albright was born in Shelby, New York September 16, 1877 and

moved to Montana in 1905. He died in 1946 at the age of 68. Albright, who practiced medicine in

Augusta, Anaconda and Great Falls, Montana, evidently maintained a serious interest in botany and

entomology. His substantial butterfly collection was donated to Montana State University, Bozeman

following his retirement (Great Falls Tribune April 8, 1946, p. 6). Some years ago, C. C. Albright’s

granddaughter, Charlotte Albright (pers. corresp.), provided some additional information. “I don’t know

when he started collecting, but he was certainly active in the 1920’s and 1930’s, and traded extensively

with other collectors. He was an eye, ear, nose and throat doctor. By the time my father, Joe, was a

teenager my grandfather was already suffering from Parkinson’s disease, so that my father and uncle did

most of the collecting—the usual technique was to ride on the running board of their car and jump off

after likely specimens. They would go to the Highwood and Little Belt Mountains on weekends”.

Again, the original description was brief, with focus on the heaviness of the upper side maculation

and the shade of green on the under hindwing. Gunder said, “Nearest to platina Skin. from Utah and

Idaho. The black designs of the upper sides are extra heavy and the light spots marking the position of the

under side silver spotting, is very lightened in contrast, like those on upper side gallatini McD. In platina

the green cast or shadow design is very delicate, an olive shade, while in albrighti this green is dark,

heavier and the brown in conjunction is more pronounced. In female albrighti this greenish cast of design

is even more developed”.

Since Gunder’s description, very few specimens of albrighti had been obtained from the

Highwood Mountains. Most specimens collected in Montana that were referred to albrighti had been

collected in the Crazy Mountains, Sweet Grass County or the Little Belt Mountains in Cascade and

Meagher Counties. Moeck (1957), in commenting on albrighti noted the peculiar suffused, sordid green

disk, making it a very attractive and distinctive insect, but said that in the Rocky Mountains they had

found the various egleis races not too easy to locate, and that they prefer fairly high altitudes. Howe

(1975) observed that egleis are largely confined to rather high, cool and moist elevations, and that

albrighti inhabits a remote area seldom visited by lepidopterists, and is a poorly known subspecies of

rather uncertain status.

For my own part, I had determined that in order to better define which populations of Montana

egleis truly represented the concept of albrighti, that I would need a long study series from the Highwood

Mountains. Unfortunately, prior to 2011, I had only been able to obtain five examples, mainly from

riparian areas at the lower elevations. In 2011, and subsequent years, hikes to the upper parts of

Highwood Baldy Mountain and the higher ridges east of Highwood Creek produced much better results,

and larger series were obtained.

This subspecies is larger in size than macdunnoughi, and on the upper side there is more basal

suffusion of dark scales and the overall black maculation is heavier. The discal area of the under

hindwing of albrighti is most often overlaid with a diffused shade of dull brownish green. The spots in

the discal area of the hindwing are always silvered (Figs. 23-44) (Table 1).

The distribution of albrighti is limited to southcentral Montana (Fig. 127). Reports of albrighti

from the western slopes of the Big Horn Mountains in Wyoming (Ferris & Brown, 1981) are most likely

referable to macdunnoughi. For this study 219 males and 22 females were examined. Average forewing

length of males was 27.0 mm, with a range of 23.0 to 31.5 mm. Average forewing length of females was

29.3 mm, with a range of 27.0 to 32.5 mm. It flies in a single brood from late June through August,

depending on elevation and progression of the season. Peak flight at the type locality is normally from

early to mid-July.

Material Studied: MONTANA: Cascade Co.: Monarch, 29 May 1934, 36, 15 June 1934, 16, 24 June 1934, 1d, H. E.

Nelson Coll.; near Kings Hill, 20 August 1935, 12, C. C. Albright Coll.; Chouteau Co.: Highwood Mts., 14 July 1999, 22, S.

Kohler Coll.; Highwood Cr., Highwood Mts., 15 July 2010, 36, S. Kohler Coll.; Highwood Baldy, 7540’, Highwood Mts., 28

July 2011, 1436 19; vic. Prospect Peak, 5880’, Highwood Mts., 27 July 2011, 406 19, S. Kohler Coll.; near jet. trails 412 &

413, 5850’, Highwood Mts., 11 July 2013, 26a 29; jct. White Wolf & Marie Spring trails, 5875’, Highwood Mts., 10 July

2014, 38d 22, S. Kohler Coll.; jct. White Wolf & Marie Spring trails, 5850’, Highwood Mts., 2 July 2015, 143, S. Kohler

Coll.; near jct. White Wolf & Marie Spring trails, 5800-5900’, Highwood Mts., 13 July 2020, 14a 22, S. Kohler Coll.;

Golden Valley Co.: Little Snowy Mts., 14 May 1978, 36, S. Kohler Coll.; Judith Basin Co.: Yogo Peak, 8800’, Little Belt

Mts., 31 July 2020, 44, S. Kohler Coll.; Meagher Co.: Kings Hill, Little Belt Mts., 20 August 1935, 12, 29 August 1935, 12,

C. C. Albright Coll.; Loco Mountain & trail 636, 7585’, Crazy Mts., 4 August 2011, 3346 62, S. Kohler Coll.; Sweet Grass

Co.: Crazy Mts., 23 July 1971, 4a, L. P. Grey Coll.; Swamp Creek Rd., Crazy Mts., 23 July 1975, 43, 24 July 1975, 73 12,

20 July 1976, 13, 12 July 1978, 4a 12, S. Kohler Coll.; Big Timber Canyon, Crazy Mts., 19 July 1973, 3¢ 12; vic. Twin

Lakes, Crazy Mts., 31 July 2007, 1°, S. Kohler Coll.

