BULLETIN OF THE ALLYN MUSEUM
THE ALLYN MUSEUM OF ENTOMOLOGY
Number 11 10 May 1973
REVISION OF THE BOLORIA EPITHORE
COMPLEX, WITH DESCRIPTION OF TWO NEW
Edwin M. Perkins, Jr.
Department of Biological Sciences, University of Southern California, Los Angeles, California 90007
W. Craig Meyer
Department of Geological Sciences, University of Southern California, Los Angeles, California 90007
Treated as a monotypic species for more than a century, Boloria epithore was
the object of an initial, somewhat cursory appraisal seven years ago (Perkins and
Perkins, 1966); thereafter, the senior author endeavored to compile additional data
in order to augment the limited and fragmentary knowledge then available. In
addition to the 254 samples employed in the original study, another 901 spread
specimens were acquired from British Columbia, Alberta, Washington, Idaho,
Montana, Oregon, and California.
The 1,155 individuals were then subjected to 1) a re-evaluation of more
complete and extensive distributional records, 2) a more critical evaluation of
phenotypic character analyses, 3) a statistical interpretation of linear measure-
ments, and their significance, and 4) a new dimension, not previously employed:
namely, an examination of the paleogeological mechanisms that possibly affected
the dispersion and geographic isolation of epithore's various conspecific
The purpose of this revision, therefore, is to present new data--and subsequent
interpretations--dealing with the epithore species group, and to integrate these with
previous knowledge (including historical background information and original
descriptions) by means of review, evaluation, summary, and rearrangement.
Boloria epithore epithore (Edwards)
Argynnis epithore Edwards, 1864, Proc. Ent. Soc. Phila., 2: 504.
The following is quoted from Edwards' original description:
"Argynnis epithore. Boisduval in litt .... Male. Expands 1 5/10 inch. Primaries rounded as in Myrina,
not angular at apex and excavated on the margin, as in Bellona, to which last it is most closely allied. Upperside
pale fulvous at the base; hind margins bordered by a slight, interrupted line, with small lunules; otherwise the
usual markings. Under side of primaries fulvous, yellowish at apex, with ferruginous sub-apical patch.
Secondaries have an angular submesial band of irregular spots, as in Bellona, each whitish, sprinkled in the
centre with ferruginous, in the cell a round black spot; beyond the band to the margin a slight violet tinge, with a
submarginal series of round spots and marginal lunules."
When the nominotypic subspecies, Boloria epithore epithore, was described
by W. H. Edwards in 1864, California was cited as the type locality; an exact locale
was not given. Hence, Brown (1965: p. 334) designated and figured a neotype of
Edwards' epithore, with the data: "Saratoga, Santa Clara Co., Calif., R. C. Winslow,
May 13, 1899."
It is debatable whether Edwards' type specimen was collected in the vicinity
of San Francisco. In Volume I of his Butterflies of North America ("Argynnis VI."),
the following statement was made by Edwards regarding "Argynnis callippe"
Boisduval: "From California. The most common or only species of Argynnis found
in the vicinity of San Francisco according to Dr. Behr...." Volume I was divided into
ten parts; each part had a different date of issuance. Part two of Volume I, which
contained "Argynnis callippe," was issued in August, 1868. This is four years after
Edwards' description of epithore was published. Surely, if Edwards' type specimen
of epithore, from Behr's collection (Brown, 1965: p. 337), had been collected in the
vicinity of San Francisco, Behr would not have made such a misleading statement.
Hence, support is lent to the theory that Edwards' epithore was collected in the Santa
Cruz Mountains, approximately 40 miles SSE of San Francisco.
The presently known range of typical epithore extends from southern San
Mateo County, south and west, through the Santa Cruz Mountains of Santa Cruz
and Santa Clara counties (Map 1). In recent years, typical nominate epithore has
not been encountered north or south of these limits. However, Williams (1910) stated
that epithore had been collected "a good many years ago" in Golden Gate Park, San
Francisco. Thus, there is reason to believe that it did occur northward, prior to the
concentrated urbanization of the San Francisco area.
Throughout its limited range, colonies of penins liar epithore are relatively
local; they are usually found in association with their fGod-plant, Viola ocellata
Torrey & Gray (Comstock, 1927).
The senior author has examined a series of 103 males and 35 females.
Pertinent statistical data is outlined in Tables 3 and 4, as are key phenotypic
characters in Tables 1 and 2.
Although Edwards' original description is essentially applicable to those
epithore that occur in the Santa Cruz Mountains, there exist two discrepancies. He
indicated that the submedian-median band of the ventral secondaries contains
"spots, as in Bellona, each whitish, sprinkled in the centre with ferruginous..." Of
138 examples inspected from the Santa Cruz Mountains, only one, weathered female
displays "whitish" spots. However, even these have a noticeably yellowish hue; the
remainder of the series possess yellowish-cream spots (Table 2). Perhaps Edwards'
connotation--"whitish"-was intended to imply an off-white color. Secondly, his
explicit comment regarding the "pale fulvous" color of the dorsal, basal regions is not
consistent with those examples used in this study, since all possess a black or
fuscous dusting on the hindwings, dorsally (usually extending outwardly as far as
the postbasal region); the forewings, dorsally, have only slight black, basal dusting
(Table 1). Perhaps the male specimen used by Edwards to describe epithore was an
extreme, which was generally very light in coloration. In size, Edwards' type (which
measured "1 5/10 inch" in expanse, or approximately 38 mm.) is smaller than the
average size of the 103 male specimens examined. Conversely, Brown (pers. comm.)
states that the designated neotype of epithore has a left forewing (LFW) measure-
ment of 24 mm. This figure is somewhat larger than e averAe measurement
derived by the senior author (Table 3).
TABLE 1. COMPARISON OF DORSAL CHARACTERISTICS OF FOUR RACES OF Boloria epithore1
Race epithore sierra chemocki borealis
Number of typical
specimens examined 1036' 35 8 2428' 105 39 260d' 217 9 118 Vd 75 9 e
Ground color bright orange bright orange yellowish-orange flat, dull orange
markings and spots sharply demarcated sharply demarcated sharply demarcated; diffuse;
black/brown-black black/brown-black black grey-black
Black basal suffusion
on FW and HP + + +++ +++/++++
Black dusting on veins
of FW and W + + +/1+- ++++
Black spots of FW and WS
median bands contiguous sFW /HWi + + +
W vein RS connects
laterally with bifed
figure in cell RARELY YES YES yes, if not obscured
Black suffusion on
marginal area of HW + -+ + +4+
Relative size of
submarginal spots on
FW and HW 4+ + 4+ +++
Based upon the examination of 1,155 spread specimens (not including material from zones of intergradation).
BOISDUVAL'S USE OF THE NAME EPITHORE
The introduction of the name epithore cannot be attributed to Edwards,
although he was the first to publish it. In his original description, he stated: "This
species, as I am informed by Dr. Behr, is undescribed and only named in letters of
Dr. Boisduval." The original description in which Boisduval used the name epithore
appeared in 1869 (Ann. Soc. Ent. Belg., 12: 58; no. 50).
In an attempt to determine the locality from which Boisduval's type of
epithore was taken, the senior author encountered certain contradictory information.
The lectotype of Boisduval's epithore (figured by Brown, 1965: p. 335) is in the
collection of the Carnegie Museum, Pittsburgh. Its superficial appearance is analo-
gous to female epithore from the Santa Cruz Mountains. The large size--25.0 mm.
LFW (Brown, pers. comm.)-is to be expected of typical epithore; however, the greatest
LFW measurement made by the senior author on 35 females was only 24.38 mm.
Table 3, therefore, indicates that the LFW radius of the lectotype of Boisduval's
epithore exceeds even the "99% Limits" (Brown, 1951) of the series used in this study.
The specimen figured by Brown as the 'Type' of Argynnis epithore
Boisduval" has an unconnected median row of black spots on the dorsal secondaries.
On the primaries these spots tend to be slightly fused or connected by transverse
black scales along the veins. This characteristic is common only to nominotypic
epithore (Table 1). Furthermore, Boisduval refers to the submedian-median area of
the ventral secondaries as being "jaune saupoudr&e de brun" (brown-powdered
yellow). This coloration is also characteristic of epithore from the Santa Cruz
Mountains (Table 2). However, in his original description, Boisduval stated:
"M. Lorquin a trouv6 cette espece dans les hautes montagnes de 1'est ou elle est fort
rare et difficile A prendre" (Mr. Lorquin found this species in the high mountains of
the east where it is extremely rare and difficult to capture). Although it would seem
that he clearly indicated that the material he had examined did come from an area
or areas in the mountains of eastern California, a statement in Volume III of
Edwards' Butterflies of North America strongly indicates that Boisduval's locality
statement may have been misleading.
From ("Argynnis VIII.") -- "Argynnis adiante" Boisduval, Edwards stated:
"The male figured on our Plate is the original type of Dr. Boisduval, sent me by
himself, and bearing his label as 'type adiante'." Edwards then quoted Boisduval:
"This beautiful Argynnis was taken in some numbers by M. Lorquin, on the edges
of the woods, in the eastern part of California." Edwards then continues: "Of late
years adiante has not been a very common species in collections, owing to its local
habits, apparently. Professor J. J. Rivers writes me that 'it is found above Los Gatos
in the Santa Cruz Mountains. It also occurs at several localities in the same range,
and in Santa Clara and San Mateo counties; but it does not appear to be found farther
south than about nine miles north of Santa Cruz city.' Apparently Dr. Bosiduval
was mistaken in the locality."Boisduval described adiante, now a synonym of
Speyeria egleis adiaste (Edwards), on page 61 of the same publication which
contained his original description of epithore. Both the adiante and epithore types
used by Boisduval in his original descriptions were collected by Lorquin. In both
original descriptions, Boisduval referred to eastern California and the high
mountains of the east as the localities from which the respective types were collected.
However, dos Passos & Grey (1947) fixed the type locality for adiante as the "Santa
Cruz Mountains, California." The type locality for Edwards' adiaste is also in the
vicinity of the Santa Cruz Mountains.
