Volume 8, Number 6
ISSN 2643-4776 (print) / ISSN 2643-4806 (online)
27 April 2020
The Taxonomic Report
OF THE INTERNATIONAL LEPIDOPTERA SURVEY
Re-evaluation of Euchloe olympia (W. H. Edwards, 1871)
(Pieridae: Pierinae) from a geographic and taxonomic perspective.
Harry Pavulaan 1
606 Hunton Place NE
Leesburg, VA 20176
ABSTRACT. The taxonomic status of regional populations of Euchloe olympia (W. H. Edwards, 1871) is re¬
evaluated here. A review of pertinent literature and a critique of Clench & Opler (1983) are provided. The taxon rosa and two
distinct populations of the Great Lakes “dune form” are each proposed for recognition at subspecific rank. The status of
synonym anniha Ebner (1970) is addressed.
Key words: Apical infuscation, dune form, inland form, isolates.
ZooBank registration: urn:lsid:zoobank.org:pub:B 12650ED-EC10-46D3-BAE2-74CA80ECE605
INTRODUCTION
William Henry Edwards first described Anthocaris olympia (W. H. Edwards, 1871) as follows:
AWTOOCAillfi OLYMMAj n,
J'riimiries long and narrow.
JMe.—Expands LG Jbclt. Upper mlo pure white, grey at ba»< of
wing-; ji large grey patcili td npex of primaries, partially replied bj
white; cwtal margin slightly specked with btiek; on arc a straight
black tar. not prolonged uhovo gub-caatril uemiro. Secondaries him*
a few black mx\& at outer angle and n small wed go-shaped black spot
nearer bitse nn costa.
ruder side white; primaries. luiw a sntnll grey eubnpicnl patch on
cdJstQ r nearly covered with gneco *c;d>w r and a klnt greenish puteh on
htiul margin in tire iliKouhd iutcrsfiacos; dt*eal spot narrow, inmile
enclosing ■ while streak, Secondaries crucoed by bands of yellow
grtin cu ti grey ground, the fir*t, near bnx forming a segment of a
circle; the second, just anterior to the middle of the wing, of irregular
width, projecting n short, truncated branch on sub-median oertule
that connects with the third band, and ending in a slurep spur a lit-
tie inside abdominal margin ; lire tliird kind, commencing cm abdo-
iniuLi] margin, is ul firet lii oud, uml after comp eting with sueumll band,
narrow* to luilf lLa previous width, then enlarges greatly and curio*
Inward* hind margin, which it strikes in three branches, borides acini.
Inga fourth to outer angle; remetjnjc&lhi» 1n*t branch Lit d]«tonm<ctud
ami forms a marginal l spot; tire bunds nre mnre or lest marked by
Kuuill white and |K>infs.
Body black covered with grey hairs; beneath greeoialMrbile; palpi
white, with grey intei>pL*n-«d; ankuiue grey above, yellowish below ;
tdtib blown above, bright yellow below mid at tip,
>Vm<i fc> —Expand* 1.7 inch.
Similar touuilej the grey apical [Hatch otic taring » dirtinci white
1 Staff Member, The International Lepidoptera Survey, Leesburg, VA.
ppm; iltol bhtiidcr, simmit?, scwVi iijt u slinn pjmr ulong $ub~
'iKMsrnl; ciwtjkl ihljaiS jiignj irwitotetl; mulcr $fle a«f in mtilu,
Fiw*l 1 & . 1 9 , lalcun ;ti tMbiindi. \\ r , in April, 1£7I, md
1 S unken by Jlr- Hull, ;it IhklluL, Tcx»b t auJ in uoHucliuu of the
Ahl?GURli ol’CVnqk AuiiL rojubridyo,
1 w\i± wry mill li aurpi'isesil nt finding this jiuir at Canlbuigh among
« l<4 of Cut at hi > tuJu h in April* 1*71. by n jroimg frk-uil whom I bad
onj*n»u?d to inko everythingUoit waK ivbiiu, I hm.l never before met with
it. mul cli niuInL ill fit li utility hail beau pretty wrcli jrfDaueri. It is idso
reinurkblac I lint the nest gpeciifcisii .should come from f-ij remote a dts-
eon nt of tbu three ha ink crushing iecoufoiries.
Interestingly, the species was described from a pair taken at Coalburgh, West Virginia and a male
taken at Dallas, Texas. While not illustrated with the original description, olympia was later illustrated in
W. H. Edwards (1884) (Fig. 1) and a male [top left image of the TTR banner for this paper] was later
illustrated in W.H. Edwards (1897). F. M. Brown (1973) designated a male lectotype for olympia from
the W. H. Edwards collection in the Carnegie Museum of Natural History (fig. 3), which is illustrated in
Fig. 1 of F. M. Brown (1973). The type locality was noted from Edwards’ original male lectotype label as
“Kan”. Opler (1966) recommended that the type locality be more precisely stated as: “banks of the
Kanawha River, near Coalburgh, West Virginia.”
Fig. 1 . Original figures of Anthocaris olympia (W. H. Edwards, 1884). Male dorsal (left), male
ventral (center), female dorsal (right), [photo credit courtesy John Calhoun]
V 2
Fig. 2. Original figures of Anthocharis rosa (W. H. Edwards, 1897). Male dorsal (left), male
ventral (center), female dorsal (right), [photo credit courtesy John Calhoun]
2
William Henry Edwards later described Anthocharis
(W. H. Edwards, 1882) as follows:
Anthocharis Rosa.
Male.— Expands 1.2 inch.
Upper side pure white, gray at bases of wings; primaries
have a snort, straight, black bar on costal margin near apex,
turned back obliquely, which, extended, would strike middle of
inner margin; another bar on hind margin on upper branch of
median ; the apical area between these almost immaculate, a few
scattered black scales only lying near costa; on arc of cell, a nar-
row black bar, bent ora little sinuous; costal edge sometimes
pure white, immaculate; sometimes with a few black scales and
points. Secondaries are of thinner texture, discovering the mark¬
ings of under side; at outer angle a slight edging of black at the
upper branch of sub-costal, and the fringe against this is black:
also, there is black fringe at end of the lower branch of same
nervurc; otherwise all hinges arc white.
Under side white, with a pink tint over costa of primaries
and all of secondaries above subcostal nervurc, deepest next
base ; costal edge either immaculate, or with five or six separate
black points; the discal and costal bars repeated, reduced, the
latter densely covered with yellow scales; the bar on hind mar¬
gin suppressed, but represented by yellow : the ncrvulcs about
a; Jx yellow, and on costal edge a black streak at end of each
nervulc.
Secondaries have three cross bands of black, sprinkled thickly
with yellow scales, with some open spaces or spots, discovering a
yellow ground; the anterior band narrow, making a circle about
base ; the second, discal, sends a short, stout branch along median
nervurc to third, then narrows and is attenuated to a point at
inner margin ; the third, starting on inner margin, is bread, and
after the junction with second, narrows greatly, then widens and
strikes hind margin in a triple fork; at outer Jingle a short, nar¬
row bar similar to the bands; the ncrvurcs and branches on mid¬
dle of the wing yellow.
Body black ; the thorax covered with long gray hairs; be¬
neath, thorax yellow, abdomen white along venter; legs pinkish,
the femora covered with long white hairs; palpi white with much
gray; antenna: white above, yellowish below : club white above,
yellow below and at tip.
Female.—Expands 1.4 inch.
Color of male, on both surfaces, similarly marked; a few
black scales are scattered between the ends of the apical bars, sug¬
gesting a cross band.
From six examples, 3 a 3 taken by the late Mr. J, Boil, in
western Texas, March, 1S7S- Mr. Boll informed me, m answer
to my inquiry, that he Look many more, and all were of the type
sent me. The species is nearest O/jmpitu Edvv . 4 and the banding
of underside is of same diameter. It differs From that species
by the immaculate apex, and by the presence of the straight, ob¬
liqu e costal bar, instead of an irregular patch.
The taxon rosa was described as a species from three pairs taken in “western Texas”. The
butterfly [top right image of the TTR banner for this paper] was later illustrated in W. H. Edwards (1897)
(fig. 2). F. M. Brown (1973) designated a male lectotype for rosa (fig. 3) from the W. H. Edwards
collection in the Carnegie Museum of Natural History and restricted the TL to “Wichita County, Texas”.
This specimen is illustrated in Fig. 2 of F. M. Brown (1973).
3
Fig. 3. Anthocaris olympia (W. H. Edwards, 1871) lectotype (left image) as illustrated in F. M.
Brown (1973). Anthocharis rosa (W. H. Edwards, 1882) lectotype (right image) as designated
by F. M. Brown (1973). [photo reproductions by permission of American Entomological
Society]
REVIEW OF PERTINENT LITERATURE, FOLLOWING
ORIGINAL DESCRIPTIONS
(including major synonymic and other works)
H. Strecker (1872): The author followed Edwards’ original treatment of olympia under genus
Anthocharis. [Rosa had not yet been described.]
W. F. Kirby (1877): The author included olympia under genus Zegris in the 1877 Supplement to
his ‘Synonymic Catalogue of Diurnal Lepidoptera’. [Rosa had not yet been described.]
H. Strecker (1878): The author again followed Edwards’ original treatment of olympia under
genus Anthocharis. [Rosa had not yet been described.]
W. H. Edwards (1884): In this edition, the author again listed Anthocaris olympia , without
mention of rosa , but rosa was listed in the Edwards (1897) edition.
G. H. French (1886, 1914): In all editions through 1914, only species Anthocaris olympia was
listed, without mention of rosa.
W. S. Blatchley (1891): The author’s report of Anthocharis olympia is the first published account
of the lower Lake Michigan lakeshore population. Blatchley states: “Occurs in small numbers about
Whitings, Lake County...It frequents cultivated grounds, gardens and meadows”, though this is certainly
not a description of the known dune habitat. There are presently only small fragmented remnants of the
ancient dunes along the lakeshore at Whiting, certainly more expansive in the late 1800’s.
H. Skinner (1896): In his discussion of Anthocharis , the author was first to suggest, contrary to
Edwards (1882): “Rosa seems to be the same, or at best a var. of olympia. I should say it represented the
southern end of the vertical distribution.”
Scudder (1889): The author treats olympia at species rank under genus Synchloe (and considers
Anthocharis and Zegris generic-level synonyms) and without mention of rosa.