Speyeria egleis morrellensis — new subspecies

(Figs. 45-92)

ZooBank registration: urn:|sid:zoobank.org:act:C79016E0-6B3 1-40D2-84EE-C184D 1287232

During the course of this study, it became obvious that the egleis populations inhabiting the higher

mountains of the northwestern portion of Montana were distinct from those of other areas of the state.

Definition: The most striking characteristic of morrellensis dorsally is the extremely dark, dusky

appearance. In both males and females, the orange ground color of the wings of many individuals is

largely obscured by basal suffusion of black scales. It is the smallest of any of the Montana subspecies

with an average forewing length of 25.1 mm for males, with a range of 21.0 to 28.5 mm, and an average

of 27.4 mm for females, with a range of 25.0 to 30.5 mm. For this study, a total of 388 males and 76

females were examined (Figs. 45-92) (Table 1).

Etymology: This subspecies is named for the type locality, Morrell Mountain. The name “Morrell” has

been tied to the peak, lake, stream and falls, and a prairie in the Seeley Lake area. Their namesake,

Charles Thomas Morrell was a pioneer in Territorial Montana and one of the area’s early homesteaders.

He owned a gun and sporting goods store in Helena and manufactured the first gun in Montana in 1889,

just before it became a state. Morrell was the youngest of five children and grew up in the state of New

York. He followed his sister to Helena after she wrote to her brother telling him it would be a fine place

for him to start a business. In 1894 he moved out into the unsettled area northwest of Helena now known

as Seeley Lake. He homesteaded the area until 1910 when he moved back to Helena. It is likely that his

homestead at the mouth of Morrell Creek was responsible for the creek and surrounding landmarks

eventually bearing his name. The Jocko Indian Trail came into the Seeley Lake area from Placid Lake to

the southwest, and the Summit Indian Trail followed Morrell Creek going northward (Barber and Rieman,

2011).

Distribution and Phenology: This subspecies is found only in the higher elevations of the Rocky

Mountains in northwestern Montana (Fig. 127). It flies in a single brood from late June through August,

depending on elevation and progression of the season. Peak flight at the type locality is normally from

early to mid-July.

Types: Holotype male: MONTANA: Powell Co.: vic. Morrell Lookout, 7600’, 10 mi. E. Seeley Lake, 9

July 2009, S. Kohler Coll. Alltoype female: MONTANA: Powell Co., below Morrell Lookout, east of

Seeley Lake, 11 July 2000, S. Kohler Coll. Paratypes (369 males, 67 females): MONTANA: Powell

Co.: Morrell Lookout, E. Seeley Lake, 15 July 1992, 13, S. Kohler Coll.; below Morrell Lookout, east of

Seeley Lake, 16 July 1977, 553 112, 11 July 2000, 453 152, S. Kohler Coll.; below Morrell Lookout,

7450’, E. of Seeley Lake, 17 July 2001, 213 32, 20 July 2001, 36a 72, S. Kohler Coll.; N. of Morrell

Lookout, E. of Seeley Lake, 9 August 2002, 63 32, S. Kohler Coll.; vic. Morrell Lookout, 7600’, 10 mi.

E. Seeley Lake, 12 July 2004, 143 12, 10 July 2007, 160, 9 July 2009, 100 12,9 July 2014,93 29,4

July 2015, 23 12,22 July 2016, 23, S. Kohler Coll.; vic. Morrell Lookout, 7300-7700’, 10 mi. E. Seeley

Lake, 7 July 2009, 193, S. Kohler, Coll.; vic. Morrell Lookout, 10 mi. E. Seeley Lake, 23 July 2011, 35 3

22,S. Kohler Coll.; vic. Morrell Lookout, 7400’, 10 mi. E. Seeley Lake, 15 July 2013, 743 152, 26 July

2013, 163 42, S. Kohler Coll.; vic. Morrell Lookout, 7700’, 10 mi. E. Seeley Lake, 29 June 2016, 3d, S.

Kohler Coll.; Morrell Lookout Rd., 7500’, 10 mi. E. Seeley Lake, 12 July 2016, 53 22,8. Kohler Coll.

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, Gainsville, Florida. The

remaining paratypes are in the author’s collection.

Type Locality: Montana: Powell County: vicinity of Morrell Lookout, 7300-7700’, 10 miles east of

Seeley Lake. The lookout is located on the ridge leading up to the summit of Morrell Mountain.