In view of this information, it becomes apparent that Boisduval's type of
epithore most probably was collected in the vicinity of the Santa Cruz Mountains
and not somewhere in "the high mountains of the east."
Because Boisduval's original description of epithore appeared five years after
Edwards' use of the name, Edwards' epithore takes priority.
Boloria epithore chermocki Perkins and Perkins
Boloria epithore chermocki Perkins and Perkins, 1966, J. Lepid. Soc., 20 (2): 109-112.
The following is quoted from the original description:
"Males: Dorsal LFW (Expanse: 21.10 + 2.53 mm.), holotype 21.40 mm."
"Females: Dorsal LFW (Expanse: 22.01 + 2.74 mm.), allotype 23.00 mm."
"Male: Upper surface: Black spots within median band on both primaries and secondaries tending to be
fused or connected, giving effect of a continuous, irregular black line; on typical epithore, these spots only slightly
connected on primaries, on secondaries without connecting scales. Black basal suffusion heavily represented,
often extending outwardly as far as submedian area; on epithore, black suffusion seldom extending beyond
post-basal region. Segment of vein RS on secondaries bordering cell noticeably accentuated by black scales
connecting this segment to angled figure in cell; rarely an indication of this on epithore."
"Undersurface: On secondaries, submedian-median row of spots chrome-yellow, this region in epithore
cream to yellow with noticeably heavy, ferruginous dusting. Postbasal area of secondaries, below denticulate
white spot bordering vein RS (infrequently invaded by yellow scales) yellow-brown to orange-brown (latter
more common), in this respect, basal and post-basal areas analogous; these areas on epithore red-brown. Post-
median band of secondaries purplish to lilac inwardly, contrasting to the lighter, outward limits of this band; this
contrast much less evident on epithore."
"Female similar in appearance to male."
A type locale was affixed to 0.5-2.9 mi. E. of Dolph, Yamhill County, Oregon.
The holotype and allotype (collected on 18 June 1962 by S. F. Perkins and E. M.
Perkins, respectively), together with 98S and 699 paratypes were deposited in the
collection of the Los Angeles County Museum of Natural History. Of the remaining
paratypes, 6SS and 499 were placed in each of the following institutions: the Calif-
ornia Academy of Sciences, San Francisco, California, The American Museum of
Natural History, New York City, New York, and the Allyn Museum of Entomology,
The authors agreed to name the new subspecies in honor of Franklin H.
Chermock of Baltimore, Maryland. He was the first to call its occurrence to their
attention, and although it was difficult at the time to justify the use of a patronymic
name, Mr. Chermock was known to be dying of a terminal illness.
The description of chermocki was based upon the examination of approx-
imately 250 specimens from various localities in southwestern British Columbia,
Washington, Oregon, and north-central California. Throughout this range, the sub-
species remained constant in appearance. Both its superficial characters and
measurements contrasted favorably to those of peninsular epithore (see Tables 1-4).
These findings are confirmed by the present study, which employed 26056 and
21799 examples of typical chermocki.
Inhabiting moist associations where its probable food-plant, Viola
sempervirens Greene and/or Viola glabella Nuttall is to be found, chermocki occurs
along the west slope of southwestern British Columbia's Coast Mountains and
Lillooet Range, on east-central and southeast Vancouver Island, in the Olympic
Mountains of northwestern Washington, and throughout the Coast Range and
Cascade Mountains of Washington, Oregon, and extreme northern California (refer
to Maps 1 and 2). Occasional demes also appear at lower elevations along the Coast,
and in the more wet, intermountain valleys.
TABLE 2. COMPARISON OF VENTRAL CHARACTERISTICS OF FOUR RACES OF Boloria eptthore1
Race epithore sierra chemocki borealis
Number of typical
specimens examined 103Lf 35 T 242 c d105 ? 260o 217*$ 118'md 7588
Color of submedian-median
row of spots yellowish-cream yellowish-cream chrome-yellow chtbme-yellow
Amount of ferruginous
submedian-median spots ++ +4+ + +/++
Color of HW postbasal
area red-brown red-brown orange-brown brown
Color of W denticulate
spot, bordering vein RS white white usually white usually cream
Postmedian band of HW
delimited inwardly by
dark, violet scaling + + ++ 4++
Numbers of ferruginous
scales on subapical and
outer marginal areas of RF +++ +++ + +
Degree to which pale, sub-
marginal locus between
veins H2-M, of H contrasts
with surrounding, darker
limbal band + + -+ +4.
1Based upon the examination of 1,155 spread specimens (not including material from zones of intergradation).
At the time of the 1966 study, the few available specimens from Clallam,
Whatcom and Okanogan counties in Washington, Umatilla and Wallowa counties
in Oregon, and Glacier County, Montana all evinced a pronounced melanotic
tendency. It was erroneously assumed that these examples merely represented
ecophenotypic variants associated with the more extreme elevations from which
they came. Hence, the distribution of chermocki was described incorrectly to
include the remainder of southcentral British Columbia, southwestern Alberta,
Idaho, and Montana.
Since then, 118 5' and 75 99 examples of B. epithore ssp. have been obtained
from the above-mentioned geographical areas. Comparison of this material to
chermocki reveals yet another unique and geographically distinct subspecies of
B. epithore, not heretofore described.
Boloria epithore borealis Perkins, new subspecies
Males: Dorsal LFW (Expanse: 20.10 + 2.74 mm.), holotype 20.20 mm.
Females: Dorsal LFW (Expanse: 21.31 + 2.81 mm.), allotype 21.30 mm.
Male: Upper surface: Ground color flat, dull orange; unlike yellowish-orange
ground color of chermocki. Like chermocki but unlike nominotypic epithore, black
spots within median band of primaries and secondaries tending to be contiguous
and uninterrupted, forming a continuous, irregular black line. Black basal
suffusion more heavily represented than in chermocki, extending laterally from
basal area of HW and FW to submedian-median areas. Segment of vein RS on
secondaries connecting laterally with bifed figure in cell, like chermocki but unlike
peninsular epithore; unlike chermocki however, figure frequently obscured by black
suffusion. Unlike both chermocki and epithore, degree of black dusting on veins
of both FW and HW prounced, as are amount of black suffusion in marginal area of
FW and relative size of submarginal spots on both wings. Pattern markings and
spots on both FW and HW tending to be greyish black and diffuse, as opposed to
those of both epithore and chermocki which are black to brownish-black and sharply
Undersurface: On secondaries, submedian-median row of spots chrome-
yellow (like chermocki), over which are imposed moderate numbers of ferruginous
scales. Postbasal area of HW brown, unlike orange-brown of chermocki or red-brown
of epithore. Denticulate spot on HW, bordering vein RS, usually cream-colored,
not white. Postmedian band of secondaries delimited inwardly by marked contrast
with dark, refractive blue-violet scaling; this contrast less evident in chermocki, and
much less evident in nominotypic epithore. Like chermocki but unlike epithore,
amount of ferruginous scaling on subapical and outer marginal areas of FW
diminished. Degree to which pale, submarginal locus between veins M,-M,
of HW contrasts with surrounding, more darkly pigmented limbal band greater than
Males and females similar in appearance.
TABLE 3. PARAMETERS OF SAMPLES OF FOUR RACES OF Boloria ep thore1
Race sierra chore cherocki borealis
c$r d5dV d' r V d q da 99
N 242 105 103 35 260 217 118 75
LF, Mean (om.) 19.21 20.62 21.93 23.06 21.24 22.12 20.10 21.31
P.em. (im.) 0.010 0.089 0.064 0.095 0.067 0.110 0.105 0.137
S. D. (im.) 0.75 0.84 0.60 0.51 0.99 1.02 1.06 1.09
v13.63 17.67 8.93 6.62 24.93 26.02 27.94 29.43
U 3.742 4.207 2.988 2.571 4.993 5.101 5.286 5.425
99% limits (am.) 19.21+1.94 20.62+2.17 21.93+1.55 23.061.32 21.24+2.55 22.12+2.63 20.104-2.74 21.31+2.81
1After Brown, 1951.
Holotype 6: Shingle Creek Road, Keremeos, BRITISH COLUMBIA, 26
June 1936 (A. N. Gartrell); placed in the collection of the Entomology Research
Institute, Canada Department of Agriculture, Ottawa, Ontario, Canada. Allotype
9: Keremeos, BRITISH COLUMBIA, 7 June 1923 (C. Garrett), ex coll. J. D. Gunder;
placed in the collection of The American Museum of Natural History, New York
City, New York.
Paratypes (40): BRITISH COLUMBIA: Keremeos, 1-VI-23, 2-VI-23, 4-VI-23,
6-VI-23, 7-VI-23, 766, 10 99 (C. Garrett, ex coll. J. D. Gunder); Hedley, 23-VII-23,
2 S6 (C. B. Garrett); Mt. Apex, vic. Hedley, 26-VII-33, 1 6, 3 99 (A. N. Gartrell);
Mt. Apex, vic. Hedley, 3-VII-67, 1 6 (N. A. Roman); Princeton Summit, 17-VII-09,
1 6, 1 9 (H. Bower); Princeton, VIII-40, 1 9 (L. I. Hewes); Big White Mtn., 40 mi. N.
Vernon, 22-VIII-67, 3 6, 1 9 (N. A. Roman). ALBERTA: Racehorse Cr., Kananaskis
Rd., 12-VII-67, 1 6, 1 9 (R. L. Anderson). MONTANA: Beaverhead Co., Polaris,
25-VI-41, 4-VII-41,14-VII-41, 2 (, 2 99 (collector unknown); Flathead Co., Lake Mac-
Donald, Glacier Nat'l Park, 7-VII-30, 2 99 (E. C. VanDyke). OREGON: Wallowa Co.,
Ice Lake Trail, Mt. Sacajawea, 29-VIII-67, 1 (E. M. Perkins). Three 68 and three 99
have been deposited with the holotype; four 38 and seven 99 have been deposited
with the allotype. Six 66 and two 99 have been placed in the Allyn Museum of
Entomology, Sarasota, Florida. Three 99 have been deposited in the collection of the
California Academy of Sciences, San Francisco, California; five 66 and five 99 have
been placed in the Los Angeles County Museum of Natural History, Los Angeles,
California. One 6 and one 9 have been retained by the author.