W. H. Edwards (1897): In this edition, the author listed Anthocharis rosa as a species-level
taxon. He makes notes similarities and differences from olympia , which he still considered a separate
species but questioned: “It may be that these are properly but forms of one species, Olympia the northern,
Rosa the southern form. As yet not much is known about either...” Interestingly, Edwards seemingly
makes reference to the dune populations of olympia : “The species [ olympia ] seems particularly abundant
at Whiting’s, Lake County, Indiana.. .One of these is represented in Fig. 5...” Edwards’ fig. 5 may be the
first actual depiction of the dune phenotype!
W. Beutenmuller (ed.) (1897): In the Proceedings of the New York Entomological Society,
meeting minutes of October 16, 1897, William Beutenmuller quotes himself: “Rosa is without much
doubt a variety of olympia .” In the meeting minutes, Beutenmuller referred to subspecies as “races”, thus
he certainly considered rosa an infrasubspecific form.
W. Beutenmuller (1898): In the author’s revision of North American Euchloe , he includes
entries for Euchloe olympia and “ Euchloe olympia var. rosa ”, providing a description of both.
Interestingly, he did not include distribution of nominotypical olympia , yet he described the “habitat” of
rosa as “West Virginia to Texas and Nebraska.” In this paper, Beutenmuller did not elaborate on his
application of the term “var.”, thus interpretation of Beutenmuller’s application of the terms “race” and
“variety” is based on his quotes from Beutenmuller (1897). Also, interestingly, the author did not
illustrate nominotypical olympia , yet he illustrated rosa. In a response to Beutenmuller (1897), Butler
(1899) states: “Euchloe olympia is undoubtedly a species of Zegris .”
H. Skinner (1898): The author treats species Anthocharis olympia at species rank with the range
of “W. Va., Ind., Neb., Col., Minn., Summit, N. W. T.” but treats rosa as “var.”, giving the range as
“West Texas”. Skinner similarly listed geographical taxa (subspecies) as “var.” and did not employ the
term “subspecies”. By giving the range of rosa , one can imply that he considered rosa as a subspecific
taxon.
S. F. Denton (1900): Lists Anthocharis olympia but makes no mention of rosa. Interestingly, the
author misidentified two specimens of what appear to be Euchloe hyantis on page 330 as “olympia”.
W. J. Holland (1899, 1900, 1902,1904, 1916a): In ‘The Butterfly Book’, in what seems to be a
case of accidental omission, olympia is merely illustrated on Plate XXXII (fig. 39) as “Euchloe olympia ,
Edwards, var. rosa Edwards”, yet there is no species entry for olympia in the text. However, Euchloe
rosa is given full species treatment (following Edwards).
H. G. Dyar (1902): The author includes olympia within the genus Synchloe and treats rosa at
subspecies rank.
J. H. Comstock & A. B. Comstock (1904): The authors include olympia within the genus
Synchloe but do not include mention of rosa.
W. G. Wright (1906): Euchloe Rosa is treated at species rank but with little natural history
information. Of great interest is the figured specimen (Plate VII, Fig. 56c), which displays a yellow
ventral ground color instead of the usual white. The specimen was reportedly taken at or near Napa
[California]. However, J. W. Tilden (1975) states that the illustrated specimen is “an off-color specimen
of Euchloe ausonides Lucas.”
M. J. Elrod (1906): The author refers only to Synchloe olympia and does not mention rosa.
C. A. Shull (1907): The author, referring only to species-rank Anthocharis (Synchloe) olympia ,
published a detailed description of the immature stages and flight characteristics of the lakeside dune
population in northern Indiana. Shull did not note subspecific differences between populations of
olympia.
W. J. Holland (1916b): In ‘The Butterfly Guide’, in what appears to be a repeat of accidental
omission from earlier works (Holland, 1900, 1902, 1916a) there again is no species entry for olympia in
the text [no doubt the work of the publisher]. However, Euchloe rosa is given full species treatment
(following Edwards) and is illustrated on Plate Cl (fig. 2).
W. Barnes & J. McDunnough (1917): In ‘Check List of the Lepidoptera of Boreal America’,
rosa was listed at subspecific rank under Zegris olympia.
C. M. Weed (1917a, 1917b, 1923): The author refers only to species-rank Synchloe olympia and
makes no mention of rosa.
S. Moore (1922): The author studied butterflies over a five-year period in northern Michigan and
documented two specimens listed as Zegris olympia rosa.
A. Seitz (1924) [text and plates separate]: The author lists Zegris olympia and treats rosa as an
aberrant form. Interestingly, his illustrated specimen of olympia (plate 28b) is a rosa phenotype.
W. J. Holland (1931, 1951): In ‘The Butterfly Book (New and Thoroughly Revised Edition)’,
olympia and rosa are each given full species treatment. Holland gives the range of olympia as
“southwestern Pennsylvania and West Virginia westward to Minnesota and Colorado.” He gives the
range of rosa as “Texas”. Euchloe olympia is illustrated on Plate LXVII (fig. 28) and Euchloe rosa is
illustrated on Plate XXXII (fig. 39).
F. C. Cross (1937): The author lists Euchloe olympia at species rank, occurring in the “eastern
part of the state and along the foothills”, but interestingly states: “Typical specimens exhibit a rosy flush
on the wings which has prompted many collectors to confuse our race with olympia rosa. It is unclear if
he considered Colorado olympia a different “race” from either rosa or nominotypical olympia.
W. D. Field (1938): The author discusses differences between the eastern subspecies olympia and
the western subspecies rosa at length.
R. A. Leussler (1938): This is the last known published treatment of olympia under genus Zegris
and treats rosa at subspecific rank.
J. McDunnough (1938): In ‘Check List of the Lepidoptera of Canada and the United States of
America’, the author maintained rosa at subspecific rank under Euchloe olympia.
E. P. Meiners (1939): The author discussed life history stages of olympia in Misouri, but did not
address taxonomic issues, thus did not treat rosa at any rank.
A. H. Clark & L. F. Clark (1939): The authors treated rosa as a form of Euchloe olympia in
Virginia. They first stated: “On May 7 and 8 of this year, although we took a few battered individuals of
more or less typical olympia , nearly all that we captured were almost, or quite, typical rosa.” They go on
6
to elaborate: “[T]he natural conclusion is that in the cool weather of early spring this species appears in
the form olympia , but later in the season after warm weather has become established it changes over to
form rosa. These two forms, therefore, are primarily cool and warm weather forms of the species, and not
geographical races.”
Examination of Appalachian specimens in the American Museum of Natural History, Field
Museum, National Museum of Natural History, and the Peabody Museum of Natural History, as well as
my own specimens from neighboring Maryland, reveals few individual variants qualifying even as
intermediates. Most individuals are typical Appalachian olympia , and have considerable apical
infuscation, though a very small percentage of individuals (<10%) do have reduced apical infuscation.
What may factor into the authors’ assertions, is the fact that olympia’ s wings tend to be very frail and tend
to show age wear very quickly within a few days of flight, thus scale loss and fading out of the dark apical
area of the fore wing. Thus, some later individuals might appear more like rosa than the earliest, freshest
olympia to emerge weeks earlier. However, the authors likely did not notice an important difference
between nominotypical olympia and Great Plains rosa : the different shape of the forewing [discussion to
follow].
A. H. Clark (1940): The author treated rosa as a form of Euchloe olympia in Virginia. The
author repeated his earlier description of finding nominotypical olympia “earlier in the spring in the same
region” and rosa later in the spring. He again went on to conclude: “On examining the matter we found
that early in spring when it is still cold the typical form, olympia , is on the wing. After it becomes hot the
butterfly changes over to the western type of coloration.” What is interesting is that Clark illustrated two
specimens: a nominotypical olympia in very fresh condition collected on April 24, 1938; and a very worn
specimen collected in nearby West Virginia on May 8, 1939, which he referred to as E. o. rosa. That
latter specimen displayed considerable scale wear from age, making it somewhat difficult to conclude that
it represented the “form characteristic of the region from Nebraska southward to Texas and New Mexico.”
Interestingly, that specimen displayed a wedge-shaped black mark midway along the outer margin of the
hindwing which is commonly seen in eastern nominotypical olympia and not in Great Plains rosa.
R. W. Macy & H. H. Shepard (1941): The authors provided considerable discussion of “race
rosa ”, which implies subspecific treatment.
A. H. Clark & L. F. Clark (1951): The authors again treated rosa as a form of Euchloe olympia
in Virginia. They stated: “[Individuals of this species are always typical olympia ”, but went on to
elaborate with the following: “If during the period of emergence the weather becomes hot and dry, this
tendency is carried to an extreme and the form rosa appears, quite like specimens from Texas...The form
rosa flies with worn individuals of typical olympia and a more or less complete series of intergrades. In
Virginia and West Virginia it is possible to capture on the same day typical olympia and typical rosa ,
though most individuals will be intermediate between the two forms.”
A. B. Klots (1951): The author treats Euchloe olympia rosa at subspecies rank, but states: “not a
very valid subspecies.” Interestingly, Klots provides illustration of a specimen of the distinctive dune
population from “N.E. Lake Co., Ill.” (Page 208, illustration 6).
F. M. Brown, D. Eff & B. Rotger (1957): The authors treat Euchloe olympia rosa at subspecific
rank.
W. T. M. Forbes (1960): The author simply states, under Euchloe olympia: “[W]estem
specimens from Minnesota to Kansas and Texas considered a race rosa Edwards, but not really distinct.”
C. F. dos Passos (1964): In ‘A Synonymic List of the Nearctic Rhopalocera’, rosa was listed at
subspecific rank under Euchloe olympia.
7
R. R. Hooper (1969): Euchloe olympia rosa is listed for Saskatchewan.
J. A. Ebner (1970): The author inadvertently published a new name, Euchloe olympia anniha
which was attributed to dos Passos & Klots in error. Ebner (personal correspondence) informed me that
when the manuscript was submitted to the publisher, the trinomial name was not included, and only
appeared when ‘Butterflies of Wisconsin’ went to print: “I have no recollection as to how or why this
name was applied”. According to Ebner, the inclusion of the trinomial anniha was certainly added by the
editorial staff of the publisher. The name is apparently a misspelling and misapplication of annickae dos
Passos & Klots, 1969, a subspecies of Anthocharis midea. Masters (1972) states: “Dos Passos and Klots
have never described any subspecies of Euchloe olympia nor have described anything with the name of
anniha. The name anniha must be credited to Ebner.” Julia Colby, Vertebrate & Invertebrate Collections
Manager, Milwaukee Public Museum (personal correspondence) indicated that there are several
specimens identified as “ anniha ” in the museum’s collection, but all were collected “at least a decade
after Mr. Ebner’s book was published.”