Additional Material Studied: MONTANA: Glacier Co.: near Two Medicine, 3 August 1978, 13, S. Kohler Coll.; Spot

Mountain, near Two Medicine, 28 July 1998, 43, S. Kohler Coll.; Mount Henry, 8500’ near Two Medicine, 2 August 2001,

33 12,8. Kohler Coll.; Mount Henry, near Two Medicine, 22 July 2005, 13, S. Kohler Coll.; Lewis & Clark Co.: Scapegoat

Mountain, 9200’, Scapegoat Wilderness, 22 July 2009, 73, S. Kohler Coll.; Flint Mountain, 8000’, Scapegoat Wilderness, 23

July 2009, 83; above Copper Camp, 8235’, 12 mi. N. of Lincoln, 4 August 2009, 3 2, S. Kohler Coll.; Red Mountain, 9000-

9400’, 13 mi. N. of Lincoln, 4 August 2009, 126 22, S. Kohler Coll.; SW slope Red Mountain, 7700-8660’, NE Lincoln, 21

July 2015, 15d 12, S. Kohler Coll.; Missoula Co.: Cottonwood Lakes Rd., 10 mi E. Seeley Lake, 21 July 1978, 13 12,S.

Kohler Coll.; near Necklace Lakes, Swan Range, 19 August 2000, 13, W. Kerling Coll.; vic. Holland Lookout trail 42, 2

August 2002, 43, S. Kohler Coll.; Teton Co.: Mount Wright, 38 mi. NW Choteau, 29 July 2005, 16, 11 July 2007, 12, S.

Kohler Coll.; Mount Wright, 8000’, 40 mi. NW Choteau, 9 August 2014, 116 19, S. Kohler Coll.

Speyeria egleis kutoyisiks — new subspecies

(Figs. 93-126)

ZooBank registration: urn:|sid:zoobank.org:act:8A673343-B55C-4DFE-AE7E-662764D9 1DC5

The “Island Ranges” of central Montana rise up from the surrounding prairies in contrast to the

main mass of the Rocky Mountains to the west where the mountains dominate the intervening valleys.

Montana’s island ranges include the Sweet Grass Hills, Bears Paw Mts., Little Rocky Mts., Highwood

Mts., Judith and Moccasin Mts., Little Snowy Mts., Big Snowy Mts., Little Belt Mts., Big Belt Mts.,

Castle Mts., and Crazy Mts. Because of their separation from the Rockies and from each other, the

possibility exists that distinct populations of species of butterflies are present in these island ranges that

that differ substantially. Subspecies albrighti is restricted to a group of island ranges including the

Highwood, Judith, Little Snowy, Big Snowy, Little Belt, Big Belt and Crazy Mts. Of all the island

ranges, the most isolated are the Sweet Grass Hills, Bears Paw Mts. and the Little Rocky Mts. It comes as

no surprise then, that another distinct subspecies of egleis would be present in this group of island ranges.

Norbert Kondla, my neighbor to the north in Canada first brought this population to my attention in 1989.

Definition: This is the largest of the Montana subspecies, with forewing length of males averaging 29.1

mm with a range of 21.0 to 32.0 mm, and females averaging 31.5 mm, with a range of 28.0 to 35.0 mm.

Dorsally, the wings have the least amount of basal suffusion of black scales, and the maculation is the

least heavy, allowing a greater amount of the orange ground color to show. The ground color of the discal

area of the under hindwing in many male individuals is a warm golden buff, overlaid with only a slightest

dull greenish brown sheen, while others display this sheen of variable intensity. Females have more

brownish tone to this sheen, but overall the ventral hindwing has a light appearance. The dorsal wing

surface has a noticeable two-tone appearance, where the orange ground color in the post-median wing

areas 1s a lighter shade. This character is present in both sexes, but is more pronounced in the females.

(Figs. 93-126) (Table 1).

Etomology: Kutoyisiks is the Blackfeet Indian word for the Sweet Grass Hills, and means “sweet pine

hills. The Sweet Grass Hills possess special significance to the Blackfeet Indians and to other tribes on

the northern Great Plains. According to legend, the creator Napi fashioned the hills in the dim past out of

the rocks left over from the formation of the Rocky Mountains. Napi liked his creation so much that the

hills became a favored resting place for the old trickster. Located in the heart of a fertile bison hunting

ground, the hills served as a vantage point for game and as a lookout for enemies trespassing in Blackfeet

territory. Because of the isolation and connection with the creation of the earth, they have deep cultural

significance to the Blackfeet as a spiritual refuge where teenage boys made vision quests to help guide

them into adulthood. Many of the Blackfeet’s traditional stories take place in and around the hills. One

of those involves the Blackfeet culture hero Kutoyis (Blood-Clot Boy) who sought to rid the world of evil

in the early history of the world. It was in the Sweet Grass Hills that Kutoyis defeated Lizard or Frog

Man after an epic battle that lasted many days. The Blackfeet paid tribute to his memory by naming the

hills “Kutoyisiks” in his honor (Montana Department of Transportation Historical Highway Marker).