TABLE 4. COMPARISON OF SERIES OF FOUR RACES OF Boloria epithore
Race sierra epithore chermocki borealis
it" score compared to:
sierra --- 26.9 17.3 6.0
epithore 26.9 --- 6.8 12.3
chermockt 17.3 6.8 --- 6.5
borealis 6.0 12.3 6.5 ---
1After Brown, 1951; where '"I" scre of at least 6 to 7 supports noticeable difference in samples.
At present, the northernmost, known record for borealis is Smithers, British
Columbia (Map 1). From here it ranges south, through the Fraser Plateau and fertile,
inland valleys of south-central British Columbia, east of the Coast Mountains and
Lillooet Range. In southeastern British Columbia, it occurs in both the Selkirk
and Purcell mountains, extending east to the Continental Divide at a latitude
comparable to that of Revelstoke. Confined in Alberta to the contiguous Rocky
Mountains, it ranges in the north from Bow Pass (north of Lake Louise, Banff
National Park), south to Waterton Lakes National Park, Alberta and Glacier
National Park, Montana. Restricted to extreme western Montana, it occupies
principally the Flathead, Big Hole, and Bitterroot ranges. Found throughout the
northeastern Bitterroot and south-central Sawtooth ranges of Idaho (the latter
representing its southernmost extension), it also occurs in Idaho's north-central
Payette and Nezperce National Forests, to the west of which it "leaks" across to
Oregon's Blue and Wallowa mountains (see Maps 1 and 2).
Records also exist for, or imply, the occurrence of borealis in northwestern
Wyoming, Colorado, and southeastern Alaska; they are, however, somewhat
dubious. Ferris (1971) lists both Yellowstone National Park and Wyoming's Big
Horn Mountains. The former locality was obtained through a collector, second-hand
to him (Ferris, pers. comm.); although the Yellowstone record is possible (Map 2),
the Big Horn Mountains locality is questionable. The senior author has examined a
specimen in the collection of the California Academy of Sciences bearing the data:
"Big Horn Mts., Wy., July, 1925 (?), L. I. Hewes collection." The example agrees
favorably with chermocki, but in no way resembles borealis. Certainly, in this case,
inaccurate locality information had been affixed to the specimen; however, as
Ferris points out (pers. comm.), it is possible that isolated Wyoming colonies of
B. toddi have been confused with B. epithore. The senior author agrees. Among
several museum collections examined during this study, the following species were
inaccurately determined epithore: toddi (Holland), frigga (Thunberg), helena
(Edwards), and kriemhild (Strecker). Such taxonomic confusion probably accounts
for the spurious Colorado records; additionally, neither Eff nor Brown (pers. comm.)
are aware of any legitimate Colorado records of B. epithore. Similarly perplexing
are the Alaska records: although southeastern Alaska has not been sampled
extensively, not one of the some 15,000 butterfly specimens assembled for the Alaska
Lepidoptera Survey has been B. epithore (Philip, pers. comm.). Again it is likely
that frigga, which is common over much of the state, has been confused for epithore;
such was probably the case regarding a St. Michaels, Alaska record (Wright, 1906).
Food-plant preferences for borealis are not known. Dethier (1937) lists none.
Shepard (in press) states: "The author has reared epithore from Viola sp. in Montana.
Overwinters as next to last instar larvae." No other information ia available,
although it is tempting to speculate that V. glabella Nuttall may serve as one of the
food-plants, since Munz and Keck (1968) cite its distribution to include Alaska and
the northern Rocky Mountains.
ab. obscuripennis (Gunder)
Brenthis epithore Bdv., ab. obscuripennis Gunder, 1926, Ent. News, 37(1): 7.
The type of this aberration, a female, was collected at Chilcotin, British
Columbia, on May 30,1915. Other examples have been reported by Jones from Alexis
Creek and Shuswap Lake, British Columbia (Occ. paper #1, Ent. Soc. B. C., June,
1951). In the original description Gunder stated: "Primaries entirely fogged over
with dark shading, obscuring and submerging maculation, especially on the inner
half with cell quite dense where only a single yellow brown spot shows; normal
row of round black spots indistinctly visible. Secondaries, outer half normal;
confused yellow brown maculation of inner half externally edged by black shading
which extends also along the costal margin, basal area quite dark." The preceding
description applies to the dorsal surfaces. Because it is a melanic aberration,
originally described as such, obscuripennis must be considered infrasubspecific
to B. epithore borealis.
Although, in the 1966 study, certain examples of B. epithore from Oregon
and California were noted to be somewhat unusual, insufficient numbers (57 $8
and 27 9 ), compounded by inadequate knowledge of their biology and the inability
to interpret the meaning of their bifid distribution in California and zones of
intergradation in both states, cast considerable doubt upon their taxonomic validity.
The authors were also reticent to affix a new taxon based solely upon the criterion
of size--even though the specimens were statistically smaller. Hence, the opportunity
to assign a subspecific rank to the possibly new entity was precluded.
Since then, 242 &3 and 105 99 have been assembled and examined. Com-
parison of this material to both chermocki and nominate epithore reveals yet another
unique and geographically distinct subspecies of B. epithore, not heretofore
S++ + + +
0+ + + +
+ + + +
0 o oo
Map 1: known distribution of the Boloria epithore complex, based upon the
determination of one or more of the 1,155 specimens examined during this study.
Various symbols correspond to respective subspecies: (*) = B. e. epithore, (o) =
B. e. chermocki, (+) = B. e. borealis, and (x) = B. e. sierra.
o %o +
Boloria epithore sierra Perkins, new subspecies
Males: Dorsal LFW (Expanse: 19.21 + 1.94 mm.), holotype 18.40 mm.
Females: Dorsal LFW (Expanse: 20.62 + 2.17 mm.), allotype 19.00 mm.
Male: Upper surface: Ground color bright orange, with sharply demarcated,
black to brown-black superimposed pattern markings and spots; black basal
suffusion and melanic dusting along veins of FW and HW mild, as in nominotypic
epithore. Like chermocki but unlike epithore, HW vein RS connecting laterally with
bifed figure in cell, and black spots of FW and HW median bands contiguous. Unlike
either epithore or chermocki, black basal suffusion on HW marginal area moderate,
and relative size of submarginal spots on both wings small.
Undersurface: On secondaries, submedian-median row of spots yellowish-
cream (like epithore), over which are imposed large numbers of ferruginous scales,
sometimes so numerous as to obscure color of spots in submedian-median row; in this
respect, unlike nominotypic epithore or any other subspecies. Like epithore but
unlike chermocki, color of HW postbasal area red-brown; postmedian band of HW
meagerly delimited inwardly by dark violet scaling; and ferruginous scales on
subapical and outer marginal areas of FW plentiful. In other respects, comparable
to peninsular epithore.
Males and females similar in appearance.
Holotype 8: Sentinel Dome, Yosemite National Park, Mariposa County,
CALIFORNIA, 3 July 1946 (F. H. Rindge, ex. colln. D. C. Ferguson); placed in the
collection of the Allyn Museum of Entomology, Sarasota, Florida. Allotype 9:
Glacier Point, Yosemite National Park, Mariposa County, CALIFORNIA, 2 July
1921 (J. A. Comstock); placed in the collection of the Los Angeles County Museum of
Natural History, Los Angeles, California.
Paratypes (42): CALIFORNIA, Yosemite Nat'l Park, Mariposa Co.: Glacier
Point, 28-VI-21, 2-VII-21, 16 8~, 7 99 (J. A. Comstock), (Note: some labels on the 23
preceding Comstock specimens simply bear the hand-written data: "nr. Yosemite,
7700' el., July 2,1921." However, because they are identical to Comstock's other data-
determination labels, and because the specimens bear the same date, elevation, and
were borrowed from the same institution as were those specimens clearly labelled:
"Glacier Point, nr. Yosemite, Cal., July 2, 1921, 7700 ft. elev., Br. epithore," it is
assumed that they indeed represent Glacier Point material, spread or labelled at a
different time or by a different individual); Tenaya Canyon, 11-VII-58, 3 SS, 1 9
(A. O. Shields); Yosemite, 21-VI-34, 2 66, 1 9 (T. Craig); Yosemite Valley, 20-VII-30,
1 (, 3 99 (E. C. Zimmermann); Hy. 120, Yosemite Nat'l Park, 15-VI-61, 3-VII-62, 5S ,
1 9 (E. M. & S. F. Perkins); Tuolumne Co.: Tioga Pass, 11-VII-31, 1 9 (J. W. Tilden);
vic. Tioga Pass, 1-VIII-52, 1 S (P. A. Opler). Five S6 and one 9 have been deposited
with the holotype; 15 S3 and six 99 have been deposited with the allotype. Three SS
and one 9 have been placed in The American Museum of Natural History, New York
City, New York; four SS and five 99 have been deposited in the California Academy
of Sciences, San Francisco, California. One S and one 9 have been retained by the
Although sympatric populations and intergradations between chermocki and
sierra typify drier, south-central Oregon (Jackson, Klamath and Lake counties) and
lava-strewn, north-central and northeastern California (Lassen, Modoc, Siskiyou
and Shasta counties), demes of typical sierra occasionally occur to the south and
west-at higher elevations--in the Coast Range Mountains (Glenn, Lake, Mendocino,
Tehama and Trinity counties) and consistently occur to the south and east,
throughout the length of the Sierra Nevada (from Plumas and Sierra counties in
the north, to Kern County in the south). See Maps 1 and 2. The southernmost
extension of sierra was discovered in 1961 by Stanford (Perkins and Perkins, 1966),
who was collecting in the vicinity of Tobias Peak, located in the Greenhorn
Mountains south of the Tulare-Kern county line. It is interesting to note, botanically,
that Tobias Peak also represents the southernmost limit of sierra's known food-
plant: V. glabella Nuttall (J. F. Emmel, pers. comm.).
