Title: Bulletin of the Allyn Museum
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Title: Bulletin of the Allyn Museum
Series Title: Bulletin of the Allyn Museum.
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Physical Description: v. : ill. ; 23 cm.
Language: English
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Dates or Sequential Designation: Began in 1971.
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BULLETIN OF THE ALLYN MUSEUM

Published by
THE ALLYN MUSEUM OF ENTOMOLOGY
Sarasota, Florida

Number 5 10 July 1972





NOTES ON CERTAIN SPECIES OF COLIAS
(LEPIDOPTERA: PIERIDAE) FOUND IN
WYOMING AND ASSOCIATED REGIONS'


Clifford D. Ferris
University of Wyoming, Laramie, Wyoming, and
Research Associate, Allyn Museum of Entomology


INTRODUCTION

There has been some confusion in the literature concerning three species
of Colias found in Wyoming. The taxa involved are Colias meadii Edwards,
Colias alexandra Edwards, and Colias gigantea Strecker. Subspecies of
alexandra and gigantea have been described from the state.
The purpose of this paper is to examine the distributions of these three
species of Colias in Wyoming with respect to range and subspecies. Inter-
grades within a given species are discussed and possible hybridization with
another Colias group is examined. As necessary, populations from other
geographic regions are treated.

Colias meadii Edwards
There are various literature references to Colias meadii elis Strecker from
Wyoming although I have yet to see typical specimens from the state. There
is a question in some workers' minds whether elis and meadii should be sepa-
rated at the subspecific level or considered simply as clinal forms. Several
workers have proposed separation at the species level. After examining large
series (in the hundreds) of C. meadii from Colorado, Wyoming, and Alberta,
and a limited number of specimens from Montana and Utah, I would tend to
make the separation into two subspecific taxa on the basis of habitat and
differences in facies.
Generally C. m. meadii in Colorado and Wyoming inhabits the windswept
tundra-like regions above the timberline (above 10,000') where the females
are reported to oviposit upon the alpine clover Trifolium dasphyllum Torr.

1 Published with the approval of the Director, Wyoming Agricultural Experi-
ment Station, as Journal Paper no. 467.












and Gray (Shields, et al., 1969). The butterfly is wary and possessed of swift
flight, usually quite close to the ground. Capture is not easy while the insect
is on the wing, and equally difficult when the butterfly alights because of the
protective green coloring of the under side which blends with the alpine flora.
Although basically a high mountain butterfly in the two states, I have taken
meadii in lush meadows below timberline in both Colorado (San Juan Co.) and
Wyoming (vicinity of Green River Lakes, Sublette Co., and near Brooks Lake,
Fremont Co.).
As we move north in latitude, meadii flies lower in elevation. During
July, 1970, I collected extensively in Alberta from the Montana border to just
south of Jasper. In this region C. m. elis was found flying in meadows at
quite low elevation (on the order of 6000') and well below timberline. Speci-
mens were taken in suitable meadows along most of the length of the Ka-
nanaskis Forest Trunk Road between Coleman and Seebee, on Plateau Moun-
tain (elevation 8200'), and elis was literally swarming in mid-July in the
valley through which Nigel Creek flows in Banff National Park. Flight pat-
terns and habits were quite different from Colorado and Wyoming meadii. In
Canada, elis flew relatively slowly, appeared rather unwary, and frequently
nectared upon flowers where it was easily netted. Considerable collecting on
the ridges above timberline (here only 6500'-7000' north of Banff) failed to
produce very many elis although other alpine species were taken, including
C. nastes streckeri Grum-Grschimailo. From my experiences, T must infer that
meadii is a high alpine species generally, while elis prefers lush meadow areas
generally below timberline. One must also consider the change in latitude with
respect to the two subspecies.
The males in a given population of meadii are rather uniform in facies.
The females show considerable variation in the amount of dark color which
composes the wing borders dorsally. With increasing altitude, m. meadii
tends to become smaller, the under side darker, and the upper side more dusted
in aspect. In northern Wyoming, a few females from any meadii population
will approach elis. Fig. 58 shows a typical female meadii and an extreme light
female (Fig. 59) is shown from Wyoming. A dark female (Fig. 62) and an
extreme light female elis (Fig. 63) are also shown.
Canadian elis is a larger insect than the American meadii (length of
costal margin of female forewing, 2.5 cm. for elis and 2.1 cm. for meadii, Al-
berta and Wyoming specimens) and the females show considerably less black
and more yellow than any meadii which I have seen. Both sexes are quite
pale on the under side when compared with the nominate subspecies.
Colorado and Wyoming populations produce albinic females (form
"medi" Gunder). A Wyoming specimen is illustrated (Fig. 60). Although
originally described from Colorado, female form "medi" is not common in that
state. White females are recorded from Fremont, Park, and Sublette Counties
in Wyoming. Specimens have also been taken in Carbon Co., Montana on the
Beartooth Plateau which extends into Park Co., Wyoming. One population in
Park Co., based upon limited study, may run as high as 25 per cent albino fe-
males.
C. meadii elis appears to produce albinistic females only rarely. Letters
from Dr. T. N. Freeman and Dr. A. B. Klots indicated that neither the Ca-
nadian National Collection nor the collection of the American Museum of
Natural History contains white females of elis. Dufrane (1947) named a
white female form, presumably of elis, as "lambillioni". No precise type lo-
cality is stated, only that the type specimen was from Canada "sans nom de
locality precise." A white female Colias (Fig. 64) was taken by the author on
Plateau Mountain in Alberta in 1970. Photographic reproduction fails to do
justice to detail. The specimen appears possibly to be an albino elis. It may











well be an atypical streckeri or as remote possibility elis x streckeri. In the
meadow where it was collected, both species were flying. The specimen is
quite distinct from any of the females in the series of streckeri which was
taken concurrently.
The type localities and dates of publication for the meadii group are
listed below:

Colias meadii meadii Edwards, 1871 [T. L. Mosquito Pass, Park/Lake Co.,
Colorado].
Y form "medi" (Gunder), 1934 [T. L. Breckenridge Peak, Empire Co.,
Colorado].
Colias meadii elis Strecker, 1885 [T. L. vicinity of Kicking Horse Pass,
10,000', Rocky Mtns., British Columbia, Canada].
2 form "lambillioni" Dufrane, 1947 [T. L. Canada].