Distribution and Phenology: This subspecies is found only in the Sweet Grass Hills, Bears Paw Mts.

and Little Rocky Mts. of northcentral Montana (Fig. 127). It flies in a single brood from late June

through July, depending on progression of the season. Peak flight at the type locality is normally from

early to mid-July.

Types: Holotype male: MONTANA: Toole Co.: Pratt Canyon, 4470-5000’, West Butte Sweet Grass

Hills, 1 July 2019, S Kohler Coll. Allotype female: MONTANA: Toole Co.: Pratt Canyon, 4460-4800’,

West Butte Sweet Grass Hills, 16 July 2019, S. Kohler Coll. Paratypes (249 males, 86 females):

MONTANA: Toole Co.: Pratt Canyon, 4705’, West Butte, Sweet Grass Hills, 9 July 2001, 13, S. Kohler

Coll.; Pratt Canyon, West Butte, Sweet Grass Hills, 10 July 2001, 23 22, 24 July 2002, 1630 32, 22 July

2003, 236 32, S. Kohler Coll.; Pratt Canyon, 4800’, West Butte, Sweet Grass Hills, 29 July 2004, 13, 12

July 2007, 103, 12 July 2007, 103, 24 June 2009, 43, 25 June 2009, 23, S. Kohler Coll.; Pratt Canyon,

4470-5000’, West Butte, Sweet Grass Hills, 26 June 2019, 46, 1 July 2019, 51d 22, S. Kohler Coll;

Pratt Canyon, 4430-4800’, West Butte, Sweet Grass Hills, 9 July 2019, 793 42, S. Kohler Coll.; Pratt

Canyon, 4460-4800’, West Butte, Sweet Grass Hills, 12 July 2019, 403 23 2, 16 July 2019, 263 422, 19

July 2019, 23 82,S. Kohler Coll.

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, Gainsville, Florida. The remaining

paratypes are in the author’s collection.

Type Locality: Montana: Toole County: Pratt Canyon, 4430-5000’, West Butte, Sweet Grass Hills.

The Sweet Grass Hills in the northern parts of Toole and Liberty counties near the Alberta-Montana

border in northcentral Montana are unique in that they are the highest isolated peaks in the United States.

Of volcanic origin, the Sweet Grass Hills are prominent landmarks, rising nearly 3000’ above the

surrounding prairie with rolling hills extending to the north almost to the Alberta-Montana border. They

are visible for more than 50 miles and consist of three distinct butte complexes with scattered grassy hills

connecting them. The three buttes are West Butte (elevation 6983’), Middle or Gold Butte (elevation

6512’) and East Butte (elevation 6958’). The stream that flows in Pratt Canyon is Simmons Creek. The

high peaks of the Sweet Grass Hills rise abruptly from the surrounding prairie and support montane plant

communities, including coniferous forests, which have their closest counterparts in the Cordilleran Rocky

Mountains over 80 miles to the west (Thompson and Kuijt, 1976). These authors conducted the first

detailed study of the montane and subalpine flora of this isolated mountain range. They found Viola

adunca and V. canadensis in the riparian forests along Simmons Creek, extending well into the montane

region, and V. canadensis in the aspen grovelands. Open groves of Populus tremuloides surrounded by

grasslands are found on grassy south-facing slopes as high as 1800m. Both of these violets could be

considered possible host plants for S. e. kutoyisiks.

Additional Material Studied: MONTANA: Blaine Co.: N. Fork Little Peoples Cr., 3800-4500’, Little Rocky Mts., SE

Hays, 26 June 2018, 3¢, 3 July 2018, 1d, 4 July 2018, 13, 6 July 2018, 1d, 10 July 2019, 13, 17 July 2019, 13, 23 June

2020, 53, 25 June 2020, 36, 15 July 2020, 16, S. Kohler Coll.; N. ridge N. Fork Little Peoples Cr., Little Rocky Mts., SE

Hays, 5 July 2018, 46 12,8. Kohler Coll.; hill W. jct. CR 303 & 304, 4300’, 4.3 mi. S. Lloyd, Bears Paw Mts., 11 July 2019,

23, S. Kohler Coll.; Chouteau Co.: Sandy Creek Rd., 4300-4800’, SE Rocky Boy, Bears Paw Mts., 24 June 2020, 12, S.

Kohler Coll.; Hill Co.: Miners Gulch, 4350-4850', E. Beaver Creek Rd., Bears Paw Mts., S. Havre, 14 July 2020, 52, S.