Map 2: projected distribution of Boloria epithore's four subspecies in western
North America, based upon all determinations and records presented in this
revision. Variously shaded areas correspond to respective subspecies: dark (re-
stricted to areas immediately north and south of San Francisco Bay) = B. e. epithore;
medium = B. e. chermocki; very dark = B. e. borealis; and light = B. e. sierra.
5 ~er 6
Plate I: Adult males of Boloria epithore complex: 1) sierra holotypee),
Sentinel Dome, Yosemite Nat'l Park, Mariposa Co., Calif., 3-VII-46 (F.H. Rindge);
2) sierra (paratype), Glacier Point, Yosemite Nat'l Park, Mariposa Co., Calif.,
2-VII-21 (J.A. Comstock); 5) epithore, Big Basin, Santa Cruz Mts., Santa Cruz Co.,
Calif., 4-V-46 (O.E. Sette); 6) epithore, Santa Cruz Mts., Santa Cruz Co., Calif.,
15-VI-46 (T.W.Davies); 9) chermocki (topotype), 2.9 mi. E. Dolph, Yamhill Co.,
Oregon, 18-V-68 (S.F. Perkins); 10) chermocki (topotype), 2.9 mi. E. Dolph, Yamhill
Co., Oregon, 18-V-68 (S.F. Perkins); 13) borealis holotypee), Shingle Creek Road,
Keremeos, B.C., 26-VI-36 (A.N. Gartrell); 14) borealis (paratype), Keremeos, B.C.,
2-VI-23 (C. Garrett). Figures to the left are dorsal; those to the right are ventral. All
specimens actual size.
3 I 4
'I. I I. = ''^f ils-
Plate II: Adult females of Boloria epithore complex: 3) sierra (allotype),
Glacier Point, Yosemite Nat'l Park, Mariposa Co., Calif., 2-VII-21 (J.A. Comstock);
4) sierra (paratype), Glacier Point, Yosemite Nat'l Park, Mariposa Co., Calif., 2-VII-
21 (J.A. Comstock); 7) epithore, Big Basin, Santa Cruz Mts., Santa Cruz Co., Calif., 4-
V-46 (O.E. Sette); 8) epithore, Gazos Creek Road, 5.2 mi. N.E. Hy. 1, San Mateo Co.,
Calif., 30-V-68 (J.F. Emmel); 11) chermocki (paratype), 2.9 mi. E. Dolph, Yamhill Co.,
Oregon, 10-VI-62 (E.M. Perkins); 12) chermocki (paratype), 2.9 mi. E. Dolph, Yamhill
Co., Oregon, 12-VI-63 (E.M. Perkins); 15) borealis (allotype), Keremeos, B. C., 7-VI-23
(C. Garrett); 16) borealis (paratype), Keremeos, B. C., 7-VI-23 (C. Garrett). Figures
to the left are dorsal; those to the right are ventral. All specimens actual size.
Map 3: proposed evolvement, dispersal and geographic isolation of Boloria
epithore complex. Arrows indicate probable, southerly movements from points
of origin, where broken arrows = B. e. epithore; large, open arrows = B. e. chermocki;
large, solid arrows = B. e. borealis; and narrow, elongated arrows = B. e. sierra.
ab. eldorado (Strand)
Brenthis epithore Bdv., cum. ab. eldorado Strand, 1914, Archiv fiir Naturgeschichte,
80 (A) pt. 11: 156.
Described on the basis of six examples from Plumas County, California
(collected June 10-17, 1913), and one specimen from El Dorado County, California
(collected between June 25 and June 28, 1913), the distinguishing features of
eldorado, as indicated by Strand in his original description, include: (1) the black
markings on the underside of the forewings are large and consequently appear to
be near to one another; (2) the two transverse spots in the middle of the field (in
cells M3and CUI) are connected by means of a median, black longitudinal line;
(3) the angled figure in the cell (discal cell) completely or almost completely touches
the discocellular spots; and (4) the three or four postmedian spots touch or almost
touch the transverse lines in the form of black flecks. Strand also mentioned that
the black design above is stronger in both wings.
From this information and a study of material from both Plumas and El
Dorado counties, the senior author concludes that Strand's name, eldorado,
represents an aberration. Under the rules of the International Code of Zoological
Nomenclature, eldorado must therefore be regarded as being infrasubspecific in
rank to B. epithore sierra.
ab. wawonae (Gunder)
Brenthis epithore Bdv., ab. wawonae Gunder, 1924, Ent. News, 35(5): 156.
The type was collected at Wawona (in Yosemite National Park), Mariposa
County, California on July 6, 1922. It is pictured in J. A. Comstock's Butterflies of
California, P1. 26, fig. 10. The distinctive feature of wawonae is found on the
secondaries where the row of postmedian spots is "lacking."
As is eldorado, wawonae is now considered an infrasubspecific entity of
B. epithore sierra.
A revision of the Boloria epithore complex would be incomplete without
briefly reviewing and summarizing the salient phenotypic characters of each of its
races. In order to avoid redundancy, the reader is referred to Plates 1-2 and Tables
1-4, which serve this purpose.
Similarly, it would be less meaningful if present knowledge regarding the
geographic distribution, zones of intergradation, ecological associations, and food-
plant preferences of B. epithore's four subspecies were not incorporated into an
overview of those paleogeological mechanisms that possibly affected the dispersion
and geographic isolation of its various conspecific populations.
During the Pleistocene, polar isotherms were depressed southward through-
out the Northern Hemisphere, causing continental glaciation in central North
America, and alpine glaciation in the Cordillera, Sierras, and Pacific Coastal
ranges. Atmospheric and physiographic control of climate may be linked to
Pleistocene tectonic and glacial events, which apparently controlled the distri-
bution of the four subspecies of Boloria epithore.
It appears that borealis represents the parent stock from which the other
subspecies evolved during southern migration (see Map 3). It is possible that the
migration of borealis, and the southward progression of the floral ecosystem
(Axelrod, 1957; Munz & Keck, 1968) upon which the species is dependent, was
initiated with the onset of Wisconsin glaciation and the associated southward
depression of isotherms. The southeasterly dispersal trend parallels the pre-
dominant structural trends of the central Cordillera, following valley-conduits
during southward migration. This structural trend eliminates wind shadow effects,
allowing penetration of moisture-laden sea winds into the continental interior.
Examination of Map 2 would indicate that borealis and the other subspecies of
B. epithore are dependent upon a wet, montane habitat; hence, their distribution is
limited to such areas.
To the south, borealis split into western and eastern tongues, separated by
the Columbia plateau (see Map 3). Underlain by Miocene and possibly Pliocene
basalt flows, the Columbia plateau is now capped by loess deposits accumulated
during successive Pleistocene glaciations (Richmond et al., 1965), producing a
foreign ecosystem not inhabitable by borealis.
A rather sharp transition between borealis and chermocki occurs in the
Fraser lowlands (see Maps 2-3). During Wisconsin glaciation, the Fraser lowlands
were occupied by an alpine glacial system (Crandell, 1965) that effectively eliminated
communication between areas north and south of it. Such a boundary would have
existed until 11,000 ybp (years before present) when glacial retreat opened the
Pacific Northwest (Bandy and Wilcoxin, 1970).
As indicated by Map 2, chermocki inhabits the rain forests of Oregon and
Washington. This peculiar climate occurs where moisture-laden marine winds,
blocked by the Klamath and Cascade Mountain ranges, are forced to rise, cool
adiabatically, and release their water content. Hence, chermocki is essentially
limited to the windward sides of these ranges, indicating that abundant rainfall
is necessary for its survival. Leeward of these ranges, the climate is considerably
drier due to rain shadow effects; therefore, the crest of these mountains acts as
an effective physiographic and ecologic barrier.
It is possible that the southward migration of borealis and its concomitant
evolvement into chermocki (Map 3) was made possible by tectonic events in the
Klamath and Cascade mountains. Because these mountains are young, probably
attaining their present altitude and relief in the Pleistocene (Wahrhaftig and
Birman, 1965), the moist climate necessary for survival of chermocki may not have
developed in this area until these coastal ranges had been sufficiently uplifed to
form a barrier to the inland penetration of moisture-laden winds.
A zone of intergradation between chermocki and sierra occurs in the area of
the Modoc plateau and adjacent volcanic terrains of south-central Oregon and
north-central California (Klamath, Lake, Siskiyou, Modoc, Shasta, and Lassen
counties). The area is characterized by an irregular basaltic plateau, 1,200-1,500
m. high. The presence of a recently active volcano, Lassen Peak (Wahrhaftig
and Birman, 1965), indicates that this transitional faunal boundary may still be in
the process of forming. Hence, an overlap of subspecies in this area might be expected
(see Maps 1-3).
The Great Valley of California represents a major ecologic boundary that
divided the further, southward dispersal of chermocki into two populations:
nominate epithore and sierra (Map 3).
The present distribution of epithore is spotty, being limited 1) to the wet
western face of the California coastal ranges immediately adjacent to the coast, e. g.,
Mendocino County, where its recorded food-plant is Viola sempervirens Greene,
and 2) to its major occurrence as an outlie--an area confined to the Santa Cruz
Mountains (600 m.), some 120 miles south of its discontinuous, Mendocino County
parent group (Maps 1-2). The area separating these two populations has been
rather extensively cultivated, thus eliminating to some extent the availablility
of V. sempervirens, upon which it was dependent. The resultant isolation of penin-
sular epithore, therefore, may have led to its subspeciation from chermocki, and
adoption of V. ocellata Torrey & Gray as its major food-plant in the Santa Cruz
As described by Peabody and Savage (1958), southern migration of Arcto-
Tertiary amphibians occurred along a coastal corridor opened by eustatic lowering
of sea level accompanying Pleistocene glaciation. So too might have epithore effected
its southward migration, from Mendocino County in the north to San Mateo, Santa
Clara, and Santa Cruz counties in the south. B. e. sierra, however, was confined to
the mountainous terrain of the North Coast Range, where it was associated with
its food-plant, V. glabella Nuttall (J. F. Emmel,pers. comm.). Because the lowering of
sea level caused the emergence of coastal lowlands only, its migration potential
remained unaffected by the emergent corridor (Maps 2-3).