Based upon my experience with meadii, I would conclude that meadii and
elis should be retained as separate subspecific taxa; elis, per se, does not occur
Wyoming; intergrades between meadii and elis occur to a limited extent in
northern Wyoming.

The Colias alexandra Complex
Three forms of C. alexandra are found in Wyoming. Before undertaking
a study of this taxon in the state, it is first necessary to devote some discus-
sion to a confusion which has developed over the years between two distinct
taxa. Both taxa, C. astraea and C. harroweri, were originally described from
Wyoming. Their principal species associations are noted below.

Colias alexandra astraea Edwards
In recent years, astraea has been placed with the C. scudderii complex
(dos Passos, 1964), most probably as a result of a paper by Klots (1937). In
this paper, Klots recorded a collecting trip made in 1935 through New Mex-
ico, Colorado, and Wyoming. On page 321 he notes: "A considerable number
of C. astraea were seen, and a small series taken, in the valley of Clear Creek
just below the 'Natural Bridge', near the head of the lower Green River Lake
on August 3." This location is in the Bridger Wilderness in Sublette Co.,
Wyoming. Klots subsequently realized that this butterfly was not astraea
and he named it harroweri (1940). In the 1937 paper, reference was made to
Salix as the possible food plant for "astraea" (= harroweri). This reference
has probably aided in the misapplication of astraea to the scudderii complex.
William Henry Edwards described C. astraea in March, 1872 based on a
single male taken by the Hayden Expedition of 1872 (Figs. 83-84). The type
locality is Yellowstone Lake, (Yellowstone National Park) Wyoming. Ed-
wards noted at the end of his description: "On the under side this species is
nearest Alexandra; on the upper side of a different shade of color from any
of our species."
Two key points of Edwards' description are the following excerpts:
"Upper side pale ochraceous, very little tinted with orange on the disks of
secondaries from cell to marginal border and from base to hind margin of
secondaries below the cell, this color being not decided but only a tint; ...
discal spot of primaries a short black streak; on secondaries wanting."
"Under side of primaries yellow nearly as above, without orange; of sec-
ondaries yellow densely covered with black scales so as to obscure the whole
surface; discal spot of primaries very narrow, black, enclosing a few yellow
scales, of secondaries white, as in Alexandra, without a ring; ...."











The specimen which Edwards had before him would appear fo be not
quite typical of the insect which flies in northern Wyoming thence to Alberta.
Specimens fitting the description occur sparingly and I have in my collection
one from Wyoming and one from Alberta. When a series of "typical" astraea
is examined, one notes a distinct discal spot on the upper surface of the sec-
ondaries. There are two extremes in dorsal color of the wings from heavily
orange tinted to nearly pure yellow. In the former case, the discal spot on
the upper side of the hind wing is obscured by the ground color (probably the
situation in the type specimen). In the latter case, it appears as a pale but
distinct orange spot. On the under side of the hind wing, the discal spot is
usually finely ringed with pink scales. This is quite different from the situa-
tion in harroweri described below.
In other respects, astraea resembles alexandra in the ventral color and
the sharp apices of the primaries. The females are white. Ventrally they are
similar to the males except for lighter ground color. The secondaries exhibit
a greenish overscaling. Dorsally, the margins are lightly black bordered in
a pattern suggesting eurytheme, but much more lightly scaled. In some ex-
amples, the dark scaling is virtually absent. The secondaries may or may not
show dark border scaling. The discal spot varies from white to pale pink-
orange to orange.

Colias gigantea harroweri Klots
This taxon was originally placed by Klots in the combination shown.
Based upon food plant association, habitat similarity, and other considera-
tions, various workers have adopted scudderii as the species. Some discussion
of this point will be found in a later section of this paper.
In his description of harroweri, Klots (1940) recognized the astraea-
gigantea confusion, as noted in his opening statement: "For many years this,
as well as other yellow Colias from western North America, has been passing
under the name Colias christina [now C. alexandra christina] form astraea
Edwards." He then continues: "A small series taken by the author in 1935
(now a part of the present type lot) was thus tentatively named (Journ. N. Y.
Ent. Soc., 1937, XLV, p. 321)." Here Klots corrected the earlier misidentifica-
tion of harroweri as astraea. The description of harroweri is quite detailed
and needs no embellishment. The key points are as follows: Males Apices of
primaries definitely rounded; dorsal ground color clear lemon yellow with
greenish tinge on secondaries; discal spot of secondaries "shows through"
from under side as a pale orange spot; ventral ground color similar to above
but lighter with some dark overscaling along costa of primaries and lightly
on disc of secondaries; secondary discal spot large and distinct, usually with
satellite and frequently with full double spot; spot or spots pearly, heavily
ringed with dark pink-brown scales which frequently invade the spot center;
satellite spot often totally dark. Females Dorsal ground color bright yellow
with orange tinting toward body; black border absent or present only as a
trace; discal spot of secondaries bold, double, and orange; ventrally the nature
of the coloring resembles that of the males, but with the base color of the fe-
males. White females are occasionally found in Wyoming, and become more
common in regions north of the state.
C. alexandra astraea and C. gigantea harroweri are quite distinct from
one another. In some areas, the two species are sympatric and synchronic
(vicinity of Slide Falls and Clear Creek, Bridger Wilderness, Sublette Co.,
Wyoming) which perhaps has caused some of the confusion. This area is
the type locality for harroweri. The term sympatric is used here in the gen-
eral sense, as harroweri is a denizen of the wettest portions of sphagnum