Kohler Coll.; Miners Gulch, 4340-4900’, E. Beaver Creek Rd., Bears Paw Mts., S. Havre, 21 July 2020, 14, 22 July 2020, 22,

S. Kohler Coll.; Liberty Co.: Breed Creek, East Butte, Sweet Grass Hills, 5 August 2003, 12, 28 July 2005, 12, S. Kohler

Coll.; Mount Royal, East Butte, Sweet Grass Hills, 30 July 2004, 1¢ 22, S. Kohler Coll.; Breed Creek, 4720-4800’, East

Butte, Sweet Grass Hills, 6-7 mi. SE Whitlash, 15 July 2009, 23, S. Kohler Coll.; Phillips Co.: Beaver Creek Rd., Little

Rocky Mts., NE Zortman, 4 July 2018, 14, S. Kohler Coll.; Toole Co.: West Butte, Sweet Grass Hills, 12 July 1980, 1d 22,

N. G. Kondla Coll.; Limekiln Creek, West Butte, Sweet Grass Hills, 10 July 2002, 13, S. Kohler Coll.

There is the possibility that either one or both of the new subspecies of egleis will be found in

Canada, due the close proximity to the border, of some of the Montana records. Layberry, Hall and

Lafontaine (1998), commented on egleis in Canada. “This species has been reported several times from

southwestern Alberta and southwestern British Columbia (Guppy et al., 1994). All of these records have

proved to be misidentifications. The Great Basin Fritillary does occur close to the Canadian Border in

Montana and may still be discovered in southern Alberta”.

Figs. 1-18. S. e. macdunnoughi. Fig. 1. Gallatin Co., MT, 3 (d). Fig. 2. Same, (v). Fig. 3. Same, (d). Fig. 4. Same,

(v). Fig. 5. Gallatin Co., MT, @ (d). Fig. 6. Same, (v). Fig. 7. Madison Co., MT,? (d). Fig. 8. Same, (v). Fig. 9.

Sweet Grass Co., MT, & (d). Fig. 10. Same, (v). Fig. 11. Sweet Grass Co., MT, 2 (d). Fig. 12. Same, (v). Fig. 13.

Beaverhead Co., MT, & (d). Fig. 14. Same, (v). Fig. 15. Same, (d). Fig. 16. Same, (v). Fig. 17. Same, (d). Fig. 18.

Same, (v). Figs. 0.8 life size and to scale. Photos S. Kohler.

Figs. 19-20. S. e. macdunnoughi. Fig. 19. Lincoln Co., WY, & (d). Fig. 20. Same, (v). Figs. 21-22. S. e. near

macdunnoughi. Fig. 21. Grant Co., OR, 3 (d). Fig. 22. Same, (v). Figs. 23-36. S. e. albrighti. Fig. 23. Chouteau

Co., MT,& (d). Fig. 24. Same, (v). Fig. 25. Same, (d). Fig. 26. Same, (v). Fig. 27. Same, (d). Fig. 28. Same, (v).

Fig. 29. Chouteau Co., MT, 2? (d). Fig. 30. Same, (v). Fig. 31. Same, (d). Fig. 32. Same, (v). Fig. 33. Same, (d).

Fig. 34. Same, (v). Fig. 35. Same, (d). Fig. 36. Same, (v). Figs. 0.8 life size and to scale. Photos S. Kohler.

Figs. 37-44. S. e. albrighti. Fig. 37. Chouteau Co., MT, 2 (d). Fig. 38. Same, (v). Fig. 39. Sweet Grass Co., & (d).

Fig. 40. Same, (v). Fig. 41. Meagher Co., MT, & (d). Fig. 42. Same, (v). Fig. 43. Meagher Co., MT, (d). Fig. 44.

Same, (v). Figs. 45-54. S. e. morrellensis. Fig. 45. Powell Co., MT, holotype 3, (d). Fig. 46. Same, (v). Fig. 47.

Powell Co., MT, paratype & (d). Fig. 48. Same (v). Fig. 49. Same, (d). Fig. 50. Same, (v). Fig. 51. Same, (d). Fig.

52. Same, (v). Fig. 53. Same, (d). Fig. 54. Same, (v). Figs. 0.8 life size and to scale. Photos S. Kohler.

Figs. 55-72. S. e. morrellensis. Fig. 55. Powell Co., MT, paratype 3 (d). Fig. 56. Same, (v). Fig. 57. Same, (d).

Fig. 58. Same, (v). Fig. 59. Same, (d). Fig. 60. Same, (v). Fig. 61. Same, (d). Fig. 62. Same, (v). Fig. 63. Same,

(d). Fig. 64. Same, (v). Fig. 65. Powell Co., MT, allotype 2 (d). Fig. 66. Same, (v). Fig. 67. Powell Co., MT.,

paratype 2(d). Fig. 68. Same, (v). Fig. 69. Same, (d). Fig. 70. Same, (v). Fig. 71. Same, (d). Fig. 72. Same, (v).

Figs. 0.8 life size and to scale. Photos S. Kohler.

Figs. 73-90. S. e. morrellensis. Fig. 73. Powell Co., MT, paratype 2 (d). Fig. 74. Same, (v). Fig. 75. Same, (d). Fig.

76. Same, (v). Fig. 77. Same, (d). Fig. 78. Same, (v). Fig. 79. Same, (d). Fig. 80. Same, (v). Fig.81. Same, (d). Fig.

82. Same, (v). Fig. 83. Glacier Co., MT, 2 (d). Fig. 84. Same, (v). Fig. 85. Teton Co., MT, & (d). Fig. 86. Same,

(v). Fig. 87. Teton Co., 2 (d). Fig. 88. Same, (v). Fig. 89. Lewis & Clark Co., 3 (d). Fig. 90. Same, (v). Figs. 0.8

life size and to scale. Photos S. Kohler.