The California coastal ranges inhabited by sierra are not high (800-2,200
m.), and would trap only a portion of the moisture borne by sea breezes. The re-
maining moisture passes over the coast ranges and is screened by the western face
of the Sierra Nevada (750-3,100 m.), east of the Great Valley. Hence, with the ex-
ception of an apparent southerly false start down the eastern coastal ranges, where
its aborted penetration terminates, north of the Napa Valley, sierra is presently
distributed largely along the wet western slope of the Sierra Nevada between
Plumas and Kern counties (Maps 1-3), where its recorded food-plant is V. glabella
(1) As of 1966, the Boloria epithore complex consisted of two distinct sub-
species: Boloria epithore epithore Edwards and Boloria epithore chermocki Perkins
& Perkins. A revised treatment of the complex was proposed. At the time, limited
numbers of available specimens, together with inadequate criteria for additional
systematic analysis, negated further nomenclatorial designation.
(2) In the present revision, 920 spread specimens have been acquired and
examined, in addition to the original 254 employed in the 1966 study. A reevaluation
of distributional records, coupled with a more critical examination of phenotypic
character analyses and statistical interpretation of linear measurements, clearly
demonstrates that the Boloria epithore complex is comprised of four distinct
subspecies: Boloria epithore epithore Edwards, Boloria epithore chermocki Perkins
& Perkins, Boloria epithore borealis Perkins, and Boloria epithore sierra Perkins.
(3) A revised treatment of the epithore complex is proposed:
603 epithore (Edwards), 1864
a e. epithore (Edwards), 1864
b e. chermocki Perkins & Perkins, 1966
c e. borealis Perkins, 1973
ab. obscuripennis (Gunder), 1926
d e. sierra Perkins, 1973
ab. eldorado (Strand), 1914
ab. wawonae (Gunder), 1924
The following records represent 1,155 examples of Boloria epithore ex-
amined by the senior author during the course of this study (collectors are listed
alphabetically at the end):
Banff, (?), (ERI coll.); Blairmore, (?), (R.H.); Carbondale, Crow's Nest Pass Forest,
14 mi. S. Coleman, 20-VII-68, (S.S.); Jet. Kananaskis Hy. & Oldman R., 24 mi. N.
Coleman, 16-VII-67, (R.E.W.); 39 mi. N. Lake Louise, Banff Nat'l Park, (?), (J.L.);
Racehorse Creek, Bow River Forest Reserve, 16 mi. N. Coleman, 10-VII-67, (S.S.);
Racehorse Creek on Kananaskis Rd., 12-VII-67, (R.L.A.); Waterton Lakes Nat'l
Park, (?)/8-VI-58, (J.B./S.S.).
Ainsworth, 3-VI-03, (G.H.F.); Alexis Creek, VII, (?); Alta Lake, 10/12-VI-26, (J.M.);
Apex Mtn., Hedley, 26-VII-33, (A.N.G.); Apex Mtn. L.O., 34 mi. W. Penticton,
3-VII-67, (N.A.R.); Big White Mtn., 40 mi. N. Vernon, 22-VIII-67, (N.A.R.); Coalmont,
nr. Princeton, 3-VII-68, (S.S.); Duncan, Vancouver Island, (?), (C.L.); Fitzgerald,
S. Shawnigan Lake, V. I., 26-V-19,15/26-VI-19, 3-VI-21, (W.R.C.); Forbidden Plateau,
V. I., 10-VII-38, (T.D.G.); Glacier, Glacier Nat'l Park, 13-VII-04, (ERI coll.); Gold-
stream, V. I., 2-VI-18, (W.D.); Grouse Mtn., (?), (S.S.); Harrison Lake, (?), (S.S.);
Hedley, 16/23-VII-23, (C.B.G.); Hope, (?), (S.S.); Hope Mts., 20-VII/8-VIII-32,
(A.N.G.); Kaslo, 7-VI-03, (J.D.G.); Kelowna, VII-20/(?), (A.T./J.L.); Keremeos,
1/7-VI-?, (C.B.G.); Keremeos, Shingle Creek Rd., 26-VI-36, (A.N.G.); Lambly Lake,
nr. Kelowna, (?), (H.F.M.); Lihumption Park, nr. Cultus Lake, 1/2-VIII-27, (C.H.Y.);
McCheam, 22-VII-16, 28-VII-15, (R.C.T.); Maillardville, N. New Westminster,
28-VI-29, 2-VII-29, (R.H.R.); Maillardville, 28-VI-29, 30-VI-29, 2/11-VII-29, (A.E.W.);
Malahat, 6-VI-23, (W.D.); Mt. Kobau, nr. Oliver, (?), (H.F.M.); Nanaimo, V. I.,
23/27-VI-20, (E.C.V.); Nanoose, V. I., 9-VI-55, (D.E.); N. Vancouver, 24-V-02, (ERI
coll.); Princeton, 17-VII-09/VIII-40, (H.M.B./L.I.H.); Quamichan, V. I., 27-IV-31,
(J.F.); Revelstoke, 17-V-70, (D.T.); Rogers Pass, (?), (J.L.); Rossland, 19-VI-00,
(W.H.D.); Royal Oak, 12-VI-17, (W.D.); Salmon Arm, vic. Shuswap Lake, (?), (D.T.);
Shuswap Lake, VII, (?); Silver Star Mtn., nr. Vernon, (?), (D.T.); Smithers, 19/31-VII-
31, (J.D.G.); Stuart Island, '33-'36, (F.H.R.); Tod Inlet, 13/20-VI-28, (W.H.A.P.);
Vancouver, 28-V-06, 30-V-03, (ERI coll.); Victoria, V. I., 20-V-93, (ERI coll.);
Wellington, V. I., 23-V-50/ 15-VI-50/ 4-VI-49/ 6-III-04/ 27-VK'/ 16-V-56/ 26-VI-02,
(T.W.D./T.W.D./R.G./E.A.D./D.E./D.E./ ERI coll.).
ALPINE CO.: Elephant Lake, 4-VII-54, (P.A.O.); Lake Alpine, 24-VII-30, (R.P.A.);
AMADOR CO.: Cook's Station, Kit Carson Pass Rd., 20-VI-40/23-VI-40, (T.W.D./
W.A.H.); 27 mi. E.N.E. Jackson, 14-VII-58, (O.E.S.); CALAVERAS CO.: nr. Big
Meadows, along Stanislaus R., 4-VII-68, (A.O.S.); EL DORADO CO.: Mt. Tallac,
20-VII-09, (F.X.W.); Echo Lake, 29-VI-61, (A.O.S.); Gillmore's Ranch, head of Fallen
Leaf Lake, 23-VII-92, (W.G.W.); Fallen Leaf Lake, VII-15, VII-16, (L.S.R.); Glen
Alpine Creek, 17/18-VII-09, (F.X.W.); Glen Alpine, 29-VI-29, (E.P.V.); Union Valley
Reservoir, (?), (N.L.L.); FRESNO CO.: Round Meadow, Huntington Lake, 3/4-VII-31/
9-VII-60, (M.L.W./O.E.S.); Huntington Lake, 14/18-VII-35/ 3-VII-19/ 18-VI-30/
4-VII-50, (M.L.W./E.P.V./R.H.R./?); GLENN CO.: Plaskett Meadows Camp,
4-VII-72, (N.L.L.); HUMBOLDT CO.: Green Point, 10/12-VI-16, (E.C.V.); Redwood
Creek, 5-VI-16, (E.C.V.); KERN CO.: AIGER Flat Campground Rd., Greenhorn
Mts., 24-VI-61, (R.E.S.); MARIPOSA CO.: Glacier Point, Yosemite Nat'l Park,
1/2-VII-21/ 28-VI-21/ 29-VI-37, (J.A.C./J.A.C./?); Yosemite Valley, 20-VII-30,
(E.O.Z.); Hy. 120, Yosemite Nat'l Park, 15-VI-61, 3-VII-62, (E.M.P.); Yosemite,
21-VI-34/ 30-VII-28, (T.C./?); Yosemite Park, VI-24, (J.A.C.); Wawona, 25-VI-21,
(J.A.C.); Wawona, 25-VI-21, (J.A.C.); Sentinel Dome, Yosemite Nat'l Park,
3-VII-46, (F.H.R.); Tioga Rd., high Sierras, Yosemite Nat'l Park, VII-19, (S.T.); Tenaya
Cy., Yosemite Nat'l Park, 11-VII-58, (A.O.S.); MENDOCINO CO.: nr. Casper, 4 mi N.
Mendocino, 30-V-70; (P.A.O.), Ukiah, Halfway House, 18-VI-94, (W.G.W.); Man-
chester, 13/16-VIII-05, (E.J.N.); Mendocino, 28-IV-85, (W.G.W.); Graveyard Rd., E.