bogs in which its food plant (Salix) grows (occasionally a female is found
nectaring at the bog edge), while astraea is found on dry land in grassy
clearings in the forest, frequently in association with sagebrush and its sus-
pected food plants in the family Leguminosae, Astragalus and Lupinus.
The specimens of C. astraea figured as types by Holland in the Butterfly
Book (Plate LXVIII, Figs. 26, 27, 1931 Edition) and illustrated in Figs. 81,
82, 85, 86, are not the types and at best represent atypical astraea. Klots re-
ferred to this situation in his 1940 paper. The male specimen which Holland
figured shows "eurytheme" spotting on the under side of the secondaries and
a partial satellite discal spot which is not characteristic of astraea. The type
specimen of astraea (Figs. 83, 84) does not exhibit "eurytheme" spots.
I have seen specimens of C. alexandra astraea from Wyoming, Utah,
Idaho, Montana, and southern Alberta. C. gigantea harroweri is recorded
from Wyoming, Montana, Idaho, and southern portions of Manitoba, British
Columbia, and Alberta. Sorensen (unpublished) has shown that specimens
from southern Manitoba are distinct harroweri as has Masters (1970a). The
Chermocks (1940) applied the name mayi to the Riding Mountains, Manitoba
population. Sorensen's work indicates that mayi, if a synonym, should be
considered synonymous with gigantea rather than harroweri as listed by dos
Passos (1964). Masters has chosen to consider mayi a valid taxon. Northern
Canadian specimens are referred to C. g. gigantea as is the Alaska popula-
tion. The ranges of the various forms of gigantea overlap considerably and
appear to represent a clinal situation. It remains to be seen whether only the
end points of the dine should carry subspecific names, or if additional sub-
specific names should be applied. The ranges of C. alexandra and C. gigantea
overlap considerably, but these species are distinct in their morphology and
their food plant associations. Their habitats are different in that gigantea is
a bog species and alexandra a dry land species. Positive separation can be
achieved by ultraviolet photography of doubtful individuals. Under ultraviolet
illumination, C. alexandra males exhibit a luminous patch (actually an inter-
ference pattern) on the dorsal surface of the secondaries; C. gigantea shows no
luminosity. For several years a series of Colias from southern British Colum-
bia and northern Idaho has resided in the author's collection as giganteaa".
These are large insects and generally resemble gigantea except that they were
collected in forest clearings rather than in bogs. Ultraviolet photography has
shown them to be alexandra. Detailed studies will be published in the near
future.
Both Klots (1940) and McDunnough (1922, 1928) have called attention
to the differences between the alexandra-astraea-christina (alexandra) com-
plex and the gigantea-scudderii complex. Apparently these works have been
overlooked in recent years by some workers.
To understand the alexandra populations found in Wyoming, it is neces-
sary to examine superficially the entire alexandra complex as it is presently
known. Three population divisions can be made with respect to dorsal color-
ation in the males: entirely yellow, entirely orange, mixed yellow and orange.
The taxa associated with these color divisions, their type localities, and dates
of publication are:

Yellow Populations
Colias alexandra alexandra Edwards, 1863 [T. L. Front Range, West of
Denver, Colorado].
Colias alexandra edwardsii Edwards, 1870 [T. L. Virginia City, Storey Co.,
Nevada].
Colias alexandra emilia Edwards, 1870 [T. L. Oregon].
Colias alexandra ssp. Arizona New Mexico segregate.












Orange Populations
(?) Colias alexandra alberta Bowman, 1942 [T. L. Wembley, Alberta, Can-
ada]. See discussion which follows below concerning placement of
alberta.
Colias alexandra krauthii Klots, 1935 [T. L. Black Hills, 12 miles west of
Custer Co., South Dakota].
Colias alexandra ssp. Manitoba segregate. Beulah, Manitoba and other
areas. Orange population perhaps referable to krauthii (?).

Yellow-Orange Populations
Colias alexandra astraea Edwards, 1872 [T. L. Yellowstone Lake, Wyo-
ming].
Colias alexandra christina Edwards, 1863 [T. L. Slave River Crossing, N.
W. T., Canada].

The yellow populations of alexandra range from the Pacific Northwest
(Oregon) south through California and east across the Great Basin and into
Arizona, New Mexico, and Colorado. Apparently undiluted yellow forms are
found in western Nebraska, and in the southern portions of Utah and Wyo-
ming. Generally speaking, emilia is associated with the Pacific Northwest,
edwardsii with eastern California and the Great Basin, and alexandra with
Colorado and the Front Range into Wyoming. The Arizona New Mexico
segregate from the Mogollon Rim through the White Mountains in Arizona
and into the Mogollon Mountains in New Mexico is interesting. It appears
to be distinct from the named subspecies. The insects are quite large and the
females very pale in color. This segregate appears closest to edwardsii. This
is true also of some southern Utah specimens.
The orange form krauthii occurs in western South Dakota and north-
eastern Wyoming (Black Hills region). Another orange form, alberta, was
described by Bowman from Canada. This taxon is the subject of a separate
discussion in a later section of this paper.
The geographic region defined by the Pacific Northwest into northern
Utah and Wyoming on the south, through Montana and Idaho into Alberta
and east to Manitoba encompasses the range of the yellow-orange populations
of alexandra.
When we examine the yellow-orange populations, it is frequently difficult
to attach the name astraea or christina to given specimens. Wyoming ma-
terial from Sheridan Co. west to Yellowstone National Park generally fits
Edwards' description of astraea with reference to the males. Based upon
specimens which I have seen and taken in Wyoming, I would arbitrarily as-
sign females which exhibit an untinted white dorsal ground color to this
taxon.
Northern Utah specimens (Salt Lake, Summit, Tooele, Wasatch Cos.)
tend toward both astraea (in the males) and christina (in the females). The
females frequently show an overwashed orange coloration. Idaho and Mon-
tana specimens represent both forms. Material from southern Alberta south
of Banff National Park exhibits the complete range from pure yellow alex-
andra-like specimens to pure orange krauthii-like specimens in the males. Fe-
males from this region range from pure white dorsally to pale orange-yellow
as in christina. I have not seen specimens with as dark an orange as in
krauthii females from the Black Hills. The name christina has been applied
to specimens taken from the Northwest Territories to the Riding Mtns. in
Manitoba.