Figs. 91-92. S. e. morrellensis. Fig. 91. Lewis & Clark Co., MT, 2 (d). Fig. 92. Same, (v). Figs. 93-108. S. e.

kutoyisiks. Fig. 93. Toole Co., MT, holotype & (d). Fig. 94. Same, (v). Fig. 95. Toole Co., MT, paratype 3 (d).

Fig. 96. Same, (v). Fig. 97. Same, (d). Fig. 98. Same, (v). Fig. 99. Same, (d). Fig. 100. Same, (v). Fig. 101. Same,

(d). Fig. 102. Same, (v). Fig. 103. Same, (d). Fig. 104. Same, (v). Fig. 105. Same, (d). Fig. 106. Same, (v). Fig.

107. Same, (d). Fig. 108. Same, (v). Figs. 0.8 life size and to scale. Photos S. Kohler.

4 ws

A how

. 111. Toole Co,

Same, (v). Fig

114. Same, (v). Fig. 115. Same, (d). Fig

allotype ? (d). Fig. 110.

’

Figs. 109-126. S. e. kutoyisiks. Fig. 109. Toole Co., MT

. 116.

Same, (v). Fig. 121. Blaine Co.,

113. Same, (d). Fig.

Same, (d). Fig. 118. Same, (v). Fig. 119. Same, (d). Fig. 120

Same, (v). Fig

MT, paratype 2 (d). Fig. 112.

(v). Fig. 117.

Same,

MT,

Hill Co, MT, & (d).

(v). Fig. 125

. Same,

Blaine Co., MT 2 (d). Fig. 124

. 126. Same, (v). Figs. 0.8 life size and to scale. Photos S. Kohler.

Same, (v). Fig. 123.

122

3 (d). Fig.

Fig

DATA DETAILS FOR SPECIMEN FIGURES

Speyeria egleis macdunnoughi. Figs. 1-2. MONTANA: Gallatin Co.: Jackson Cr. trailhead, Bridger Mts., NE Bozeman, 23

June 2014, S. Kohler Coll. Figs. 3-4. Same, above Fairy Lake, 7600’, Bridger Mts., 24 July 2013, S. Kohler Coll. Figs. 5-6.

Same, above Fairy Lake, Bridger Mts., 19 July 2007, S. Kohler Coll. Figs. 7-8. Madison Co.: S. slope Lone Mountain,

Madison Range, 2 August 2006, S. Kohler Coll. Figs. 9-10. Sweet Grass Co.: Picket Pin Mountain, 10000’, west of Nye, 17

July 2014, S. Kohler Coll. Figs. 11-12. Same, S. slope Picket Pin Mountain, 9000-9700’, W. of Nye, 31 July 2015, S. Kohler

Coll. Figs. 13-14. Beaverhead Co.: Baily Hill, Jackson, 3 July 1942, H. A. Howland Coll. Figs. 15-16. Same, Polaris, 18

July 1944, H. A. Howland Coll. Figs. 17-18. Same, 16 July 1944, H. A. Howland Coll. Figs. 19-20. WYOMING: Lincoln

Co.: Swift Cr., nr. Afton, 29 June 1973, L. P. Grey Coll.

Speyeria egleis near macdunnoughi. Figs. 21-22. OREGON: Grant Co.: Aldrich Mtn. Lookout, Aldrich Mts., Rd. 7, 1-7.5

mi. E. jct. Fields Cr. Rd., 16 July 2016, R. L. Romeyn Coll.

Speyeria egleis albrighti. Figs. 23-24. MONTANA: Chouteau Co.: jct. White Wolf & Marie Spring trails, 5875’, Highwood

Mts., 10 July 2014, S. Kohler Coll. Figs. 25-26. Same, Highwood Baldy, 7540’, Highwood Mts., 28 July 2011, S. Kohler

Coll. Figs. 27-28. Same, jct. White Wolf & Marie Spring trails, 5875’, Highwood Mts., 10 July 2014, S. Kohler Coll. Figs.

29-30. Same, nr. jct. trails 412 & 413, 5850’, Highwood Mts., 11 July 2013, S. Kohler Coll. Figs. 31-32. Same, vic. Prospect

Peak, 5880’, Highwood Mts., 27 July 2011, S. Kohler Coll. Figs. 33-34. Same, Highwood Baldy, 7540’, Highwood Mts., 28

July 2011, S. Kohler Coll. Figs. 35-36. Same, jct. White Wolf & Marie Spring trails, 5875’, Highwood Mts., 10 July 2014, S.

Kohler Coll. Figs 37-38. Same, nr. jct. White Wolf & Marie Spring trails, 5800-5900’, Highwood Mts., 13 July 2020, S.

Kohler Coll. Figs. 39-40. Sweet Grass Co.: Swamp Cr. Rd., Crazy Mts., 12 July 1978, S. Kohler Coll. Figs. 41-42.

Meagher Co.: Loco Mountain & trail 636, 7585’, Crazy Mts., 4 August 2011, S. Kohler Coll. Figs. 43-44. Same.