Russian Gulch State Park, 30-V-70, (J.F.E.); Chipmunk Spgs., N.W.Hull Mtn., 31-V-
70, (J.F.E.); Russian Gulch State Park, 28-III-70, (R.L.L.); MODOC CO.: nr. Eagle-
ville, 24-VII-37, (J.A.C.); MONO CO.: Tioga Lodge, Mono Lake, 22-VI-29, (E.P.V.);
NEVADA CO.: Donner Summit, nr. Truckee, 23/30-VI-60, (J.F.E.); PLACER CO.:
Deer Park, 2 mi. W. Lake Tahoe, 5-VII-63, (E.M.P.); Deer Park, 7/14-VII-09,
(E.J.N.); McKinney Creek, 3 mi. S.W. Tahoma P.O., 10-VII-48, (O.E.S.); Lake Tahoe,
22-VII-53, 29-VI-30, (LACM coll.); nr. Lake Tahoe, 8-VIII-22, (LACM coll.);
Ward Creek, 2 mi S. Tahoe City, VII-66, (N.W.); PLUMAS CO.: Gold Lake, 15-VIII-53,
(T.W.D.); Buck's Lake, 23-VI-49/ 4-VII-57, (C.I.S./R.L.L.); vic. Quincy, (?), (T.W.D.);
SAN MATEO CO.: Portola State Park, 19-VI-60, (C.N.S.); Wood Haven Girl Scout
Camp, nr. La Honda, 13-VI-54, (D.H.B.); San Lorenzo Woods, (?), (O.E.S.); Gazos
Creek Rd., 5.2 mi. N.E. Hy. 1, 30-V-68, (J.F.E.); SANTA CLARA CO.: Santa Clara,
29-V-04, (E.A.D.); Saratoga, 7-VI-44, (ERI coll.); SANTA CRUZ CO.: Big Basin,
Santa Cruz Mts., 10/19-VI-45, 4-V-46, 4-V-47, (O.E.S.); Big Basin Redwoods State
Park, 19-VI-50, (D.M.); 7 mi. N.W. Boulder Creek, 4-V-47, (O.E.S.); 6 mi. N.W.
Boulder Creek, 16-V-63, (O.E.S.); Santa Cruz Mts., 15/16-VI-46/ 14/18-V-95/VII-01/
?, (T.W.D./CAS coll./LACM coll./CAS coll.); Felton, 20-V-56, (J.W.T.); Santa Cruz,
18-V-24, (J.F.S.); Santa Cruz, 12-V-35, (J.W.T.); Santa Cruz, 2-VI-19, (E.P.V.);
Hecker Pass, 22-IV-56, (P.A.O.); Soquel, 12-V-29, (C.E.P.); Santa Cruz, 7-VI-92,
(W.G.W); Boulder Creek, Big Basin State Park, Santa Cruz Mts., 17-VI-61, (J.S.
& P.E.); Boulder Canyon, 28-IV-55, (D.E.); China Grade, 2-VI-62, (D.E.); Bear Creek,
4 mi. E. Boulder Creek, 28-V-72, (D.S.); SHASTA CO.: Lassen Nat'l Park, 21-VII-37,
(J.A.C.); SIERRA CO.: Gold Lake, 15/16-VII-35/ 12-VII-41/ 1-VII-41, (R.H.R./
D.E./D.E.); nr. Gold Lake Lodge, 18/19-VII-65, (A.O.S.); Yuba Pass, 15-VII-62,
(R.L.W.); Bassetts, 12/13-VI-60, (P.A.O.); Yuba Pass, 30-VI-61, (A.O.S.); SISKIYOU
CO.: vic. McCloud, S. base Mt. Shasta, 24-VI-67, (E.M.P.); Bigelow Meadows, 16 mi.
S.E. Mt. Shasta, 27-VI-59, (O.E.S.); Mt. Shasta, VII-04, (F.X.W.); 0.25 mi. S. Toad
Lake, 19 mi. W. Mt. Shasta, 22-VII-67, (E.M.P.); Castle Lake, 9-VII-60/ 9-VII-41/
18-VII-50/ 28-VI-47/ 22-VII-67/ 24-VII-48/ 24-VI-48, (?/ A.J.U./ R.F./ T.W.D./
E.M.P./D.E./D.E.); TEHAMA CO.: Mill Creek, 22-VI-61, (J.W.T.); Mill Creek, 8 mi.
S.W. Mt. Lassen, 1/4-VII-56, (O.E.S.); Mineral, 26/27-VI-30, (?); TRINITY CO.:
Trinity Center, 6-VI-31/ 6-V-31, (H.M.B./ ERI coll.); Carrville, 18-VI-31, (E.C.V.);
Forest Glen, (?), (D.B.); TULARE CO.: Sequoia Nat'l Park, 9-VII-29/ 7/8-VII-39/
21-VII-37, (G.H./?/A.J.U.); Mineral King, 26-VII-36, (G.H.); TUOLUMNE CO.:
Tioga Pass, Yosemite Nat'l Park, 11-VII-31/ 1-VIII-52, (J.W.T./ P.A.O.); Pinecrest,
BLAINE CO.: Beaver Creek, nr. Alturas Lake, 28-VII-44, (D.E.); CUSTER CO.:
S. Stanley, 10-VII-44, (D.E.); Inlet Campground, Stanley Lake, Challis Nat'l
Forest, 8-VII-72, (S.E.); IDAHO CO.: S. Fork Clearwater R., vic. Golden, 13-VI-44,
(L.I.H.); Lolo Pass, 11-VIII-61/ 28-VII-63, (S.F.P./ D.E.); KOOTENAI CO.:
Trapper Creek at Bunco Rd., 8-VII-67, (R.L.C.); LATAH CO.: Idaho Hy. #3,
Shoshone Co. Line, 12-VII-72, (S.E.); SHOSHONE CO.: Wallace, 20-V-16, 17-VI-17,
7-VII-17, 12-VII-17, 19-VII-17, 20-VII-17, 9-VI-18, 15-VI-18, 1-VII-18, 9-VII-18, (O.H.);
Pine Creek, Bitterroot Range, (?), (O.H.); VALLEY CO.: Forest Rd. 082, Fir Creek, nr.
Fir Creek Campground, 8-VII-72, (S.E.); Deadwood, Deadwood R., 8-VII-72,
(S.E.); McCall, 12-VII-67, (R.L.C.).
BEAVERHEAD CO.: Polaris, 25-VI-41, 4-VII-41, 14-VII-41, (LACM coll.); Big Hole
National Monument, 10-VII-59, (T.W.D.); Polaris, 13-VII-42, 5-VII-48, 14-VII-43,
26-VI-43,11-VII-42, (D.E.); FLATHEAD CO.: Sperry Chalets, Glacier Nat'l Park,
24-VIII-25, (C.E.P.); Lake McDonald, Glacier Nat'l Park, 7-VII-30, (E.C.V.);
GLACIER CO.: St. Mary Lake, Glacier Nat'l Park, 1-VII-61, (E.M.P.); E. boundary
Glacier Nat'l Park, 9-VII-56, (R.L.A.); MINERAL CO.: Randolph Creek, 5-VII-72,
(S.K.); Lookout Pass, 11-VIII-61, (S.F.P.); 2 mi. S. St. Regis, Hy. 10, 10-VIII-61,
(S.F.P.); MISSOULA CO.: Miller Creek, 12-VI-71, 19-VI-71, (S.K.); Ninemile
Creek, 3-VII-72, 10-VII-72, (S.K.).
BENTON CO.: McDonald Forest, 5 mi. N.W. Corvallis, 29-V-60/ 28-V-65/ 21-V-58/
25-VI-62/ 20-V-59/ 16-V-58, (R.E.W./E.M.P./E.M.P./E.M.P./E.J.D./E.J.D.); Mary's
(E.J.D./E.J.D./E.J.D./E.J.D./F.F.H., OSU coll./ F.F.H., OSU coll./A.H.M./
E.G.); Alsea Fish Hatchery, 31-V-64/25-V-30, (E.J.D./L.G.H., OSU coll.); Hoskins,
1-VI-58, (E.J.D.); Corvallis, 11-VI-25, (E.C.V.); CLACKAMAS CO.: Clackamas R.,
Rd. 284, 7-VI-62, (E.M.P.); 6 mi. W. Timothy Lake, 7-VI-62, (E.M.P.); Mt. Hood,
27-VI-26/16-VI-61, (L.I.H./R.J.A.); Clackamas Lake, 3-VII-61, (E.J.N.); Austin
Hot Spgs., 16-VI-57, (S.G.J.); Big Eddy, 18-V-58, 1-VI-52, (S.G.J.); Eagle Fern Park,
4-V-58, 2-VI-57, 4-VI-61, (S.G.J.); Barton, 15-V-60, (S.G.J.); CLATSOP CO.:
Saddle Mtn. State Park, 27-V-64, 12-VII-64, (E.M.P.); Saddle Mtn., 8-VII-61,
(S.G.J.); COLUMBIA CO.: Vernonia, 20-V-38, (OSU coll.); 4 mi. N. Vernonia,
6-VI-70, (C.W.N.); CURRY CO.: Winechuck R., 17-VI-67, (G.M.H.); DESCHUTES
CO.: Deschutes R. Bridge, W. Terrebonne, 8-VII-53, (S.G.J.); 3 Creeks Meadow,
6-VII-72, (E.J.D.); DOUGLAS CO.: Diamond Lake, 13-VII-61/1-VII-47/8-VII-?/
28-VII-53, (A.O.S./T.W.D./E.A.D./S.G.J.); Drew, 7-V-39, (L.I.H.); Scottsburg,
24-V-31, (L.I.H.); Reedsport, 15-V-26, (E.M.); Bradley Creek Meadows, 2-VIII-62,
(E.J.N.); Jet. Muir Creek & Rogue R., 19-VII-56, (A.H.M.); HOOD RIVER CO.:
Cloud Cap Rd., Mt. Hood, Hy. 35, 12-VI-62, (E.M.P.); W. Dee, 5-VII-70, (J.H.);
JACKSON CO.: Mt. Ashland Loop Rd., 8-VII-61/9-VII-61, (A.O.S./D.D.); Tubb
Spgs., 4 mi. W. Pinehurst, 19-VI-58, 17-VI-58, (E.J.D.); Kane Creek, 5 mi. W. Gold
Hill, 23-VI-37, (S.G.J., OSU coll.); Butte Falls, 5-V-68, (D.E.); Dead Indian Soda
Spgs., 17-V-62, (J.H.S.); Mt. Ashland, 16-VI-68, 19-VI-70, (E.J.D.); French Gulch
Rd., 22-V-64, (E.J.D.); JEFFERSON CO.: Camp Sherman, Metolius R., 21-VI-64/
21-VI-66/ 27-V-66/ 14-VI-64/ 28-V-50, (E.M.P./E.M.P./E.M.P./C.W.N./S.G.J.);
Summit Santiam Pass, 31-VII-53, 8-VII-59, (S.G.J.); JOSEPHINE CO.: vic. O'Brien,
15-VII-58, (R.J.A.); KLAMATH CO.: Gilchrist, 15-VI-58/ 24-V-59/ 15-VI-60/
10-VII-60/ 15-VI-53, (E.J.D./E.J.D./D.E./D.E./D.E.); 8-10 mi. E. Beaver Marsh,
14-VII-61, (A.O.S.); Davis Lake, 15-VII-56/ 22-VII-34, (A.H.M./ S.G.J., OSU coll.);
Crater Lake Nat'l Park, Annie Creek, 16-VII-56, (A.H.M.); Crescent Creek at
Hy. 58, 27-VI-62, (K.G.); Skookum Meadows, 23-VII-61, 30-VI-62, (E.J.N.); 6 mi.