There appears to be one characteristic in the males of the alexandra com-
plex which can be used reliably to separate the populations. This is the
dorsal discal spot on the secondaries. The original descriptions of the taxa
associated with the yellow populations note that this spot is absent (hence
concolorous with the yellow of the wings). In the orange and the yellow-
orange populations, this spot is orange, although sometimes masked by the
orange color of the wing. Extreme examples of astraea, although appearing
pure yellow, will show the orange discal spot and may thus be separated from
alexandra alexandra and other yellow populations.
It would appear that "emilia" from north-central Washington (Okanogan
Co.) intergrades with the yellow-orange populations to the east. I have seen
specimens which fit the original description of emilia in all respects save that
the discal spot is orange. Specimens of alexandra from southern British Co-
lumb;a and northern Idaho, alluded to previously, are quite large and show an
orange discal spot in the males. They are quite distinct from astraea. F. M.
Brown is currently preparing a paper which treats "emilia" and edwardsii, as
part of his continuing work on the Edwards types. The alexandra complex
will be treated in detail in a subsequent paper.
Females of the alexandra complex are quite variable and range from
white through yellow to orange. The quantity of dorsal dark bordering
varies from lacking to prominent. The discal spot separation technique is
not reliable with this sex. Normally members of the yellow populations have
the spot absent or appearing lighter than the wing color. Occasionally the
spot is pale orange. Females of the orange and yellow-orange populations
normally exhibit an orange spot, if the females are yellow or orange, but the
spot may be almost white (or sometimes a very pale pink) in white females.
The ventral surface of the males and to a lesser extent in the females is
usually diagnostic. Members of the yellow populations exhibit a cold gray-
greenish color on the secondaries. This is produced by a heavy dusting of
dark scales over the yellow base color. In the orange and yellow-orange
races, the secondaries generally show a warm orange overtone, although
heavy dark dusting may be present. Ultraviolet photography is diagnostic
for males of alexandra.
In western Wyoming and northern Utah, a diminutive alexandra is found.
It is not clear whether this is an altitudinal form or genetically related. The
specimens are considerably smaller than normal alexandra alexandra and the
dorsal black borders of the males are very narrow (as in emilia). The discal
spot is concolorous with the yellow of the wings.
C. alexandra is usually associated with relatively open areas, either sage-
brush zones (frequently in aspen-pine zone clearings) or forest clearings. The
food plant is generally thought to be the Leguminosae (Lupinus and Astrag-
alus) and has been proved in certain instances (McDunnough, 1922; Shields,
et al., 1969). The larvae of the yellow-orange population in Utah pupate at
the bases of sagebrush plants, generally in the debris at ground level (Tid-
well, 1970).

Colias alberta Bowman
Bowman, when he described C. alberta, placed the taxon with C. eury-
theme in the combination Colias eurytheme alberta Bowman (1942). Hovanitz
(1943) in a brief discussion assigned alberta to alexandra and dos Passos
(1964) followed. The position and validity of this taxon has been in question
ever since. Some workers have stated that Bowman simply re-described C.
alexandra christina; others feel that C. eurytheme was re-described. Recently,
Masters (1970b), based upon examination of a single paratype, has unequivo-











cally stated that alberta is C. alexandra christina. The author has not been
able to ascertain what specimens Hovanitz examined in preparation for his
1943 paper.
According to the International Code of Zoological Nomenclature (Ar-
ticles 72 and 73), when a holotype is designated, as Bowman did, that
specimen is the reference for the taxon. Thus one runs a risk in making pro-
nouncements based upon examination of paratypes or single specimens within
the type series.
Recently I was able to locate the type specimens of alberta in the ento-
mological collection of the University of Alberta at Edmonton. These are
figured on the color plate. In addition, seven paratypes from the original
series were examined as well as the holotype and one paratype of the female
form "pall;d;ssima" Bowman. Photographs of additional paratype material
from the Los Angeles County Museum of Natural History were also examined.
The type specimens have been placed in the Canadian National Collection.
The seven paratypes of alberta and the holotype and paratype of "pallidis-
sima" are figured (Figs. 7-24.)
Based upon careful examination of this material and comparison of it
with populations of C. eurytheme and C. alexandra, it would appear that al-
berta represents a hybrid situation and is not really a valid taxon.
The maculation of the upper surface is characteristic of C. eurytheme
Boisduval in the following respects: males- ground color, width and dentation
of dark margin (margin about half again as wide as in alexandra), size,
shape and color of both primary and secondary discal spots. Based upon ex-
amination of the upner surfaces only, one would place alberta with eurytheme
as Bowman did. The white female forms, pallidissima, are not unlike the
white females of C. philodice vitabunda Hovanitz; they bear little resemblance
to the white female forms in the alexandra complex.
The facies of the under surface of alberta are generally characteristic of
eurytheme, but with some exceptions, as shown in Figs. 3-24. Figures 3-6
illustrate typical eurytheme from the Rocky Mountain regions. These two
specimens were selected for their similarity to the holotype and allotype
(dorsal surface) of alberta. The overall coloring of the under sides of alberta
specimens is lighter than eurytheme from the Middle Atlantic Region and the
spotting is not so strongly defined. Three characteristics of eurytheme are
present. These are submarginal spots on either the primaries or secondaries,
or both, or satellite discal spot to main discal spot on the secondaries. The
"eurytheme" spots are generally present to some degree, although very pale
to the point of being almost absent in some of the material examined, es-
pecially in the holotype and allotype. The discal spot on the secondaries is
light and pearly or almost silvery as in eurytheme. The submarginal row of
spots on the secondaries, although generally light, is typical in its form to
eurytheme. The pink fringes on the wings are again more typical of eury-
theme than alexandra.
Actually the specimens are not very different from the forms of eurytheme
taken at higher elevations in Colorado and Wyoming. Wyoming material, in
particular, is frequently pale on the under surfaces with spotting almost ob-
solete. If we equate altitude to latitude, as can sometimes be done with re-
spect to maculation, we would expect lighter spotting in northern popula-
tions of eurytheme.
The specimens shown in Figs. 7-14 have been photographed also under
ultraviolet illumination. The interference patches which occur are closer in
form to those produced by eurytheme than those produced by alexandra.
Patches occur on both the primaries and secondaries as is the case with the
orange races of alexandra.