Reference here is made to phenotype and range: “It is essentially white, although a few black
markings accent the fore wings. The subspecies anniha seems to be prevalent throughout most breeding
colonies in the state [Wisconsin]. Here and there an individual with more extensive black markings also
turns up, placing it more closely to typical olympia ”. Reference to the butterfly being “essentially white,
although a few black markings accent the forewings” draws up comparison closer to subspecies rosa ,
though some individuals are described as being closer to nominotypical olympia. I will add here that rosa
specimens from central Wisconsin populations compare well with Great Plains specimens as far south as
Texas. I consider “ anniha ” as a junior subjective synonym of rosa , based on the description above, and
on the specimen image in Ebner (1970).
Though a holotype was not designated, nor intended, a male specimen was illustrated (Fig. 4),
placing it very closely to the rosa phenotype. Ebner (personal correspondence) indicated that the
disposition of that specimen is unknown, nor was the location of that specimen documented. The only
reference to a Wisconsin location for olympia in Ebner (1970) is “Chippewa County”. I have selected an
image of a very similar male specimen (Fig. 5) from the National Museum of Natural History to serve as a
representative for the illustrated phenotype, from Nekoosa, Wood County, the closest location to
Chippewa County (southeast corner) by approximately 62 miles.
The hostplant of anniha is described as “Rockcress”, which can apply to several species of Arabis ,
thus it is not known if the illustrated specimen was associated with either Arabis lyrata , Arabis
drummondii , or another species.
Fig. 4. Euchloe olympia u anniha ” Ebner (1970). Male upper (left), male under (right). [Original images
reproduced by permission of Milwaukee Public Museum]
WI., May 5, 1918, leg. E. Beer. Dorsum (left), venter (right). Photograph by permission of the
Smithsonian National Museum of Natural History.
C. D. Ferris (1971): The author treats rosa at subspecific rank but also suggests that
“...subspecies olympia (Edwards) also may occur in our area [Wyoming].”
K. Johnson & J. M. Malick (1972): The authors listed subspecies ‘ Euchloe olympia anniha dos
Passos & Klots’ for central Wisconsin, apparently following the treatment of Ebner (1970). The only two
records listed are for Portage County, and the only image, one of the rosa phenotype, is presumably from
the same county.
K. Johnson (1972): The author treats rosa at subspecific rank.
J. H. Masters (1973): Interestingly, the author, despite assertion over the inadvertent naming of
anniha (Masters, 1972), included this subspecific name and wrongly attributed it as “ Euchloe olympia
anniha dP. & Kts.” [This paper was issued in the J. Res. Lepid. 1972(1973), thus was likely submitted for
review prior to Masters (1972).] This is the only other literature reference to anniha that could be found.
R. R. Hooper (1973): The author treats rosa at subspecific rank.
W. H. Howe (1975): In ‘The Butterflies of North America”, the author treats rosa at subspecific
rank and illustrates rosa specimens on plate 71 (no’s 24 & 25).
W. H. Wagner (1977): The author published a detailed description of what he referred to as the
distinctive “dune form” of Euchloe olympia. Wagner’s first encounter with this phenotype was in the
dunes area at the south end of Lake Huron in the Ipperwash Area of Lambton County, west of Grand
Bend. His preliminary observations in 1975 suggested that this distinct population “might even represent
a distinct subspecies, so numerous were the differences”. Later observations of this diminutive dune
phenotype were made in similar habitat at the south end of Lake Michigan in Berrien County, Michigan.
Wagner went on to describe differences between the Michigan “Inland Form” and “Dune Form”, noting
that: “The Dune Form is readily distinguished from the Inland Form by its smaller size, whiter ground
color, and more complex pattern of markings on the secondaries below”, though he did acknowledge that
there is some overlap in characters. Wagner also stated: “The Lake Michigan Dune populations are not
quite so extremely differentiated as the Lake Huron”.
Of interest is Wagner’s observations regarding differing hostplant preferences of the “Dune” and
“Inland” Forms in Michigan. He noted that “Inland Form” populations of olympia in Allegan, Montcalm
9
and Roscommon Counties utilize Arabis drummondii while the “Dune Form” populations of Berrien
County, ML and Lambton County, ON. utilize Arabis lyrata. Wagner indicated an initial impression that
Arabis lyrata was “primarily a plant of dunes and shoreline sandy strips” (though it is also found
throughout much of Michigan), thus being a valid differentiating biological character (in Michigan).
Later plotting revealed that A. lyrata also occurred commonly throughout much of Wisconsin, especially
in the “Driftless Area” in the southwestern part of the state. He concluded: “On the basis of the
phytogeographical data, therefore, it seems unlikely that it is foodplant preference alone that governs the
differentiation of the Dune populations.”
S. Kohler (1980): The author listed county distribution of “olympia f. “rosa”” in Montana but
commented: ““rosa” is best considered a form rather than a subspecies.”
C. D. Ferris & F. M. Brown (1981): The authors consider rosa a form: “Two sspp. Have been
described, but one appears invalid as discussed below. The nominate ssp...generally occurs in our
region...Throughout this butterfly’s range, and even in a given locality, both phenotypes can be collected.
For this reason, “rosa” must be considered as a form, not a ssp.” I would like to address these statements
here. One specimen illustrated (“fm. “rosa” m, D, V.”) is indeed rosa. The first specimen (“ olympia m,
D, V”) does not appear to be of the nominotypical phenotype as suggested in the authors’ statements. It is
a rosa primarily by wing shape, but the gray subapical bar is characteristic of intermediate forms found in
Wisconsin. Not knowing where the illustrated specimens originated from, what series the authors
examined, or what their experience was, in regard to regional variation in olympia , I feel compelled to
regard their conclusion as insufficient, given the lack of Rocky Mountain specimens available for
examination in collections.
L. D. Miller & F. M. Brown (1981): In ‘A Catalogue/Checklist of the Butterflies of America
North of Mexico’, rosa is now listed as a junior subjective synonym of Euchloe olympia.
R. A. Layberry, J. D. Lafontaine & P. W. Hall (1982): In ‘Butterflies of the Ottawa District’,
the authors list subspecies rosa as occurring in the Ottawa area (including adjacent Quebec).
R. W. Hodges (1983): In ‘Check List of the Lepidoptera of America North of Mexico’, rosa is
again listed as a subspecies of Euchloe olympia.
H. K. Clench & P. A. Opler (1983): The authors published a study describing geographic
variation in Euchloe olympia , in which several aggregate population groupings, or biogeographic
“isolates”, were identified and statistically analyzed for several adult wing characters, with the result as
possible alternative taxonomic treatments. These isolates were also grouped based on distinctive traits
relating to hostplants and habitat types. The authors concluded that rosa best be treated as a synonym of
nominotypical olympia , though offering: “[Ojne might restrict nomenotypic olympia to isolates with dark
apical infuscation.. .while applying the subspecific name rosa to all lightly marked isolates”.
Y. Sedman & D. Hess (1985): The authors simply state: “Specimens from Mason County...are
recognized as the form “rosa”.
J. W. Tilden & A. C. Smith (1986): In ‘A Field Guide to Western Butterflies”, the authors treat
“rosa” at subspecific rank and illustrate a specimen designated at subspecific rank (plate 25, illustration
4),
J. A. Scott (1986b): In ‘The Butterflies of North America’, the author does not recognize any
subspecies under Euchloe olympia. Interestingly, he gives the common name as “Rosy Marble”.
R. A. Royer (1988): Describes the ventral “rosy flush” and “much reduced DFW apical dark
markings” as a variable character and states: “this character in the extreme defines form “rosa”.
10
C. A. Bridges (1988). The author treats rosa as a junior synonym of olympia , citing Ferris &
Brown (1981): “Note - Ferris does not consider this a valid subspecies.” [Bridges bibliographic
reference: bl386, bl396]
P. Klassen, A. R. Westwood, W. B. Preson & W. B. McKillop (1989): The authors state that
“no subspecies are recognized”.
B. D’Abrera (1990): Placed olympia back into genus Zegris. The author stated: “This species has
been shifted around from Zegris to Euchloe , and back again. The only features it does not share with
Palaearctic Zegris are the hirsute larvae and pupa within a girdled cocoon. However, the shape of the
costal margins of both wings...together with a typical (for Euchloe) h.w.v. pattern...all make its
placement here more logical. The pinkish ground colour of olympia (especially in fresh specimens) also
make it a ‘bad’ fit with Euchloe , but a ‘good’ one with Zegris .” The author illustrates specimens taken in
Kansas. This view of including olympia in genus Zegris did not gain popular acceptance post-1990. The
name ‘rosa’ was not mentioned.
D. C. Iftner, J. A. Shuey & J. V. Calhoun (1992): The authors treat rosa as a form:
“Individuals with rosy markings on the ventral hindwing are known as form ‘rosa’ (W. H. Edwards,
1882). These rosy markings fade and disappear on pinned specimens.”
R. A. Layberry, P. W. Hall & J. D. Lafontaine (1998): The authors treat olympia only at
species rank with no mention of rosa.
M. C. Nielsen (1999): The author does not treat rosa , but in two plates (page 49), he illustrates
dorsal differences between a male of “inland” olympia and what he refers to as a “Coastal male”.
D. K. Parshall (2002): The author produced a conservation assessment of Euchloe olympia for
the USDA Forest Service, Eastern Region. A considerable amount of published and new (previously
unpublished) information was integrated into this report. Parshall erroneously stated: “No subspecies has
ever been described” and cited Edwards (1882) for describing “form” rosa. To correct the record here,
Anthocharis rosa (W. H. Edwards, 1882) was described as a full species, as indicated by the original
binomial treatment, making the name available for the Great Plains subspecies. Parshall also summarized
variation in olympia as: “remarkably uniform in appearance throughout its range in the western and
eastern U.S.”, which I find to be inaccurate, and also contradicts the findings of Clench & Opler (1983).