S. La Pine, 3-VII-68, (E.J.D.); LAKE CO.: Camas Creek, Summit Prairie, S.E.
Warner Cy., 12-VI-62, (E.J.N.); Lakeview, 27-VII-30, (OSU coll.); LANE CO.:
3 mi. W. Willamette Pass, 2-VII-60, (R.E.W.); Willamette Pass, 3-VII-59, (E.J.D.);
Oakridge, 11-VI-55, (E.J.D.); Hills Creek Dam Rd., 13-VI-64, (E.J.D.); Blue Pond
Forest Camp, 13-VI-64, (E.J.D.); Mule Prairie, 13-VI-64, 18-VI-57, 22-VI-58,
3-VII-59, (E.J.D.); Mule Prairie, 2-VIII-62, (E.J.N.); LINCOLN CO.: Elk City,
30-V-59, (E.J.D.); LINN CO.: Lost Prairie, 28-VI-60/ 15-VII-65/ 11-VIII-64/
20-VI-65/22-VII-64/ 12-VI-66/ 15-VII-65/2-VII-67/ 21-VI-59/ 30-VI-61/ 14-VI-62/
(D.E./D.E./ E.M.P./E.M.P./E.M.P./E.M.P./E.M.P./E.M.P./ E.J.D./ E.J.N./
E.J.N.); Tombstone Prairie, 17-VI-67/ 22-VII-61/ 22-VII-60/ 29-VI-62/ 22-VII-64/
16-VI-67/ 23-VI-61/ 19-VII-58/ 16-VIII-71/ 4-VIII-60/ 11-VIII-64, (E.M.P./
doo Bowl, S. Santiam Pass, 6-VII-68, (S.F.P.); Cascadia State Park, 13-VI-61/
23-V-59/ 10-VI-62/ 28-VI-60/ 19-V-35, (E.J.D./E.J.D./E.J.D./E.J.D./H.A.S., OSU
coll); Big Meadows, Santiam PUASSE'&IVII-56, 18-VII-48, 11-VIII-47, (R.J.A.);
Monument Peak, (?)/ 16-VII-60/ 12-VII-72, (C.W.N./F.F.H., OSU coll./E.J.D.);
Front Creek Camp, S. Santiam Hy., (?), (E.J.D.); Marion Mtn., 11-VII-64, (D.R.S.);
Trout Creek Camp, S. Santiam Hy., 19-V-40, (H.A.S., OSU coll.); 9 mi. S. Marion
Falls, Hy. 22, 22-VI-62, (K.G.); MARION CO.: Elk Lake, 6-VII-58, (S.G.J.); POLK
CO.: Valsetz, 30-V-64, (D.V.M.); Falls City, Valsetz Rd., 6-VII-64, (D.V.M.); 5
mi. S. Falls City, 17-IV-65, 11-V-65, (D.V.M.); TILLAMOOK CO.: 1 mi. E. Lee's
Camp, 18-VI-62, 16-VI-63, (E.M.P.); Bell Mtn., (?), (C.W.N.); UMATILLA CO.:
Blue Mts., N.E. Tollgate, 11-VII-64, (E.M.P.); UNION CO.: nr. Spout Spg., 15-
VII-70, (E.J.D.); WALLOWA CO.: Wallowa Lake, 29-VII-54/ 26-VII-67, (A.O.S./
E.J.D.); Ice Lake Trail, Mt. Sacajawea, 29-VIII-67, (E.M.P.); Lostine R., 15-VII-64,
28-VII-65, (E.J.D.); Hurricane Creek, 23-VII-67, (E.J.D.); Aneoid Lake Trail,
22-VII-67, (E.J.D.); Wallowa R., N. Wallowa Lake, 25-VII-53, (R.L.C.); WASCO
CO.: Bear Spgs. Camp Ground, 20-VI-62, 13-VI-64, 30-V-65, 25-VI-62, (E.M.P.);
Wapanitia, 17-VI-47, (R.J.A.); Bear Spgs., 3-VII-61, (E.J.N.); WASHINGTON CO.:
Lee Falls Rd., 2-V-70, (J.H.); YAMHILL CO.: 2.9 mi. E. Dolph, Hy.22, Siuslaw
Nat'l Forest, 18-V-68, 10-VI-62, 18-VI-62, 12-VI-63, 27-V-65, 29-V-66, (E.M.P. &
S.F.P.); 0.5 mi. E. Dolph, U.S.F.S. Rd. #S-581, Siuslaw Nat'l Forest, 18-VI-62,
12-VI-63, 27-V-65, (E.M.P. & S.F.P.); Willamina, 18-V-29, (?); Baker Creek Valley,
17-V-30, 4-VI-30, (K.M.F.); McMinnville, (?), (K.M.F.).
CHELAN CO.: Stevens Pass, (?), (V.C.); CLALLAM CO.: Hurricane Ridge, Olympic
Nat'l Park, 27-VII-67, (S.F.P.); Deer Park Camp Ground, Olympic Nat'l Park, (?),
(S.F.P.); FERRY CO.: O'Brien Creek, E. Republic, 27-VI-61, (E.J.N.); JEFFERSON
CO.: Hoh R., Olympic Peninsula, 22-VI-59, (R.L.C.); KING CO.: Stevens Pass,
24-VII-60/ 9-VIII-59, (R.E.W./V.C.); Stevens Pass, White Rock Spgs., 14-VII-30,
(E.C.V.); McBean's Peak, N.W. Snoqualmie, (?), (E.C.V.); vic. Snoqualmie Pass,
26-V-68, (D.E.); KITTITAS CO.: Morp Mtn. L.O., 13-VII-71, (E.G.); Ravens Roost
L.O., 8-VIII-71, (E.G.); Tamarack Spgs., W. Clay Elum, 7-VII-65, (E.J.N.); KITSAP
CO.: Bremerton, (?), (D.F.); KLICKITAT CO.: Bowman Creek, N.E. Goldendale,
(?), (E.J.N.); LEWIS CO.: Longmire Spgs. Wash, Ranier Nat'l Park, 27/28-VII-20,
(E.C.V.); White Pass, (?), (E.J.N.); MASON CO.: Belfair, 22-V-49, (D.F.);
OKANOGAN CO.: Salmon Meadows, 16-VII-57/ 12-VII-55/ 4-VIII-46/ 24-26-
VI-60/ 8-VII-60/ 21/22-VI-65/ 7-VII-64/ 6-VII-64, (J.C.H./J.C.H./T.W.D./
J.C.H./J.C.H./D.E./D.E./R.E.W.); Tiffany Lake, 1-VIII-37, (T.W.D.); Alta Lake
State Park, (?), (J.C.H.); Brewster, 15-VII-59/ 23-VI-60/ 7/8-VII-60/ 2-VII-44,
(J.C.H./J.C.H./J.C.H./D.E.); vic. Harts Pass, 24-VII-68, 28-VII-68, (R.E.W.);
Harts Pass, N.W. Mazama, 25-VII-62, 4-VIII-71, (E.J.N.); PIERCE CO.: Sunrise
Peak, Ranier Nat'l Park, 26-VII-36, (E.C.V.); Carbonado, 18-V-58, (D.E.); SAN JUAN
CO.: Rosario, Orcas Island, (?), (D.F.); SNOHOMISH CO.: Sloan Peak, (?),
(C.W.N.); Darrington, 4-VII-37, (ERI coll.); THURSTON CO.: Tenino, (?), (D.F.);
WHATCOM CO.: Mt. Baker Lodge, (?), (R.E.W.); YAKIMA CO.: 10 mi. E.N.E.
Chinook Pass, American R., 8-VIII-64/ 8-VII-58, (E.M.P./R.L.L.); Oak Creek,
13-VI-60, (E.J.N.); S.E. slope Mt. Adams, Bird Creek Meadows, 16/19-VII-64,
Additional records have been obtained for Butte Co., California and Lincoln
Co., Montana. Although received too late to be incorporated into the manuscript
proper, these locales are represented in Maps 1-2.
INDEX OF COLLECTORS
(R.J.A.) R. J. Albright; (R.P.A.) R. P. Allen; (R.L.A.) R. L. Anderson;
(D.H.B.) D. H. Bartholomew; (J.B.) J. Belicek; (H.M.B.) H. M. Bower; (V.C.)
V. Calkins; (W.R.C.) W. R. Carter; (R.L.C.) R. L. Chehey; (J.A.C.) J. A. Comstock;
(W.H.D.) W. H. Danby; (T.W.D.) T. W. Davies; (D.D.) D. Dirks; (E.A.D.) E. A. Dodge;
(E.J.D.) E. J. Dornfeld; (W.D.) W. Downes; (D.E.) D. Eff; (S.E.) S. Ellis; (J.F.E.)
J. F. Emmel; (P.E.) P. Engelder; (K.M.F.) K. M. Fender; (G.H.F.) G. H. Findlay;
(J.F.) J. Fletcher; (R.F.) R. Ford; (D.F.) D. Frechin; (E.G.) E. Gage; (C.B.G.) C. B.
Garrett; (A.N.G.) A. N. Gartrell; (K.G.) K. Goeden; (T.D.G.) T. D. Greyson; (J.D.G.)