In view of the preceding discussion, I feel that alberta represents a hy-
brid situation and th ename should be suppressed rather than sunk in synonymy
with another taxon. The hybrid would appear to be C. alexandra christina
x C. eurytheme, and possible backcross generations. Populations of C. philo-
dice eriphyle Edwards are sympatric with christina, but it is felt that this is
not the hybridizing species. The philodice group does not exhibit ultraviolet
interference patterns while eurytheme does. This situation would also indi-
cate that philodice and eurytheme are separate species, although hybridiza-
tion does appear to occur naturally. These problems are currently being
studied by 0. R. Taylor, J. T. Sorensen, and R. Silberglied. P. A. Opler (un-
published) has placed eurytheme as a subspecies of philodice.
There is considerable evidence for natural hybridizing between eurytheme
and members of the alexandra complex. Some discussion is presented in sub-
sequent portions of this paper. Dr. S. A. Ae (Japan) has done studies on
laboratory hybrids between C. eurytheme and C. interior, C. philodice and C.
alexandra.

The Colias scudderii-gigantea Complex
The exact relationships of the members of this complex is not completely
clear. The Wyoming taxa involved are Colias gigantea harroweri Klots,
Colias pelidne skinneri Barnes, and Colias s. scudderii Reakirt. Klots (in
Ehrlich and Ehrlich, 1961) provisionally combined the taxa pelidne and scud-
derii and indicated that these may be conspecific with gigantea. dos Passos
(1964) lists gigantea and scudderii as conspecific and pelidne separately. It
has been demonstrated that the larvae of all three feed upon Salix (Klots in
Ehrlich and Ehrlich, 1961 ; Klots 1937, 1940 ; Shields, et al., 1969). I do not
feel that a common food plant genus is sufficient evidence for conspecificity.
C. pelidne skinneri and C. gigantea harroweri are sympatic in Wyoming. The
range of C. s. scudderii does not appear to overlap the ranges of the other two
species.
My inclination is to treat pelidne as a species distinct from gigantea and
scudderii. Based upon one seemingly consistent character, I would also sepa-
rate gigantea and scudderii, although they appear very closely related. This
character is the discal spot on the ventral surface of the secondaries. Only
very rarely in scudderii does this show a suggestion of a satellite spot. In
gigantea, this spot invariably shows a trace of a satellite spot (except in a
very few far northern specimens) and generally the spot is distinctly double,
as shown in Figs. 73-78, 87, 88.
There is one habit difference between gigantea and scudderii in Wyoming:
gigantea harroweri is distinctly a deep bog species and seldom strays very
far from the center of its habitat; scudderii on the other hand may range
quite far from willow bogs. Both s. scudderii and pelidne skinneri may be
found along forest roads and frequenting open grassy meadows, although
these two species inhabit different areas of the state.
In Wyoming, the females of all three species are dimorphic, being either
yellow or white. The dominant female form for scudderii is white, for har-
roweri yellow, and about equally divided between yellow and white for skin-
neri. In northern Canada, gigantea females tend toward white rather than
yellow.
Based upon the previous discussion, I would be inclined to group these
Salix feeders as follows:

Colias scudderii scudderii Reakirt, 1865 [T. L. probably near Empire,
Clear Creek Co., Colorado].
Colias scudderii ruckesi Klots, 1937 [T. L. Windsor Creek Canyon, west












of Cowles, San Miguel Co., New Mexico].
Colias gigantea gigantea Strcker, 1900 [T. L. west coast of Hudson Bay
above Ft. York, now taken as Churchill, Manitoba, Canada. For the
present, mayi is lumped here.
Colias gigantea harroweri Klots, 1940 [T. L. vicinity Clear Creek and Slide
Falls, Sublette Co., Wyoming].
Colias pelidne group. Four subspecies listed by dos Passos 1964.
Colias pelidne skinneri Barnes, 1897 [T. L. Yellowstone National Park,
Wyoming].

Dr. A. B. Klots has raised a question that the Alaska population of gi-
gantea might be quite different from the Churchill, Manitoba (type locality)
population. Compare the specimen shown in Figs. 73-74 with the specimen
shown in Figs. 87-88. Note the double discal spot on the underside of the
Alaskan specimen.
Further research may well show that the taxa gigantea, pelidne, and scud-
derii should all be grouped under pelidne which bears publication priority.
Morphology would incline me to separate gigantea at least from the other
two taxa as was suggested by Klots (in Ehrlich and Ehrlich, 1961).