Speyeria egleis morrellensis. Figs. 45-46. MONTANA: Powell Co.: vic. Morrell Lookout, 7600’, 10 mi. E. Seeley Lake, 9

July 2009, S. Kohler Coll. Figs. 47-48. Same, vic. Morrell Lookout, 7400’, 10 mi. E. Seeley Lake, 15 July 2013, S. Kohler

Coll. Figs. 49-50. Same, vic. Morrell Lookout, 7400’, 10 mi. E. Seeley Lake, 26 July 2013, S. Kohler Coll. Figs. 51-52.

Same, vic. Morrell Lookout, 10 mi. E. Seeley Lake, 23 July 2011, S. Kohler Coll. Figs. 53-54. Same, below Morrell Lookout,

7450’, E. of Seeley Lake, 20 July 2001, S. Kohler Coll. Figs. 55-56. Same. Figs. 57-58. Same, vic. Morrell Lookout, 10 mi.

E. Seeley Lake, 23 July 2011, S. Kohler Coll. Figs. 59-60. Same, vic. Morrell Lookout, 7300-7700’, 10 mi. E. Seeley lake, 7

July 2009, S. Kohler Coll. Figs. 61-62. Same, vic. Morrell Lookout, 10 mi. E. Seeley Lake, 23 July 2011, S. Kohler Coll.

Figs. 63-64. Same, below Morrell Lookout, east of Seeley Lake, 16 July 1977, S. Kohler Coll. Figs. 65-66. Same, 11 July

2000, S. Kohler Coll. Figs. 67-68. Same, below Morrell Lookout, 7450’, E. of Seeley Lake, 17 July 2001, S. Kohler Coll.

Figs. 69-70. Same, vic. Morrell Lookout, 10 mi. E. Seeley Lake, 23 July 2011, S. Kohler Coll. Figs. 71-72. Same, below

Morrell Lookout, 7450’, E. of Seeley Lake, 20 July 2001, S. Kohler Coll. Figs. 73-74. Same, below Morrell Lookout, east of

Seeley Lake, 16 July 1977, S. Kohler Coll. Figs. 75-76. Same. Figs 77-78. Same, below Morrell Lookout, east of Seeley

Lake, 11 July 2000, S. Kohler Coll. Figs. 79-80. Same, vic. Morrell Lookout, 7400’, 10 mi. E. Seeley Lake, 15 July 2013, S

Kohler Coll. Figs. 81-82. Same, below Morrell Lookout, east of Seeley Lake, 16 July 1997, S Kohler Coll. Figs. 83-84.

Glacier Co.: Mount Henry, 8500’, near Two Medicine, 2 August, 2001, S. Kohler Coll. Figs. 85-86. Teton Co.: Mount

Wright, 38 mi. NW Choteau, 11 July 2007, S. Kohler Coll. Figs. 87-88. Same, Mt. Wright, 8000’, 40 mi. NW Choteau, 6

August 2014, S. Kohler Coll. Figs. 89-90. Lewis & Clark Co.: SW slope Red Mtn., 7700-8660’, NE Lincoln, 21 July 2015,

S. Kohler Coll. Figs. 91-92. Same.

Speyeria egleis kutoyisiks. Figs. 93-94. MONTANA: Toole Co.: Pratt Canyon, 4470-5000’, West Butte, Sweet Grass Hills, 1

July 2019, S. Kohler Coll. Figs. 95-96. Same, Pratt Canyon, 4470-5000’, West Butte, Sweet Grass Hills, 26 June 2019, S.

Kohler Coll. Figs. 97-98. Same. Figs. 99-100. Same, Pratt Canyon, 4470-5000’, West Butte, Sweet Grass Hills, 1 July 2019,

S. Kohler Coll. Figs. 101-102. Same, Pratt Canyon, 4470-5000’, West Butte, Sweet Grass Hills, 26 June 2019, S. Kohler

Coll. Figs. 103-104. Same, Pratt Canyon, 4430-4800’, West Butte, Sweet Grass Hills, 9 July 2019, S. Kohler Coll. Figs.

105-106. Same, Pratt Canyon, 4470-5000’, West Butte, Sweet Grass Hills, 1 July 2019, S. Kohler Coll. Figs. 107-108. Same,

Pratt Canyon, 4430-4800’, West Butte, Sweet Grass Hills, 9 July 2019, S Kohler Coll. Figs. 109-110. Same, Pratt Canyon,

4460-4800’, West Butte, Sweet Grass Hills, 16 July 2019, S. Kohler Coll. Figs. 111-112. Same. Figs. 113-114. Same, Pratt

Canyon, 4460-4800’, West Butte, Sweet Grass Hills, 12 July 2019, S. Kohler Coll. Figs. 115-116. Same, Pratt Canyon, 4470-

5000’, West Butte, Sweet Grass Hills, 1 July 2019, S. Kohler Coll. Figs. 117-118. Same, Pratt Canyon, 4460-4800’, West

Butte, Sweet Grass Hills, 12 July 2019, S. Kohler Coll. Figs. 119-120. Same, Pratt Canyon, 4430-4800’, West Butte, Sweet

Grass Hills, 9 July 2019, S. Kohler Coll. Figs. 121-122. Blaine Co.: N. ridge North Fork Little Peoples Cr., 4500-5000’,

Little Rocky Mts., SE Hays, 5 July 2018, S. Kohler Coll. Figs. 123-124. Same. Figs. 125-126. Hill Co.: Miners Gulch,

4350-4850’, E. Beaver Cr. Rd., Bears Paw Mts., S. Havre, 14 July 2020, S. Kohler Coll.