J. D. Gunder; (R.G.) R. Guppy; (W.A.H.) W. A. Hammer; (F.F.H.) F. F. Hashrosek;
(G.M.H.) G. M. Hawk; (G.H.) G. Heid; (L.I.H.) L. I. Hewes; (J.H.) J. Hinchliff;
(R.H.) R. Hooper; (L.G.H.) L. G. Hudson; (O.H.) O. Huellemann; (S.G.J.) S. G. Jewett;
(S.K.) S. Kohler; (N.L.L.) N. L. LaDue; (R.L.L.) R. L. Langston; (J.L.) J. Legge;
(C.L.) C. Livingston; (H.F.M.) H. F. Madsen; (D.V.M.) D. V. McCorkle; (J.M.)
J. McDunnough; (E.M.) E. McKinney; (D.M.) D. Meadows; (A.H.M.) A. H. Moeck;
(C.W.N.) C. W. Nelson; (E.J.N.) E. J. Newcomer; (P.A.O.) P. A. Opler; (E.M.P.)
E. M. Perkins; (S.F.P.) S. F. Perkins; (C.E.P.) C. E. Peterson; (W.H.A.P.) W. H. A.
Preece; (R.H.R.) R. H. Reid; (F.H.R.) F. H. Rindge; (N.A.R.) N. A. Roman; (L.S.R.)
L. S. Rosenbaum; (J.S.) J. Schosanski; (H.A.S.) H. A. Scullen; (O.E.S.) O. E. Sette;
(J.H.S.) J. H. Shepard; (A.O.S.) A. O. Shields; (S.S.) S. Shigematsu; (C.N.S.)
C. N. Slobodchikoff; (C.I.S.) C. I. Smith; (D.R.S.) D. R. Smith; (D.S.) D. Sorg;
(R.E.S.) R. E. Stanford; (J.F.S.) J. F. Strohbeen; (A.T.) A. Tate; (D.T.) D. Threatful;
(J.W.T.) J. W. Tilden; (S.T.) S. Towns; (R.C.T.) R. C. Treharne; (A.J.U.) A. J. Urban;
(E.C.V.) E. C. VanDyke; (E.P.V.) E. P. VanDuzee; (M.L.W.) M. L. Walton; (R.L.W.)
R. L. Wescott; (N.W.) N. Westerland; (A.E.W.) A. E. Whitehouse; (F.X.W.) F. X.
Williams; (R.E.W.) R. E. Woodley; (W.G.W.) W. G. Wright; (C.H.Y.) C. H. Young;
(E.O.Z.) E. O. Zimmerman.
Museum collections are indicated as follows: (AME coll.) Allyn Museum of
Entomology, Sarasota, Florida; (AMNH coll.) American Museum of Natural
History, New York City, New York; (CAS coll.) California Academy of Sciences,
San Francisco, California; (ERI coll.) Entomology Research Institute, Canada
Department of Argriculture, Ottawa, Ontario, Canada; (LACM coll.) Los Angeles
County Museum of Natural History, Los Angeles, California; (OSU coll.) Oregon
State University, Corvallis, Oregon.
Sincere appreciation is expressed to the following contributors for their
part in loaning specimens, supplying field data and other pertinent information,
reviewing and offering suggestions relative to this manuscript, and/or the pre-
paration of photographic plates; without their able assistance and cooperation,
this work would not have been possible: Norman L. Anderson (Dept. Zoology &
Entomology, Montana State University, Bozeman), R. L. Anderson (Natural
History, Riveredge Foundation, Calgary, Alberta), Paul H. Arnaud, Jr. (Curator &
Chairman, Dept. of Entomology, California Academy of Sciences, San Francisco),
James H. Baker, Joseph Belicek (Dept. of Entomology, University of Alberta,
Edmonton), F. Martin Brown, R. H. Carcasson, Robert L. Chehey, Thomas W.
Davies, Julian P. Donahue (Assistant Curator, Dept. of Entomology, Los Angeles
County Museum of Natural History), Ernst J. Dornfeld (Dept. of Zoology, Oregon
State University, Corvallis), J. Gordon Edwards (Dept. of Entomology, California
State University, San Jose),J. Donald Eff, Scott L. Ellis, John F. Emmel, Clifford
D. Ferris (Bioengineering, University of Wyoming, Laramie), Ed Gage, John S.
Garth (Dept. of Biological Sciences, Universtiy of Southern California, Los
Angeles), Richard Guppy, Peter Herlan (Nevada State Museum, Carson City),
Charles L. Hogue (Senior Curator, Dept. of Entomology, Los Angeles County
Museum of Natural History), Richard Holland, Ronald R. Hooper, Willard E.
Ireland (Provincial Archives, Victoria, British Columbia), Steve Kohler, Noel E.
LaDue, Robert L. Langston, John Legge, George Lewis (Entomology Research
Institute, Ottawa, Ontario), Harold F. Madsen (Head, Entomology Section, Re-
search Station, Summerland, British Columbia), Sterling O. Mattoon, Edward
McMackin, Lee D. Miller (Curator, Allyn Museum of Entomology, Sarasota),
Randall Mita, Robert P. Nelson, E. J. Newcomer, Kennelm W. Philip (Institute of
Arctic Biology, University of Alaska, College), Frederick H. Rindge (Curator of
Lepidoptera, The American Museum of Natural History, New York City), A. Oakley
Shields, Steve Shigematsu, Ray E. Stanford, David Threatful, John F. Tibbs
(Director, Biological Station, University of Montana, Missoula), and Robert
Axelrod, D. I., 1957. Late Tertiary floras and the Sierra Nevadan uplift. Bull.
Geol. Soc. Amer., 68: 19-45.
Bandy, O. L., and J. A. Wilcoxin, 1970. The Pliocene-Pleistocene boundary, Italy
and California. Bull. Geol. Soc. Amer., 81: 2939-2948.
Boisduval, J. A., 1869. L6pidopter6s de la Californie. Ann. Soc. Ent. Belg., 12(50): 58.
Brown, F. M., 1951. Simple statistics for the taxonomist I. Lepid. News, 5(1-2): 4-6.
--------- 1965. The types of nymphalid butterflies described by William Henry
Edwards. Part 1. Argynninae. Trans. Amer. Ent. Soc., 91: 332-338.
Brown, F. M., D. Eff, and B. Rotger, 1957. Colorado Butterflies. Proc. Denver Mus.
Nat. Hist., Denver, 368 pp.
Comstock, J. A., 1927. Butterflies of California. Published by the author; Los
Angeles, 334 pp.
Crandell, D. R., 1965. The glacial history of western Washington and Oregon. In:
The Quaternary of the United States. (ed. by Wright, H. E., and D. G. Frey).
Princeton University Press, New Jersey, 922 pp.
Dethier, V. G., 1937. Gustation and olfaction in lepidopterous larvae. Biol. Bull.,
Dornfeld, E. J., and J. Hinchliff, 1971. Check list of Oregon Rhopalocera, with
county records and flight periods. Published by the authors; Corvallis,
Oregon, 14 pp.
Edwards, W. H., 1864. Descriptions of certain species of diurnal Lepidoptera
found within the limits of the United States and British America. Proc. Ent.
Soc. Phila., 2(5): 503-504.
------ 1868-1872. The Butterflies of North America. Vol. 1. (Argynnis VI.)
Entomological Society of America, Philadelphia, Pa., and Houghton,
Mifflin & Co., Boston, Mass.
----- 1887-1897. The Butterflies of North America. Vol. III. (Argynnis VIII.)
Entomological Society of America, Philadelphia, Pa., and Houghton, Mifflin
& Co., Boston, Mass.
Ferris, C. D., 1971. Key to the Rhopalocera of Wyoming. Science monograph #23,
U. Wyo. Ag. Expt. Sta., Laramie.
Gunder, J. D., 1924. Several new aberrant Lepidoptera (Rhopalocera). Ent. News,
------ 1926. Several new aberrant Lepidoptera (Rhopalocera). Ent. News, 37(1): 7.
(International Commission on Zoological Nomenclature), 1961. International Code
of Zoological Nomenclature. London, Internatl. Trust Zool. Nomen.:
ii-xx + 176 pp.
Munz, P. A., and D. D. Keck, 1968. A California Flora. University of California
Press, Berkeley, 1681 pp.
dos Passos, C. F., 1964. A synonymic list of Nearctic Rhopalocera. Mem. Lepid.
Soc., (1): v + 145 pp.
dos Passos, C. F., and L. P. Grey, 1947. Systematic catalogue of Speyeria (Lepi-
doptera, Nymphalidae) with designations of types and fixations of type
localities. Amer. Mus. Novitates, 1370, 30 pp.
Peabody, F. E., and J. M. Savage, 1958. Evolution of a Coast Range corridor in
California and its effect on the origin and dispersal of living amphibians
and reptiles. Zoogeography, (8): 159-186.
Perkins, E. M., and S. F. Perkins, 1966. A new race and discussion of the Boloria
epithore complex (Nymphalidae). Jour. Lepid. Soc., 20(2): 103-117.
Richmond, G. M., R. Fryxell, G. E. Neff, and P. L. Weiss, 1965. The Cordilleran ice
sheet of the northern Rocky Mountains and related Quaternary history of the
Columbia Plateau. In: The Quaternary of the United States. (ed. by Wright,
H. E., and D. G. Frey). Princeton University Press, New Jersey, 922 pp.
Shepard, J., (in press). Boloria. In: The Butterflies of North America. (ed. by Clench,
H. K.). Doubleday & Co., Inc., Garden City, N. Y.
Strand, E., 1914. Nordamerikanische, insbesondere californische Lepidoptera.
Archiv fiir Naturgeschichte, 80(A), pt. 11: 156.
Wahrhaftig, C., and J. H. Birman, 1965. The Quaternary of the Pacific mountain
system in California. In: The Quaternary of the United States. (ed. by Wright,
H. E., and D. G. Frey). Princeton University Press, New Jersey, 922 pp.
Williams, F. X., 1910. The Butterflies of San Francisco, California. Ent. News,
Wright, W. G., 1906. Butterflies of the West Coast. 2nd ed. Published by the author;
San Bernardino, 255 pp.