Possible Hybridization
Genitalic studies and chromosome examination (Kodo and Remington,
1960) indicate that from a structural standpoint, hybrids are possible be-
tween various species of Colias. Field collected specimens tend to bear this
out. Several specimens which appear to be hybrids are figured in the plates.
Various subspecies of alexandra consistently show "eurytheme" spots (sub-
marginal row) ventrally on the secondaries and occasionally a satellite spot
to the discal spot. This is especially true in populations of astraea, christina,
and krauthii: the orange and yellow-orange populations. C. alberta has been
treated previously. It is interesting to note that the alexandra complex and
C. eurytheme have members of the Leguminosae as larval foodplants. C.
eurytheme is an aggressive species and one might expect mating attempts
with other species. Studies are being conducted by O. R. Taylor, S. A. Ae
and others.
The author has undertaken extensive ultraviolet photographic studies of
the genus Colias to determine which species have similar appearance under
ultraviolet illumination. These studies will be published subsequently. Sev-
eral species do appear similar under ultraviolet illumination which does give
support to hybridization attempts.
The possible hybrid situations for Wyoming species are presented. Fig.
46 shows a male C. alexandra krauthii with typical under side "eurytheme"
spots. Fig. 48 shows a probable krauthii x eurytheme specimen. The under
side of this female is typical of alexandra, but the upper side resembles eury-
theme. Figs. 79-80 show a probable cross between meadii and pelidne skinneri.
The dark border and general facies resemble pelidne, but the color of the dorsal
surface is orange as in meadii.

Distribution in Wyoming
Figure 1 shows the probable general distribution of the alexandra com-
plex with designations for the various color populations. The distribution
will be treated in greater depth in a subsequent paper. Figure 2 illustrates
the distributions in Wyoming for the various species mentioned in this paper.











ACKNOWLEDGEMENTS

The author is indebted to a number of people who aided in this effort.
F. M. Brown and Dr. A. B. Klots kindly read and commented upon a prelim-
inary draft of this paper. The author carried on extensive correspondence
with them. Drs. D. F. Hardwick and T. N. Freeman supplied information on
material in the Canadian National Collection. Mr. George Lewis of the Ento-
mological Research Institute, Ottawa, Ontario, Canada kindly provided the
color transparencies used in making the color plate. Dr. G. E. Ball and D. R.
Whitehead of the University of Alberta arranged for the loan of material
from the Bowman Collection at the University. Mr. Julian P. Donahue gra-
ciously provided photographs of alberta paratypes in the Los Angeles County
Museum of Natural History. Dr. H. G. Rodeck provided access to the Univer-
sity of Colorado Museum Collection.
J. R. Heitzman provided numerous specimens for examination and pho-
tography. K. B. Tidwell supplied additional specimens of Utah Colias alex-
andra. J. D. Eff provided numerous specimens for examination. Dr. John
S. Nordin provided specimens for examination, Wyoming records, and infor-
mation on collecting sites. Additional collection data was supplied by D. R.
Groothuis and J. R. Mori.
P. J. Conway and J. T. Sorensen supplied Manitoba material for exami-
nation. J. A. Ebner supplied material from northern Canada, W. V. Krivda
sent material from The Pas, Manitoba, and Dr. R. E. Woodley provided ma-
terial from the Pacific Northwest.
F. M. Brown kindly provided the photographs of the type specimens and
other material in the Carnegie Museum Collection. Dr. A. B. Klots was most
helpful in providing specimens for photography from his collection, and col-
lection records from specimens in the collection of the American Museum of
Natural History.
I am also indebted to the Bureau of Indian Affairs, Ottawa, the Park Su-
perintendent, and the Park Naturalist of Banff National Park for permission
to collect material in the Park. The main studies of Colias meadii elis were
conducted in Banff National Park.
Additional material used in this study was drawn from the literature.
The various papers examined will be found in the "Literature Cited". The
author would like to express his appreciation to Dr. Lee D. Miller and Mr.
A. C. Allyn for making publication of this paper possible.

LITERATURE CITED

Barnes, W., 1897. Some new species and varieties of Lepidoptera from the
western U. S. Can. Ent., 29: 39-42.
Bowman, K., 1942. A note on Colias eurytheme Bdv., with description of a
new race (Lepidoptera, Pieridae). Can. Ent., 74: 25.
Chermock, F. H. and R. L. Chermock, 1940. Some new diurnal Lepidoptera
from the Riding Mountains and the Sand Ridge, Manitoba. Can. Ent.,
72: 81-83.
Dufrane, A., 1947. Pieridae. Bull. Ann. Soc. ent. Belgium, 83: 46-73.
Edwards, W. H., 1863. Description of certain species of diurnal Lepidoptera
found within the limits of the United States and British America. (1),
Proc. Ent. Soc. of Phila., 2: 14-22. No. (2), Proc. Ent. Soc. of Phila., 2:
78-82.
1870. Descriptions of new species of diurnal Lepidoptera found within
the United States. Trans. Amer. Ent. Soc., 3: 10-22.
1871. Descriptions of new species of North American butterflies.
Trans. Amer. Ent. Soc., 3: 266-277.