J. P. Pelham (2008): In ‘A Catalogue of the Butterflies of the United States and Canada’, rosa is
listed as a junior subjective synonym of Euchloe olympia.
J. A. Scott (2008): In the ‘Biological Catalogue of North American Butterflies’, the author lists
rosa as a synonym of E. olympia.
M. S. Fisher (2012): The author stated: “rosa was used as a form name but is presently
considered a synonym.”
G. R. Pohl, G. G. Anweiler, B. C. Schmidt & N. G. Kondla (2010) treat rosa at subspecific
rank, stating: “...AB [Alberta] populations appear to be distinct from nominate olympia and are more
properly aligned with the subspecies rosa (Edwards).”
G. R. Pohl, J. Landry, C. Schmidt, J. D. Lafontaine, J. T. Troubridge, A. D. Macaulay, E. J.
Van Nieukerken, J. R. Dewaard, J. J. Dombroskie, J. Klymko, V. Nazari & K. Stead (2018): The
authors treat rosa as a junior synonym of olympia.
J. A. Scott (2020): The author discusses life history aspects of E. olympia in Colorado but does
not mention rosa in the discussion. However, he refers to the common name as “Rosy Marble (Olympia
Marble)”.
Some of the more recent works, mainly state or provincial treatments and regional field guides
treat Euchloe olympia at species rank only, do not mention rosa , nor do they provide assessment of
regional variation: Ehrlich & Ehrlich (1961); Clarke (1962); Irwin & Downey (1973); Pyle (1981); Opler
& Krizek (1984); Ely, Schwilling & Rolfs (1986); E. M. Shull (1987); Heitzman & Heitzman (1987);
Holmes, Hess, Tasker & Hanks (1991); Opler & Malikul (1992); Acorn (1993); Bird, Hilchie, Kondla,
Pike & Sperling (1995); Neck (1996); Glassberg (1999); Coveil (1999); Opler & Wright (1999);
Bouseman & Sternburg (2001); Glassberg (2001); Marrone (2002); Opler & Warren (2003); Brock &
Kaufman (2003); Royer (2003); Dole, Gerard & Nelson (2004); Daniels (2004, 2005); Douglas &
Douglas (2005); Schlicht, Downey & Nekola (2007); Betros (2008); Chu & Jones (2011); Patterson
(2011); Glassberg (2012); Belth (2013); Spencer (2014); Venable (2014); Jeffords, Hall, Jones, Guidotti
& Hubley (2014); Post & Wiker (2014); Hardesty (2015); Acorn & Sheldon (2016).
DISCUSSION
I initially examined the specimen series at the Smithsonian National Museum of Natural History,
with the intent of better understanding the regional variation as described in Clench & Opler (1983).
Utilizing my own reference series, those of the American Museum of Natural History (New York, N.Y.),
Field Museum (Chicago, IL.), the National Museum of Natural History (Washington D.C.), Peabody
Museum of Natural History (New Haven, CT.) and also a broad range of published and internet sources, it
was possible to update and supplement what is now known of the distribution of olympia and its regional
variation. Despite its broad range and varied habitat choices, the butterfly appears to be highly localized
in occurrence (though somewhat more widespread in the Great Plains), with widely scattered colonies
consisting of relatively small numbers of observed adults.
Clench & Opler (1983) recognized several regionalized population groupings (isolates) as follows.
These were based on the following characters: (1) forewing length (base to apex); (2) width of forewing
discal black bar; and (3) degree of black infuscation on forewing apex above.
•Appalachian isolate : Ranging from Pennsylvania south through West Virginia. This is among the other
isolates with smaller individuals, but with the heaviest apical infuscation (Fig. 6). This “isolate” refers to
nominotypical olympia.
• Great Lakes isolates : Series of populations surrounding the Great Lakes including the diminutive dune
populations. The authors divided their ecology discussion into two [sub]isolates:
• Great Lakes inland isolates : The authors include here “a mixed lot of populations with quite
differing ecological features”. These group with isolates with the smaller individuals; however,
Michigan specimens averaged with heavier apical infuscation, thus placing specimens closer to the
Appalachian isolate while Wisconsin specimens averaged with lighter apical infuscation (thus
being more similar to Plains isolates).
• Great Lakes dune isolate : This isolate is found in two disjunctive areas, restricted to lakeside
dunes along the south end of Lake Michigan in Wisconsin, Illinois, Indiana and Michigan; also at
the “upper edge of Lake Huron” [corrected here to the SOUTH END of Lake Huron]. This isolate
has the smallest-sized adults (Wagner, 1977; Clench & Opler, 1983). Apical infuscation is heavy,
more like Appalachian isolate.
12
• St. Louis isolate : A grouping of river bluff colonies in the St. Louis area. This isolate has large
individuals, but with heavy apical infuscation, thus identifying the phenotype with the Appalachian
isolate. A portion of individuals are intermediate to rosa in nature (Fig. 7), thus I place this as a grouping
of intermediate populations. Two Missouri colonies that I have studied, one in St. Francois county, the
other in Wayne County, similarly have individuals with very dark apical infuscation, placing those closer
to the Appalachian populations.
• Texas isolate : A grouping of several central and eastern Texas colonies near several major rivers (Fig.
8). This isolate groups with the other Plains isolates by having large individuals, and with the lightest
apical infuscation. This grouping also encompasses the rosa TL.
• Arkansas River isolate : A series of populations extending from Arkansas through Oklahoma and
Kansas; also along the Canadian River in north Texas. This isolate groups with the other Plains isolates
by having large individuals, and with the lightest apical infuscation.
• Missouri River isolate : Essentially applicable to all northern Great Plains populations in the Missouri
River watershed from Missouri to North Dakota and then north into Manitoba. This isolate groups with
the other Plains isolates by having large individuals, and with the lightest apical infuscation.
• Front Range isolate : Populations residing in the Colorado Front Range and along the western edge of
the Great Plains into Wyoming. Specimens of this isolate are small, like the Appalachian and Great
Lakes isolates, and in general having light apical infuscation as in the Plains isolates. Occasional
individuals with heavy apical infuscation, giving the appearance of nominotypical olympia , may be due to
elevational influence.
Based on the above, Clench & Opler (1983) left us with three taxonomic choices: (a) apply
separate subspecies status to each isolate; (b) consider isolates with dark infuscation (Appalachian, Great
Lakes dunes and St. Louis) as nominotypical olympia ; and all isolates with light infuscation as subspecies
rosa; (c) not apply subspecific status to any regional isolate. The authors chose option (c). In the present
paper, I choose a modified option (b) as discussed below.
HABITATS AND HOSTS
Since publication of the Clench & Opler (1983) paper, new distributional data has filled in many
of the regional gaps between these described isolates, thus reason to question their viability as identifiable
phenotypes in some regions, from a morphological perspective. I suggest here that, while the 1983 study
maintains valid points, the biological aspects of olympiad distribution is apparently more fragmented and
complex than that study suggests, yet regional variation based simply upon the degree of gray apical
clouding, shape of the forewing, and wing size point to a somewhat more simplified solution. Instead of 8
isolates as proposed by Clench & Opler (1983) the current mapped distribution (Map 1) shows three main
population clusters: Appalachian Mountains; Great Lakes; and Great Plains.
First and foremost, it is apparent that olympia flies in a very broad range of habitats that varies
from region to region and even within regions. For example, Howe (1975) describes the butterflies as
favoring “open woodland or nearby meadows” in the east; “river woodlands and on river bank bluffs” in
the Great Plains. Pyle (1981) describes the habitat as: “Open woods and meadows in eastern portion of
range; watercourses and nearby fields and bluffs, foothill ridges and open grasslands farther west, sandy
flats and dunes in Great Lakes area.” Clench & Opler (1983) state that olympia occurs in “relatively open
native situations with well-drained rocky or sandy substrates”. Scott (1986b) describes the habitat as
“usually in fairly dry clearings with few plants in E U.S., also prairie...” and notes that the range is
expanding in Ontario. NatureServe (2015) summarizes the habitat as “open woods, barrens, very dry
meadows in eastern part of range and open grasslands to the west”. Following are published habitat types
13
and hosts by state or province. Hostplant synonymy and common names are adapted from the USDA
Plants Database (2017).
United States :
• Arkansas: Rouse (1969) reports olympia to be found in open woodland and nearby meadows. Masters
reported the habitat in northwestern Arkansas to be bottomland deciduous forest (Clench & Opler, 1983).
This was identified as part of the “Arkansas River isolate” (Clench & Opler, 1983). Opler & Krizek
(1984) describe the habitat as “open river forests”. Spencer (2014) describes the habitat as “open areas
such as glades and forest edges”.
• Colorado: Brown, Eff & Rotger (1957) state that olympia “seems to prefer open, dry, grassy areas a the
tops of the foothill ridges and on the open rolling plains”. Howe (1975) gives the habitat as “low, open
foothills”. Ferris & Brown (1980) describe the habitat as “open prairies and the slopes of the low
foothills”, being found in localized pockets. Brown also reported olympia as “found on open ridges or
low hills in short grass prairie habitat, often not too distant from cottonwood-lined streams (Clench &
Opler, 1983). Opler & Krizek (1984) describe the habitat as “rocky foothills”. Scott (1986b) lists
“foothill chaparral”. Chu (2009) reported olympia from short grass prairie. Chu & Jones (2011) include
meadows, rocky ridges and ponderosa pine woodlands, and males are described as patrolling hilltops.
Fisher (2012) describes olympia as a “true prairie species”, occurring “on the eastern plains, especially the
higher prairie and into the foothills and lower mountains along the Front Range where prairie-like sites
often persist.” Scott (2020) gives the habitat as “foothills chaparral/open woods, and Great Plains prairie
especially with some trees.” Scott (1986a, 1992, 2006, 2020) provides a considerable number of
documented host records for the state: Arabis glabra (Tower Rockcress or Tower Mustard), Descurainia
pinnata (Western Tansy Mustard), Descurainia richardsonii (Mountain Tansy Mustard) [synonym of
Descurainia incana \, Descurainia sophia (Flixweed or Herb Sophia), Lepidium campestre (Field
Pepperweed), Lepidium virginicum (Virginia Pepperweed) and Sisymbrium altissimum (Tall
Tumblemustard). Wolfe, Harry & Stout (2010) recorded Arabis fendleri (Fendler’s Rockcress).