Fig. 127. Distribution of Speyeria egleis subspecies in Montana.

Speyeria egleis (Behr)

@ S. e. morrellensis

@ S. e. kutoyisiks

Table 1. Characteristics of Montana Speyeria egleis subspecies

Subspecies FW Length Ventral HW

| males | females | maculation | basal suffusion submarginal band

macdunnoughi | 26.7 28.7 light to light to moderate medium brown silvered or yellowish buff,

moderate unsilvered | caps on median and

submarginal spots

medium brown

— moderate moderate dull brownish green, always yellowish buff,

browner in females silvered | caps on median and

submarginal spots

greenish brown

27.4 darker dull brownish always dull yellowish buff,

green, browner in silvered | caps on median and

females submarginal spots

darker greenish

brown

27.

warm golden buff with always golden buff, caps

slight to moderate silvered on median and

greenish brown sheen, submarginal spots

browner in females lighter greenish

brown

ACKNOWLEDGMENTS

Appreciation is given to Norbert G. Kondla for his review of this paper and for providing

specimens. I am grateful to Jonathan P. Pelham for information on the distributions of the populations

involved and for encouragement and review on this project. Thanks is also given to Charlotte Albright

for information about her grandfather, and to Harry Pavulaan for help in organizing and formatting this

paper.

LITERATURE CITED

Barber, K. and B. Rieman. 2011. Morrell Creek—An important place with links to the past and future.

Clearwater Resource Council Report, Dec. 30, 2011, Seeley Lake, Montana, 2 pp.

Behr, H. 1862. Our California Argynnides (II). Proceedings California Academy of Natural Sciences

2:174.

Dornfeld, E. J. 1980. The Butterflies of Oregon. Timber Press, Forest Grove, Oregon, 276 pp.

dos Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalocera. The Lepidopterists’ Society

Memoir (1): 145 pp.

dos Passos, C. F. and L. P. Grey. 1947. Systematic catalogue of Speyeria (Lepidoptera, Nymphalidae)

with designation of types and fixations of type localities. American Museum Novitates (1370): 30 pp.

Ferris, C. D. and F. M. Brown. 1981. Butterflies of the Rocky Mountain States. University of

Oklahoma Press, Norman, Oklahoma, 442 pp.

Gunder, J. D. 1932. New Rhopalocera (Lepidoptera). Canadian Entomologist 64(12): 276-284.

Hinchliff, J. 1994. An Atlas of Oregon Butterflies. The Distribution of the Butterflies of Oregon. The

Evergreen Aurelians, The Oregon State University Bookstore, Inc., Corvallis, Oregon, 176 pp.

Howe, W.H. 1975. The butterflies of North America. Garden City, New York, Doubleday & Co., 633

Pp.

James, D. G. and D. Nunnallee. 2011. Life Histories of Cascadia Butterflies. Oregon State University

Press, Corvallis, Oregon, 447 pp.

Kohler, S. 1980. Checklist of Montana Butterflies (Rhopalocera). Journal of the Lepidopterists’ Society

34(1): 1-19.

Layberry, R. A., J. D. Hall and J. D. Lafontaine. 1998. The Butterflies of Canada. University of Toronto

Press, Toronto, Ontario, 280 pp.

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

1 Macrolepidoptera. Memoirs of the Southern California Academy of Science 1: 3-272.

Miller, L. D. and F. M. Brown. 1981. A catalogue/checklist of the butterflies of America north of

Mexico. The Lepidopterists’ Society Memoir (2): 280 pp.

Moeck, A. H. 1957. Geographic variability in Speyeria. Comments, records and description of a new

subspecies (Nymphalidae). Milwaukee; Milwaukee Ent. Soc.: 48 pp.

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

the Lepidoptera 40: 1-658.

Pelham, J. P. 2020. Catalogue of the Butterflies of the United States and Canada. Revised 7 Aug. 2020.

Accessed 19 Nov. 2020. http://www.butterfliesofamerica.com/US-Can-Cat.htm.

Thompson, L. S. and J. Kuijt. 1976. Montane and subalpine plants of the Sweetgrass Hills, Montana, and

their relation to early postglacial environments of the northern Great Plains. The Canadian Field-

Naturalist 90: 432-448.

Warren, A. D. 2005. Lepidoptera of North America 6: Butterflies of Oregon Their Taxonomy,

Distribution, and Biology. Contributions of the C. P. Gillette Museum of Arthropod Diversity. Colorado

State University, Fort Collins, Colorado, 408 pp.

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