1872. Descriptions of new species of diurnal Lepidoptera found
within the United States. Trans. Amer. Ent. Soc., 4: 61-70.
Ehrlich, P. R. and A. H. Ehrlich, 1961. How to know the butterflies. Dubuque,
W. C. Brown Co.: intro.+262 pp.
Gunder, J. D., 1934. Various new butterflies (Lepid., Rhopalocera). Can.
Ent., 66: 125-131.
Holland, W. H., 1931. The butterfly book. New York, Doubleday and Co.:
xii + 424 pp.; ill.
Hovanitz, W., 1943. The nomenclature of the Colias chrysotheme complex in
North America. (Lepidoptera, Pieridae). American Mus. Novitates, 1240.
- 1950a. The biology of Colias butterflies. I. The distribution of the
North American species. Wasmann J. Biol., 8:49-74.
-- 1950b. The biology of Colias butterflies. II. Parallel geographical
variation of dimorphic color phases in North American species. Wasmann
J. Biol., 8: 197-219.
1951. The biology of Colias butterflies. III. Variation of adult flight
in arctic and subarctic. Wasmann J. Biol., 9: 1-9.
Klots, A. B., 1935. A new Colias from South Dakota (Lepidoptera: Pieridae).
American Mus. Novitates, 767.
-- 1937. Some notes on Colias and Brenthis (Lepidoptera, Pieridae and
Nymphalidae). J. N. Y. Ent. Soc., 45: 311-333.
1940. New butterfly subspecies from Wyoming (Nymphalidae, Pie-
ridae). American Mus. Novitates, 1054.
1951. A field guide to the butterflies. Boston, Houghton Mifflin Co.:
xvi + 349 pp. ill.
--- 1961. in Ehrlich, P. R. and A. H. Ehrlich, op. cit.
Kodo, M. and C. L. Remington, 1960. Studies of the chromosomes of North
American Rhopalocera. 2. Hesperiidae, Megathymidae, and Pieridae.
J. Lepid. Soc., 14: 35-37.
Masters, J. H., 1970a. Concerning Colias christina mayi Chermock & Cher-
mock. J. Res. Lepid., 9 (4): 227-232.
- 1970b. Concerning Colias eurytheme alberta Bowman (Pieridae). J.
Res. Lepid., 9 (2) : 97-99.
McDunnough, J., 1922. Notes on the Lepidoptera of Alberta. Can. Ent.,
54: 134-141.
-- 1928. Notes on Canadian diurnal Lepidoptera. Can. Ent., 60: 266-
275.
dos Passos, C. F., 1964. A synonymic list of Nearctic Rhopalocera. Memoir
No. 1. The Lepidopterists' Society: vi+145 pp.
Reakirt, T., 1865. Observations upon some American Pierinae. Proc. Ent.
Soc. of Phil., 4: 216-222.
Remington, C. L., 1954. The genetics of Colias (Lepidoptera). In Advances
in Genetics, Vol. VI. New York, Academic Press: pp. 403-450.
1956 (1958). Genetics of populations of Lepidoptera. Proc. Tenth
Int. Congr. of Ent., 2: 787-806.
Shields, O. et. al., 1969. Butterfly larval foodplant records and a procedure
for reporting foodplants. J. Res. Lepid., 8 (1): 21-36.
Strecker, F. H. H., 1885. Description of a new Colias from the Rocky Mtns.,
and an example of polymelianism in Samia cecropia. Proc. Acad. Nat.
Sci. Phila., 37: 24-27.
1900. Lepidoptera, Rhopalocera and Heteroceres, indigenous and
exotic. Supplement No. 3. Reading, Penna. 4 to., pp. 13-27.
Tidwell, K. B., 1970. Personal communication.






13




















































Color plate. Colias "alberta" Bowman; Holotype a (upper figures) and
Allotype 9 (lower figures). Both specimens are from Wembley, ALBERTA,
CANADA. These specimens are now placed in the Canadian National Col-
lection. Color transparencies by George Lewis.















* 0


Fig. 1. Distribution of Colias alexandra in North America south of the
60th parallel. Open circles=yellow populations, solid dots orange popula-
tions and half-open circles =yellow-orange populations. Dotted line is Con-
tinental Divide. Solid dots in Alberta and British Columbia refer to C. al-
berta (see discussion in text). Only subspecies of C. alexandra as listed in
dos Passos (1964) are indicated. Compare with Hovanitz (1950a: fig. 6).







15














N.P.e .) 0 .


I '^ .-^ ; ^ *.--- --. ;
------------
r--'-
,-. - - -- - - -



SH" Srg
t. 0d L-., ~---^------- 1-------^--------- ------





n-- -i A L



I Io .
S. a ie


I AlbOy"
d. f
















Fig. 2. Distribution of certain Colias species in Wyoming. Solid dots=
C. a. alexandra, open circles= C. alexandra krauthii, vertical half-open circles
=C. alexandra astraea, horizontal half-open circles with solid bottom= C. m.
meadii, horizontal half-open circles with solid top = C. m. meadii 9 form medi,
open circles with diagonal line= C. pelidne skinneri, open triangles = C. s. scud-
derii and solid triangles C. gigantea harroweri. All county records to date
are indicated; individual collection sites within a county are not shown.










i
I ;
I
c I
r


8 9








12 13


--a


U.-


A1


- 4 /
K I


.--"
9-
a


10
-C~ "


15 16 17 18
Figs. 3-18: Colias. 3-4, C. eurytheme, S upper (3) and under (4) sur-
faces; MONTANA: Sanders Co. 5-6, Same, 9 upper (5) and under (6) sur-
faces; NEW MEXICO: Grant Co. 7-8, C. "alberta", Paratype S upper (7)
and under (8) surfaces; ALBERTA: Wembley. 9-10, Same, Paratype &
upper (9) and under (10) surfaces; ALBERTA: Wembley. 11-12, Same,
Paratype S upper (11) and under (12) surfaces; ALBERTA: Fort Vermilion.
13-14, Same, Paratype 3, upper (13) and under (14) surfaces; ALBERTA:
Beaverlodge. 15-16, Same, Paratype 9, upper (15) and under (16) surfaces;
ALBERTA: Wembley. 17-18, Same, Paratype 9, upper (17) and under (18)
surfaces; ALBERTA: Wembley.