• Illinois: Irwin & Downey (1973) stated that the Waukegan Dunes population is “well established,
common to abundant. Sedman (1983) described the habitat as “sand and loess hill prairies”. Sedman &
Hess (1985) state that olympia is “virtually limited...to sand dune areas”, but does not occur in all such
areas, even when the hostplants are present. The authors further state that the butterfly occurs “along
trails or roads, and also in open woodlands and clearings”. One of the listed records was taken in “Sand
Dune-Scrub Oak Prairie.” Bouseman & Stemburg (2001) describe the habitat as “open woods and nearby
fields with low vegetation, in arid or semi-arid locales, often sandy”, adding: “Within its general range,
scattered populations occur. The species is usually scarce, but in some years may be locally common
within a restricted area.” Parshall (2002) describes olympia “in small numbers in widely separated
populations”. Jeffords, Post & Wiker (2014) state that olympia occurs in “dry, sandy, prairie remnants;
oak savannas; and sand dunes” but under ‘Status’, describe the species as “locally encountered in sandy
areas and dry hill prairies along the Mississippi and Illinois rivers.”
• Indiana: W. S. Blatchley (1891) described the habitat at Whiting, in Lake County, as “cultivated
grounds, gardens and meadows”. Whiting is at the lower end of Lake Michigan, thus possibly the first
published record of the dune population. C. A. Shull (1907) and Wagner (1977) both reported the dune
population from lakeside dunes, utilizing Arabis lyrata (Lyre-leaved Rockcress or Lyrate Rockcress).
Belth (2013) adds sand savannas as habitat in Benton and Newton counties. E. M. Shull (1987) describes
olympia as local and rare, utilizing both Arabis lyrata and Sisymbrium officinale, though it is questionable
if reference to the latter was specific to Indiana.
• Iowa: Schlicht, Downey & Nekola (2007) describe the habitat in western Iowa as “dry gravel prairies”
and “sandy bedrock glades” in northeastern Iowa. Orwig (1990) describes olympia as “a woodland edge-
14
savannah species...common on a few of the steepest loess bluffs” along the course of the Missouri River,
but generally rare elsewhere in the state.
• Kansas: Field (1938) describes the habitat as “meadows and woods”, particularly in moist woods along
waterways. Sisymbrium officinale (Hedge Mustard) is given as the hostplant. Ely, Schwilling & Rolfs
(1986) describe the habitat as: “grasslands, woodland edge, roadsides, fields, riparian border.” They note
the butterfly as being uncommon statewide.
• Kentucky: Co veil (1999) describes olympia as “very local and uncommon”. Parshall (2002) describes
populations as very small and widely separated.
• Maryland: Relatively open or thinly wooded deciduous shale barren habitats; overgrown utility line
right-of-ways through mixed deciduous forest (personal observations). I have observed them in very
small numbers within a very open patch of deciduous forest with a thin understory of scrub oak on a
southeast-facing shale slope and on a shrubby east-facing hillside along a utility right of way, where they
utilize both Arabidopsis thaliana (Mouse-ear Cress) and to a lesser extent Cardamine pensylvanica
(Pennsylvania Bittercress). Freshly-laid eggs are reddish orange. Older larvae eat only seedpods of A.
thaliana , as the flowers and leaves are generally withered at that stage. Opler & Krizek (1984) list shale
barrens as the habitat. Parshall (2002) also reports olympia as being found in low numbers and comments
that Gypsy Moth spraying has severely impacted the species. Maryland populations are nominotypical
olympia.
• Michigan: Nielsen (1966, 1967) first reported utilization of Arabis drummondii (Drummond’s
Rockcress) as host in Montcalm County, later in Cheboygan County (Nielsen, 1979). Wagner (1977)
reported the “Inland Form” from prairies in Allegan County, Michigan, while Bill Bouton (personal
communication) describes habitat in Allegan County as advanced second-growth (Oak and Pine)
abandoned fields with openings. Opler (1974), Wagner (1977), Bess (1983), and Ronda Spink (personal
communication) all reported use of the host Arabis drummondii in that area. Bess (1983) also reported
Arabis lyrata from Allegan County but it is not clear whether this represents a lakeside dune population.
Nielsen reported the inland isolate from “openings within scrub oak-jack pine habitat” and an “old pine
burn”, both in the northern part of the Lower Peninsula (Clench & Opler, 1983). The “inland isolate” was
reported as “restricted to prairies” in several central Michigan counties (Clench & Opler, 1983). Opler &
Krizek (1984) list “sand plains” as the habitat. Nielsen (1999) reports the habitat as “Great Lakes dunes
and open oak-pine barrens on sandy soils”. Parshall (2002) indicates that the butterfly is “widespread,
and locally uncommon to common in much of northern Lower Peninsula, and in the Upper Peninsula”.
The dune isolate was reported by Wagner (1977), also by Nielsen from lakeside dunes and was found
utilizing Arabis lyrata (Opler, 1974; Clench & Opler, 1983). Nielsen (1999) notes: “Coastal specimens
are noticeably smaller than those found inland.”
• Minnesota: Macy & Shepard (1941) describe the habitat as “dry, stony hillsides near the banks of lakes
and streams” and also “scattered woods”. They report the hostplant along the St. Croix River in
Washington County as Arabis lyrata. Amhold (1952) describes olympia occurring on barren or thinly-
wooded rocky ridges above the Mississippi River. Masters reported the habitat to be deciduous woods in
east-central counties (in Clench & Opler, 1983), whereas Clench and Opler (1983) reported that the
species occurs on “open hilltops, surrounded by deciduous woods, overlooking the Mississippi River”.
Clench and Opler attribute those populations to the Great Lakes “inland isolate”. Birch, in Blackwater
Birds (2010) lists “sandy pine/oak barrens”. An online photo by A. Birch (2010) is captioned as “sandy
pine/oak barrens”. Patterson (2011) adds “sandy grasslands” to the list of habitats. A series of
specimens at the American Museum of Natural History represents a very small phenotype in Minnesota,
intermediate between nominotypical olympia and rosa , and approximate the size of individuals of the
Great Lakes Dune population (noting that Macy & Shepard (1941) reported utilization of A. lyrata in a
population of olympia in Washington County, MN. lends call for future investigation).
15
• Missouri: Meiners (1939, 1956) reports utilization of Arabis viridis (Green Rockcress) [synonym of
Arabis missouriensis (Missouri Rockcress)] in St. Louis county, and the habitat is described as tops of
high ridges with cherty soil and sparse vegetation, appearing barren in early spring, with adults rarely
straying to lower slopes (presumably forested habitat). Meiners (1939) also commented: “Later authors
are nearly agreed that rosa is merely a varietal form of olympiad Arnhold (1952) describes olympia
occurring “on the crests of narrow rocky ridges” in the Meramec Highlands near St. Louis. Opler &
Krizek (1984) list “river bluffs” as the habitat. Heitzman & Heitzman (1987) describe olympia as
occurring in scattered colonies in open woodlands, cedar glades and nearby meadows. I have found
olympia at a site in dense oak forest (early spring state) on a southward-facing hillside in St. Francois
county. In Wayne county, I found them in numbers on a shrubby, gravelly river flat, though I question
whether they are able to breed directly on the flat due to occasional flooding conditions.
• Montana: Barron (via butterfliesandmoths.org, accessed Dec. 1, 2018) described the habitat as mixed
prairie with ponderosa pines.
• Nebraska: Johnson (1973) describes olympia as a woodland species; occurring in pine forests from
Rock county westward; eastward in deciduous forests.
• New York: The habitat is known to be alvar barrens with sparse, stunted vegetation on limestone
pavement in Jefferson county. Dirig reports utilization of Arabis divaricarpa (Spreading-pod Rockcress
or Hybrid Rockcress) (in Parshall, 2002). Daniel (2018) photographed and posted to iNaturalist, a photo
of a larva feeding on what is identified as Boechera grahamii.
• North Carolina: The habitat in far western North Carolina is described as “typically open woods or
small openings within woods, in uplands, often on mountaintops” and “somewhat dry wooded slope”
(LeGrand & Howard, 2019). Cook, in Carolina Nature (2012) photographed larvae on Boechera
laevigata (Smooth Rockcress).
• North Dakota: Royer (1988, 2003) describes the habitat in western North Dakota as open prairies and
sage flats. He also reports them in “great numbers” some years, and uncommon in others.
• Ohio: The habitat is described as “dry ridgetops in and adjacent to open oak forests” (Iftner, et. al.
1992). Ohio specimens are reported to be nominotypical olympia.
• Oklahoma: McCoy reported the habitat to be bottomland deciduous forest (Clench & Opler, 1983).
This was identified as the Arkansas River isolate (Clench & Opler, 1983). Opler & Krizek (1984)
describe the habitat as “open river forests”. Harp (1986) reported larvae on Thelypodium wrightii
(Wright’s Thelypody). I have found them in central Oklahoma in a dry prairie remnant.
• Pennsylvania: Opler (1985) described the habitat as Devonian shale barrens, being found in “somewhat
open areas in the barrens and is usually found flying along ridge-tops or through clearings.”
• South Dakota: Reported to be local and uncommon, mostly in western South Dakota; isolated
occurrences in the east where native prairies remain; preferring short-grass to mid-grass prairies,
badlands, dry meadows, river breaks and rocky ridges in open pine forest (Marrone, 2002). Marrone
reports the hosts to be Arabis and Chorispora tenella (Blue Mustard or Crossflower).
• Tennessee: Venable (2014) indicates: “[T]hey seem to need areas with rocky outcrops, exposed
shale...”
• Texas: Freeman (1959) states that olympia was found on limestone outcroppings in the Dallas, TX.
area. These are identified as the Texas isolate by Clench and Opler (1983). Wauer (2006) describes the
habitat as open prairies and “juniper-dotted grassland”. I have found them in central Texas in a rather
ordinary weed field surrounded by deciduous forest. Neck (1996) lists the hosts as “ Arabis, Descaurania,
16
Sisymbrium ”; whereas the website of The Dallas County Lepidopterists’ Society (2015) specifically lists
Boechera canadensis (Sicklepod), Descurainia pinnata and Sisymbrium officinale.
• Virginia: Clark & Clark (1951) describe the habitat as “open woods, especially in the higher portions,
and along the rocky exposed crests of high ridges” (primarily in northern Virginia). In southwestern
Virginia, olympia has been found in limestone pavement habitat characterized by boulders and stunted
trees, especially Juniperus virginiana , and utilize the host Boechera laevigata (Clench & Opler, 1983).
Opler & Krizek (1984) list shale barrens as the habitat. Parshall (2002) also lists shale barrens and
indicates that Gypsy Moth spraying has impacted olympia. Virginia populations are nominotypical
olympia.
• West Virginia: Clark & Clark (1939) stated that olympia “kept to the crests of the ridges and to the
higher elevations generally, where it was very common...But in the bottoms of the valleys...absent.”
Clench and Opler (1983) describe the habitat as: “in or near open, low pine or pine-oak forests on south or
southwest facing rocky shale slopes”. Opler & Krizek (1984) list shale barrens as the habitat. Allen
(1997) refers to the primary habitat as shale barrens, specifically “semi-open, scrub oak-pine
habitats...where exposed shale slopes occur”, also in “semi-open wooded areas”. Arabis serotina (Shale
Barren Rockcress) is documented as a host in the shale barrens (Clench & Opler, 1983; Allen, 1997).
Allen (1997) also lists Boechera laevigata and, “to a lesser degree” Cardamine hirsuta (Hairy
Bittercress). W. H. Edwards (1897) reported a species of Sisymbrium [now split into many genera].
Populations in the northeastern part of the state have been heavily impacted by Gypsy Moth spraying
(Allen, personal communication). West Virginia populations are nominotypical olympia.
• Wisconsin: Amhold (1952) reports olympia occurring along the sides and top of a steep lakeside bank
within a “prairie-like” area north of Chippewa Falls. Ebner (1970) describes the habitat as “sandy
wastelands”. Masters (1973) reported olympia in “mixed forest areas of Transition zone character”.
Balogh (1980) reported olympia from sandy pine barrens, primarily dominated by Jack Pine ( Pinus
banksiana). Opler & Krizek (1984) list “river bluffs” as the habitat. The habitat is also described as Jack
Pine woods within a matrix of sand prairie (Minnesota Butterfly Walks, 2015). Swengel & Swengel
(1997) report olympia occurring in oak savannas or “barrens”, which consist of open woodlands with an
understory of plants associated with dry sandy prairies. Ebner (personal communication, 2010) described
in detail his experience collecting the dune isolate in 1953 at Kohler Andrae State Park, which he found in
close association with Arabis lyrata. Ferge (2011) describes the habitat as “dry sandy areas”. Macy and
Shepard (1941) report the hostplant along the Wisconsin side of the St. Croix River as Arabis lyrata.
Balogh (1980) also reports Arabis lyrata as the host in pine barrens.
• Wyoming: Ferris (1971) lists the habitat (or “ecological zone”) as moist meadows.
• Great Lakes region (Indiana/Illinois/Michigan/Wisconsin/Ontario): Douglas & Douglas (2005)
describe habitats in the Great Lakes region as: “dry areas, including prairie remnants, oak savannas, sand
dunes and dry lake plains...dry woodland forests or rocky prairies”. Distribution is described, as “highly
localized and small populations are the rule.”
• Kansas City region (Kansas/Missouri): Betros (2008) lists the following habitats in the greater
Kansas City region: woodlands/edges, overgrown fields, prairies and cedar glades.
• Oklahoma, Kansas, North Texas: Dole, et al. (2004) list “prairies, foothills, and open woodlands.
Canada :
• Alberta: Acorn (1993) describes olympia as being “most common in the valleys of the big prairie
rivers.” Bird, et al. (1995) give the habitat as “native prairie grassland and badlands.” Kondla (2004)
reports olympia as a “prairie butterfly...from a mid-elevation grassland adjacent [to] mountains” and also
17
specifically from “a site that had been very heavily grazed by livestock.” An anonymous member of the
flickr.com online photo gallery reported the habitat as “mountain meadows”. Kondla (pers. corr., 2020)
states that olympia is not known to occur in the Rocky Mountains of Alberta.
• Manitoba: Masters reported olympia on a hilltop in Jack Pine forest in the Riding Mountains (in Clench
& Opler (1983)). Klassen, et al. (1989) describe the habitat as “dry knolls and hills in prairie areas,
riverbank bluffs, woods and forests” and list the following hostplants: Sisymbrium officinale (Hedge
Mustard), Arabis glabra , Arabis lyrata and Arabis drummondi (Drummond’s Rockcress). [However, the
authors indicate that only three adult specimens were known from different localities in Manitoba, thus I
question whether the listed hosts were interpreted from prior published works.] Murray (2018) described
the habitat as “dry mixed grass sand prairie, pasture.”
• Ontario: Wagner (1977), reports the “inland form” from “Garden Township” [sic - correctly Carden
twp.] in open limestone barrens. Layberry, et al. (1982) describe the habitat in the Ottawa District as
“dry, barren, open areas”. Holmes, et al (1991) list “dry open areas in or near woodlands”. Layberry, et
al (1998) describe the habitat in Ontario as dry meadows, open woodlands, alvars and old dunes. Hall, et
al (2014) describe the habitat as: “Mostly very dry habitats, such as alvars, Canadian Shield rock barrens
and sandy meadows, and also open woodlands”. Lindsay (1971), Handfield (2011) and Hall, et al (2014)
reported utilization of Arabis lyrata. Layberry, et al (1982), Handfield (2011) and Hall, et al (2014) list
Arabis glabra. Layberry (1986) reported the host as Arabis divaricarpa. Handfield (2011) adds Arabis
drummondii. Ken Thome (pers. comm.) describes the dune habitat at the lower end of Lake Huron as
“sand dunes with Oak, Pine and Cedar”.
Data provided by the Ontario Butterfly Atlas Online (2016) provides considerable, more detailed habitat
and hostplant information. Various habitat descriptions can be summarized as: primarily open, dry
habitats with grassy areas and scattered, stunted shrubs, often in areas with exposed granite, limestone or
other rocks; rock outcrops and rock barrens; granite barrens; limestone alvars; semi-open pine-oak rock
barrens; sandy shrubland (sand barrens); wooded Canadian Shield country; and former pastureland. Host
records include: Arabis albida (Alpine Rockcress) [synonym of Arabis alpina ] and Arabis drummondi
both reported from Peterborough County (W. J. D. Eberlie), Arabis divaricarpa reported from Ottawa
City (Ross Layberry), Arabis glabra reported from Renfrew County (Ross Layberry); an unspecified
species of Boechera reported from Manitoulin District (Manitoulin Island) (Steven Daniel). These
records all apply to the eastward-expanding population of intermediate phenotypes. Additional records of
Arabis lyrata apply to the dune population in Lambton County (Nick Escott, Quimby Hess, Ken Thorne).
• Quebec: Layberry (1985) first reported the host as Arabis divaricarpa. Layberry, Hall & Lafontaine
(1998) describe the habitat in “Ontario and Quebec” as dry meadows and open woodlands, alvars and old
dunes [the “dunes” reference certainly pertains only to the dune-associated populations in Lambton
County, Ontario]. Handfield (2011) gives the habitat as uncultivated, open, dry places; also rocky
limestone or shale peaks and summits, more indicative of rosa habitats. Arabis glabra is listed as host.
Leboeuf & Le Tirant (2012), similarly list the habitat as: “open and dry habitats: rocky slopes, hilltops,
limestone outcrops” [translated from French text]. Handfield (2011) reports the first observation of
olympia in Quebec occurred on 5/29/1971.
• Saskatchewan: Hooper (1973) states that olympia “flies at the top of sharp knolls and badland hills.”
The hostplants are described as “Hedge Mustard” [Sisymbrium officinale ] and “Rockcress”, which can
apply to several species of Arabis or related genera.
• Eastern Canada: Acorn and Sheldon (2016) describe the habitat as “dry forests on sand or limestone
soils”.
18
• Canada: Layberry, et al. (1998) describe olympia as: “mainly a species of dryland areas. On the
Prairies, it is usually found on grassy knolls or around badlands.” The authors also document the
eastward range expansion [through southern Ontario and Quebec] since the 1970’s.
Wolfe, Harry & Stout (2010) recorded Isatis tinctoria (Dyar’s Woad) as an acceptable lab host.
Scott (1986b) noted that larvae refused Barbarea vulgaris (Wintercress or Yellow Rocket) in the lab.
One can readily conclude that the Appalachian and eastern Missouri populations have a greater
affinity for woodlands, while the Great Plains populations prefer prairie habitats. The Great Lakes dune-
associated populations stand out as having the most restricted range and being extremely habitat-specific.
Additionally, it might be noted that eastern populations in general are more localized and uncommon
while the western (Great Plains) population is somewhat more common and widely distributed.
PROPOSED TAXONOMIC REVISION
I propose the following subspecific divisions in Euchloe olympia. Subspecies olympia and rosa
are differentiated primarily by the degree of dark infuscation on the forewing apex and shape of the
forewing. Nominotypical olympia is essentially an Appalachian isolate, ranging westward into the Ohio
River drainage. Most of the isolates identified by Clench & Opler (1983) comprise the range of
subspecies rosa in the western (primarily Great Plains) portion of the species’ range. The Great Lakes,
Ontario, eastern Missouri and Ozark Regions encompass variable, intermediate populations. Great Lakes
Region variants more closely allied to nominotypical olympia are predominant east of Lake Michigan
while rosa phenotypes dominate west of the lake. Intermediates extend to the east through Ontario, into
Quebec and northern New York in a recent range expansion from the upper Great Lakes region.
Synonym anniha is attributed to the Great Lakes grouping, and is considered an available synonym of
rosa. The previously documented “Dune Isolate” or “Dune Form” (of authors) is differentiated as two
phenotypically distinct population groupings, primarily by their diminutive size and very limited range.
The lower Lake Michigan dune population is more closely allied with the rosa phenotype, while the lower
Lake Huron population is more closely allied with the nominotypical olympia phenotype; both are
proposed here as two distinct subspecies.
The county distributions below list where each actual phenotype was recorded, in an attempt to
define the range of the phenotypes, thus the subspecies. Percentages of each phenotype gleaned from
specimen series, published and internet resources were not recorded or estimated, as sampling error may
factor in. Therefore, one will note the nominotypical olympia and western rosa phenotypes may be listed
among the Great Lakes intermediate populations.