1 J

Pair.-













20 21


19


24 25


28 29


* A.


ft.


r`


'ipA
1 r


31 32 33 i 34
Figs. 19-34: Colias. 19-20, C. "alberta", Paratype 9 upper (19) and
under (20) surfaces; ALBERTA: Beaverlodge. 21-22, C. "pallidissima", Par-
atype 9 upper (21) and under (22) surfaces; ALBERTA: Fort Vermilion.
23-24, Same, Holotype 9 upper (23) and under (24) surfaces; ALBERTA:
Fort Vermilion. 25-26, C. a. alexandra, S upper (25) and under (26) sur-
faces; WYOMING: Albany Co. 27-28, Same, 9 upper (27) and under (28)
surfaces; WYOMING: Albany Co. 29-30, C. alexandra christina, & upper
(29) and under (30) surfaces; ALBERTA: Bow River Forest. 31-32, Same,
9 upper (31) and under (32) surfaces; ALBERTA: Bow River Forest. 33-
34, C. alexandra ssp., & upper (33) and under (34) surfaces; UTAH: Tooele
Co.


t


'4


I I


23






























Figs. 35-50: Colias. 35-36, C. alexandra ssp., 9 upper (35) and under
(36) surfaces; UTAH: Wasatch Co. 37-38, C. alexandra ssp. (yellow form with
orange discal spot), a upper (37) and under (38) surfaces: BRITISH CO-
LUMBIA: nr. Golden. 39-40, Same, 9 upper (39) and under (40) surfaces;
BRITISH COLUMBIA: nr. Golden. 41-42, C. alexandra christina, 9 with
"eurytheme" under surface spots, upper (41) and under (42) surfaces; AL-
BERTA: Bow River Forest. 43, C. alexandra krauthii, & upper surface;
SOUTH DAKOTA: Lawrence Co. 44, Same, 9 upper surface; SOUTH
DAKOTA: Lawrence Co. 45-46, Same, under surfaces of two $ ; SOUTH
DAKOTA: Lawrence Co. 47, C. alexandra astraea, & upper surface; WY-
OMING: Sheridan Co. 48, C. alexandra krauthii x C. eurytheme hybrid
(?), 9 upper surface; SOUTH DAKOTA: Lawrence Co. 49-50, C. alex-
andra astraea, & upper (49) and under (50) surfaces; WYOMING: Sublette
Co.








I '~
F'
1~


36 37


4-!
em,
'1"


40 41


44
1!


dV-"

*'4ix


t
I
(9


i;g


39





43





47


I //


CI




























Figs. 51-66: Colias. 51-52, C. alexandra astraea, 9 upper (51) and
under (52) surfaces; ALBERTA: Bow River Forest. 53-54, Same, 2 upper
(53) and under (54) surfaces; WYOMING: Johnson Co. 55-56, Same, 2
upper (55) and under (56) surfaces; ALBERTA: Bow River Forest. 57,
C. m. meadii, S upper surface; WYOMING: Albany Co. 58, Same, 2 upper
surface; WYOMING: Albany Co. 59, Same, 2 upper surface; WYOMING:
Fremont Co. 60. Same, form "medi", ? upper surface; WYOMING: Fre-
mont Co. 61. C. meadii elis, 8 upper surface; ALBERTA: Plateau Mtn.
62, Same, 9 upper surface; ALBERTA: Plateau Mtn. 63, Same, 2 upper
surface; ALBERTA: Plateau Mtn. (light form). 64, Same, possibly form
"lambillioni", 2 upper surface; ALBERTA: Plateau Mtn. 65-66, C. cunning-
hami Butler, S upper (65) and under (66) surfaces; erroneously labelled
"Vic Estes Park, Larimer Co., COLORADO"; probably from either Ecuador
or Peru.
























r
51' 0 52 53 54







55 56 57 58






59 60 61 62



A; *. i..


64 65


, .1








C

(


.7


19


72 73


76 77


ID'


r ~

S.


Figs. 67-80: Colias. 67-68, C. cunninghami, 9 upper (67) and under (68)
surfaces; data as in Figs. 65-66. 69-70, C. s. scudderii, 8 upper (69) and
under (70) surfaces; WYOMING: Albany Co. 71-72, Same, 9 upper (71) and
under (72) surfaces; WYOMING: Carbon Co. 73-74, C. g. gigantea, 8 upper
(73) and under (74) surfaces; MANITOBA: Churchill. 75-76, C. gigantea
harroweri, topotypic 8 upper (75) and under (76) surfaces; WYOMING:
Sublette Co. 77-78, Same topotypic 9 upper (77) and under (78) surfaces
WYOMING: Sublette Co. 79-80, C. m. meadii x C. pelidne skinneri hybrid (?),
8 upper (79) and under (80) surfaces; WYOMING: Sublette Co.


1>


67 f'


r


r7

1


















N -

S7 82 83
... : ,,,,, -.. "- : .'.',.,. ,**
^ -**iLJ- _- _'ae f 41 -,- t_ ,- -f "" *.....*"
^.^tl~~~~~~~~~~~~~ IM't~KA- ~-~_ -- __._ C.L ff-l |


- b -'.
r fl


86 ;
r, *.%< .


y,; 1
aFt.ih


Figs. 81-88: Colias. 81-82, C. alexandra astraea, the Holland "type" S
upper (81) and under (82) surfaces; ALBERTA. 83-84, same, the Ed-
wards type & upper (83) and under (84) surfaces; WYOMING: Yellowstone
Lake. 85-86, C. alexandra astraea, the Holland "type" 2 upper (85) and
under (86) surfaces; MONTANA: Judith Mtns. 87-88, C. g. gigantea, a
upper (87) and under (88) surfaces; ALASKA. Photographs by F. M.
Brown.


*T^'"




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