Euchloe olympia olympia (W. H. Edwards, 1871)
Description: Male FW length 15-21 mm (17.6 mm average); Female FW length 14-21 mm (18.8
mm average). I defer to the original description provided by W. H. Edwards (1871). The large gray patch
at the apex of the dorsal forewings characterizes the subspecies, and in Appalachian populations there is
little tendency for the patch to variably fade to white as in the Plains populations. The marbled markings
on the ventral hindwing tend to be more extensive than in Great Plains populations. Also, the forewings
tend to be slightly more rounded (the outer margin is convex) than Great Plains populations and many
specimens have dark marks in the shape of “V’s” along the dorsal hindwing outer margin where the wing
veins end (Fig. 6). The lectotype of Olympia is illustrated in F. M. Brown (1973) (Fig. 3).
19
I restrict the nominotypical subspecies to the
Appalachian region from southern Pennsylvania to
western North Carolina, eastern Tennessee, west
through central Kentucky to southern Indiana. This
corresponds with Clench & Opler’s Appalachian
isolate. Michigan, eastern Missouri and Ontario
populations, while consisting of variably mixed
phenotypes ranging from typical olympia to rosa ,
tend toward nominotypical olympia but are treated
separately as intermediate populations, below. The
common name “Olympia Marble” applies.
Confirmed distribution and/or specimens of nominotypical phenotype viewed or examined:
Kentucky: Bullitt Co., Edmonson Co. Illinois: Mason Co. Indiana: Clark, Morgan Co’s. Maryland:
Allegany, Washington Co’s. Michigan: Allegan, Benzie, Berrien, Cheboygan, Huron, Iosco, Kalkaska,
Montmorency, Muskegon, Newaygo, Oscoda, Otsego, Wexford Co’s. Missouri: St. Francois Co., St.
Louis Co., Wayne Co. New York: Jefferson Co. North Carolina: Madison Co. Ohio: Lawrence Co.
Ontario: Algoma Dist., Durham Regional Muni., Frontenac Co., Hastings Co., Kawartha Lakes Div.,
Lanark Co., Leeds and Grenville United Co., Lennox and Addington Co., Manitoulin Dist., Ottawa Div.,
Parry Sound Dist., Peterborough Co., Simcoe Co. Pennsylvania: Bedford Co. Quebec: Les Collines-
de-l’Outaouais Co., Temiscamingue Regional Co. Muni. Tennessee: Cocke, Greene, Jefferson, Sullivan
Co’s. Virginia: Frederick Co. A well-defined example can be found in Clark & Clark (1951) on plate
16, in which the entire apical area of the fore wings is very strongly marked, almost black. West
Virginia: Hampshire, Hardy, Kanawha, Mineral, Pendleton, Putnam Co’s. It is well illustrated in Allen
(1997), Plate 5, c?D, $D, c?V. Wisconsin: Columbia Co., Sauk Co.
Great Lakes, Ontario, eastern Missouri and Ozark region
intermediate and mixed populations
This is a difficult mix of populations, displaying a range of variation in the degree of infuscation
on the fore wing apex, and the shape of the fore wing. Many specimens display an intermediate character,
with a small gray subapical bar crossing the forewing, and the more angular wing shape of rosa.
Michigan “inland” populations, though highly variable, have a large proportion of specimens that are
closer to nominotypical (Appalachian) populations. Ebner (1970) interestingly noted: “Michigan
specimens are intermediate between the typical olympia and the western race rosa”. Wisconsin
populations vary considerably as well, though most specimens align more closely with rosa. Ebner’s
(1970) general description of Wisconsin populations also places them closer to rosa (and includes
synonym anniha ), but Ebner indicates: “Here and there an individual with more extensive black markings
also turns up, placing it more closely to typical olympia ”. The specimen illustrated on page 58 (Ebner,
1970) is clearly a rosa phenotype (fig. 4). Michigan (lower peninsula) specimens: Male FW length 16-20
Fig. 6. Typical Euchloe olympia olympia male
from Green Ridge State Forest Allegany Co. =
MD. 4/20/1985 : leg. Harry Pavulaan.
20
mm (18.0 mm average); Female FW length 15-21 mm (19.1 mm average). Wisconsin specimens: Male
FW length 15-18 mm (17.0 mm average); Female FW length 17-20 mm (18.0 mm average).
The eastern Missouri (St. Louis area) populations are classified primarily as intermediates but
contain a high percentage of individuals with very
heavy apical infuscation similar to Appalachian
populations, and also individuals with development
of a distinct diagonal gray subapical band (fig. 7).
Interestingly, the eastern Missouri olympia are the
largest that I have examined, among all E. olympia
populations. Male FW length 17-20 mm (18.9 mm
average); Female FW length 18-21 mm (19.5 mm
average). Clench & Opler (1983) noted these
characters for the ‘St. Louis isolate’. Specimens
and images from the Ozark region were too few to
conclude on regional alignment with either
subspecies, but appear to range from nominotypical
olympia to rosa.
Inclusion of the northern New York
population with this grouping is based on the
presence of a mix of nominotypical olympia and
rosa phenotypes, indicative of the well-documented, rapid range expansion of olympia from the northern
Great Lakes region eastward across southeastern Ontario (first reported expanding eastward in 1966
at Killaloe, Ontario) and then into southern Quebec by 1971. This range expansion extended as far east as
Portneuf, Quebec by 1978 and then southward into northern New York by 1986. I exclude the Great
Lakes “dune” populations from this grouping.
Not considered part of the Great Lakes, Ontario, eastern Missouri and Ozark Region intermediate
population grouping, are additional scant records of intermediate phenotypes mainly at the eastern edge of
the range of rosa , or at high elevations in Colorado, which are essentially rosa variants. These are listed
separately, below.
Confirmed distribution and/or specimens of intermediate and mixed phenotypes viewed or
examined:
Arkansas: Garland, Hot Spring, Pulaski Co’s. Illinois: Mason Co. Kentucky: Edmonson Co.
Michigan: Allegan, Arenac, Crawford, Huron, Iron, Kalkaska, Montcalm, Muskegon, Oscoda,
Roscommon Co’s. Minnesota: Anoka, Hubbard, Sherburne, Wabasha Co’s. Missouri: St. Francois, St.
Louis, Wayne Co’s. New York: Jefferson Co. Ontario: Algoma Dist., Durham Regional Muni.,
Frontenac Co., Kawartha Lakes Div., Leeds and Grenville United Co., Manitoulin Dist., Ottawa Div.,
Parry Sound Dist., Renfrew Co., Simcoe Co. Quebec: Communaute-Urbaine-de-l’Outaouais Co., Les
Collines-de-l’Outaouais Co. Wisconsin: Adams, Bayfield, Burnett, Columbia, Dodge, Dunn, Grant,
Jackson, Juneau, Marinette, Marquette, Sauk, Waushara, Wood Co’s.
Confirmed distribution of intermediate and mixed phenotypes outside the Great Lakes and
eastern Missouri region:
Occasional specimens of intermediate character, essentially rosa based primarily on the more
angular forewing shape, but with a heavier degree of apical infuscation occur throughout much of the
range of rosa. These are considered individual variants of rosa. However, a higher percentage of
intermediates of diminutive size occur with greater frequency in central Colorado, including some with
Fig. 7. Intermediate phenotype (male) from
Patterson. Wayne Co., MO. 3/26/1988 ; leg. Harry
Pavulaan.
very dark, almost blackened apical patches, likely affected by elevation. Clench & Opler (1983)
described these as the ‘Front Range Isolate’ but are primarily rosa in nature.
Colorado: Boulder, Jefferson, Park, Pueblo Co’s. Iowa: Fremont Co. Kansas: Greenwood Co.
Missouri: Jefferson, St. Francois Co., St. Louis, Wayne Co. Nebraska: Dawes Co., Otoe Co. South
Dakota: Lyman Co. Texas: McLennan Co.
Euchloe olympia rosa (W. H. Edwards, 1882) - revised status
Description: Male FW length 14-20 mm (18.0 mm average); Female FW length 15-22 mm (18.5
mm average). I defer to the original description provided by W. H. Edwards (1882). The subspecies is
characterized by the mostly white apex of the dorsal
forewings, in which the gray apical clouding that is
characteristic of nominotypical subspecies olympia
is reduced to two small dark patches situated along
the costal and outer wing margins, at both ends of
the subapical area. A variable gray subapical bar in
some specimens may connect these patches, but this
character is more frequent in the Great Lakes region
west of Lake Michigan. The marbling on the
ventral hindwing tends to be slightly reduced in
comparison to nominotypical olympia. Also, the
forewings are somewhat more angular (pointed)
than nominotypical olympia , with the outer margin
being either straight, or having a very slight
concavity in many specimens (fig. 8). The
lectotype of rosa is illustrated in F. M. Brown
(1973) (Fig. 3).
I restrict subspecies rosa to the Great Plains region from Texas to Alberta. This corresponds with
Clench & Opler’s ‘Texas’, ‘Arkansas River’, ‘Missouri River’ and ‘Front Range’ isolates. Of interest are
two distinctly rosa specimens from Arizona in the National Museum of Natural History collection
labeled: Chiricahua Mtns., Cochise Co., 4/29/1979 (J. Legge); and Baboquivari Mtns., Pima Co., June
1923 (Barnes collection). I have no reason to doubt they are authentic but further inquiry is needed to
determine the circumstances of these specimens. I propose the common name “Rosy Marble” be
restricted to subspecies rosa , primarily to reflect the latin name and not the ventral slightly “rosy” tint
which also occurs in nominotypical olympia.
Confirmed distribution and/or specimens viewed or examined:
Alberta: Brooks, Calgary, Cavendish, Champion, Drumheller, Empress, Finnegan, Jenner, Lethbridge,
Medicine Hat, Redcliff, Waterton Park, Veteran, Writing-on-Stone Provincial Park. Arkansas: Carroll,
Hot Spring, Pulaski Co’s. Arizona: Cochise, Pima Co’s. Colorado: Arapahoe, Boulder, Denver,
Dolores, Douglas, El Paso, Jefferson, Larimer, Pueblo, Weld, Yuma Co’s. Illinois: Mason Co. Iowa:
Fremont, Woodbury Co’s. Kansas: Cowley, Douglas, Franklin, Greenwood, Harper, Johnson,
Leavenworth, Pottawatomie, Reno, Riley, Rooks, Scott, Sumner Co’s. Manitoba: Aweme, Spruce
Woods Provincial Park. Michigan: Cheboygan, Huron, Montmorency, Muskegon, Otsego Co’s.
Minnesota: Crow Wing, Hubbard, Mille Lacs, Sherburne, Wadena, Washington Co’s. Missouri:
Benton, Jackson, Reynolds, St. Louis Co’s. Montana: Blaine, Cascade, Custer, Fallon, Fergus, Gallatin,
Golden Valley, Hill, Musselshell, Park, Prairie, Rosebud, Sweet Grass, Yellowstone Co’s. Nebraska:
Cass, Cherry, Dodge, Douglas, Lancaster, McPherson, Nemaha, Otoe, Platte, Saunders, Sioux, Thomas
Co’s. North Dakota: Billings, Slope Co’s. Oklahoma: Cleveland, Comanche, Le Flore, Murray,
Oklahoma, Payne, Texas, Tillman, Washita, Woodward Co’s. Ontario: Algoma Dist., Hastings Co.,
22
Wise Co. : TX. 3 / 23 / 2015 , leg. Harry Pavnkan.
Leeds and Grenville United Co’s., Muskoka Dist., Ottawa Div., Parry Sound Dist., Peterborough Co.
Quebec: Communaute-Urbaine-de-l’Outaouais Co., Les Collines-de-l’Outaouais Co. Saskatchewan:
Big Beaver, Cypress Hills, Divide, Regina. South Dakota: Brookings, Lyman, Pennington Co’s. Texas:
Carson, Comanche, Cottle, Crosby, Dallas, Ellis, Hale, Kent, Lynn, Mason, McLennan, Motley, Randall,
Smith, Wichita, Wise Co’s. Wisconsin: Adams, Burnett, Clark, Dane, Douglas, Dunn, Eu Claire, Grant,
Jackson, Juneau, Marinette, Menominee, Oneida, Polk, Sauk, St. Croix, Washburn, Waushara, Wood
Co’s. Wyoming: Natrona.
Wagner’s “Dune Form”
Wagner’s (1997) description: I refer here to the original description provided by Wagner (1997),
which differentiates the “Dune Form” from the “Inland Form” of Michigan, the latter of which I consider
to be highly variable and intermediate between nominotypical olympia and subspecies rosa. Wagner first
gives a brief comparison: “The Dune Form is readily distinguished from the Inland Form by its smaller
size, whiter ground color, and more complex pattern of markings on the secondaries below. The Lake
Michigan Dune populations are not quite so extremely differentiated as the Lake Huron.” This latter
comment is important. Wagner’s original full description is as follows:
KEY TO DUNE AND INLAND FORMS OF EUCHLOE OLYMPIA
Smaller, forewing averaging 16-18 mm in length; ground color chalky white; forewing
slightly more rounded, the fringe distinctly checkered dorsally from veins M3 to R2; dark
markings above more “smudged,” more confluent; basal dark areas on dorsal surfaces
darker and more extensive, running to and commonly beyond the medial transverse dark
band of the ventral surface (as visible through the wing); black patch at end of M3 on dorsal
surface of forewing usually similar to those on M2 and Mi; dark bands on ventral surface of
hind wings below usually darker and greener, wider and more irregular with more
projections and hair-line; Cu2 usually with a dark terminal patch 1-2 mm long
.DUNE FORM
Larger, forewing averaging 18-20 mm in length; ground color pale ivory-white; forewing
slightly more pointed, the fringe usually only weakly checkered to lacking any black scales
dorsally from veins M3 to R2; dark markings above not so “smudged,” more discrete; basal
dark areas on dorsal surfaces paler and smaller, running only as far as the postbasal
transverse dark band on the ventral surface (as visible through the wing); black patch at end
of M3 on dorsal surface of forewing usually darker and more conspicuous than those on M2
and Mi; dark bands on ventral surface of hind wings below usually paler and yellower,
narrower and more regular with fewer projections and hair-lines; Cu2 usually with terminal
patch vestigial, less than 1 mm long.INLAND FORM
Wagner’s key to the “Dune Form” serves as the baseline description of what I propose as two
distinct, allopatric “dune” subspecies, each representing the lower Lake Michigan and lower Lake Huron
population groupings. Wagner, however, grouped both the Lake Michigan and Lake Huron populations
together as a single “Dune Form” entity, while each is decidedly differentiated from the other in extent of
ventral markings. Most of the biological information provided by Wagner and earlier workers pertains to
the lower Lake Michigan population cluster. Less is known of the Lake Huron population.
Wagner noted greater similarities to subspecies olympia [than to the “Inland Form”] except that
the ventral markings in subspecies olympia are not as intricate and extensive, and that nominotypical
olympia in West Virginia is “of decidedly larger dimensions”. Wagner & Hansen (1980) noted that “the
dunes form of the marbled white, Euchloe olympia (Edwards) runs smaller in Berrien County [Michigan]
(also in Lambton County, Ontario) than does the inland form of savannahs and prairies which flies at the
same time (late April, May). The smaller size of the dunes form of E. olympia may be a consequence of
23
unusual climatic and edaphic conditions in the dunes: exposure, winds, permeable sands, and lowered
growth potential in food plants.” There has yet been no published account of any attempts to rear “dune”
vs. “inland” populations under identical, artificial conditions to determine if this is a factor. Interestingly,
according to Wagner (1977), the closest that the dune populations come, in distance, to inland populations
of olympia in Michigan is 50 miles! He suggested that, with future efforts, populations of the inland form
might be found closer to the dune populations, possibly coming into contact and intergrading. To date, no
such contact has been documented.
Of interest are previous illustrations of the Dune populations in the literature: Klots (1951)
illustrates a specimen of olympia from “northeastern Illinois” on Page 208 (no. 6), most certainly based
on specimens with similar labels in the NMNH (collected by H. M. Bower) which are determined to be of
dune habitat origin. Klassen, et. al. (1989) illustrate a pair of specimens taken at Ipperwash, Ontario
(lower Lake Huron population) on plate 9 (figs. 15 & 16). Opler & Malikul (1992) illustrate a specimen
of olympia from Lake County, IL. on plate 17. Nielsen (1999) illustrates a “Coastal male” in the plate on
Page 49 (collected at Grand Mere State Park, Berrien Co. ML). Most recently, the specimens illustrated
in Belth (2013) on pages 15, 133 (egg; fig. 4), 226 and 227 were photographed in Porter County, IN.
Jeffrey Belth (personal comm.) reported that the Porter County images were taken at Indiana Dunes
National Lakeshore, thus representing the lower Lake Michigan population grouping.
Euchloe olympia wagneri - new subspecies
ZooBank registration: urn:lsid:zoobank.org:act:7922ElE7-E749-49D3-8E4F-lF627B41481F
Description: (Figs. 10, 11, 12). Male FW length 14-18 mm (15.8 mm average); Female FW
length 15-19 mm (17.0 mm average). Primary phenotypic difference is consistently smaller size in
comparison to either subspecies olympia and rosa. Similar in size to subspecies huron (described below).
Similar to the ventral hindwing pattern of rosa\ and dorsally in the extent of black markings at the
forewing apex, though many individuals have slightly greater apical infuscation. The basal dark areas on
the dorsal surfaces are considerably darker and more extensive than in rosa. The ventral hind wing
“marbling” pattern is relatively narrow and well-defined. The color of the marbling is yellowish-gray
(appearing green). Interestingly there appears to be less individual variation within the lower Lake
Michigan populations than within adjacent “inland” populations, with few individuals approaching the
olympia form in extent of black markings at the FW apex.
Host: The only documented host for this subspecies, in Indiana (C. A. Shull, 1907) and Michigan
(Nielsen, 1999), is Arabis lyrata (Fig. 13). Ebner (personal communication, 2010) indicated a close
association of Wisconsin lakeside dune populations with Arabis lyrata , though no direct ovipositions
were recorded. Hostplant choice may be further indicative of ecological differences from inland
populations that might be considered diagnostic. Wagner (1977) however notes that the range of Arabis
lyrata extends over a broad area covering Michigan, Wisconsin, northern Indiana and northern Illinois,
and concludes: “therefore, it seems unlikely that it is foodplant preference alone that governs the
differentiation of the Dune populations”.
Immature stages: C. A. Shull (1907) described the immature stages of the dune population at the
southern end of Lake Michigan in Indiana. He described the eggs as “orange yellow”, deposited singly
“on the sepals of the younger, centrally-located buds, rarely on the peduncle just below the bud”; usually
only one, very rarely more than two, eggs per plant. When ready to hatch, the eggs turn “dull yellowish
brown”. C. A. Shull collected eggs on May 12, 1905. Belth (2013) illustrates an egg on page 133 (fig. 4).
The newly hatched larvae measure “a little over 1 mm” but less than 1.5 mm., which eat away
only one side of the egg shell, and immediately begin feeding on the host Arabis lyrata. C. A. Shull noted
the length of time between moults, and based his observations on a captive larva that hatched on May 13.
The first molt occurred on May 16, the second on May 18, the third on May 20 and the fourth on May 22.
24
The fully developed larva measured 28 mm., and on May 25, left its host and began searching for a place
to pupate. Pupation occurred on May 26. C. A. Shull noted that the entire larval period of his captive
stock measured 12-14 days, though he indicated that by May 26, wild larvae were found in various stages
from second to fifth instars. He attributed this to the time period over which eggs were laid, cooler nights
in the habitat, and “less abundant food” in nature. By the time of a visit to the habitat on June 1, a single
fully grown larva was located.
C. A. Shull noted changes in larval color as they grew. Newly hatched larvae were lemon-yellow
with a black head. After the first molt, they were green with the head being greenish-gray. After the
second molt, the larval colors become “fully developed” with dorsal and lateral lines and become brighter
and more contrasted in the fourth and early fifth instar. Toward the end of the fifth instar, the “brilliancy
of the color pattern is lost” and the larvae develop a purplish color, becoming very dark just prior to
pupation. The freshly formed pupa is described as rosy purple, changing to various shades of brown
within a few hours.
C. A. Shull removed the fully-grown larva of June 1 from its host, placed it on the ground and
followed it to see where it would pupate. The larva entered a clump of Andropogon scoparius grass,
abundant in the habitat, and chose a pupation site within the grass clump. A return visit on June 9 found
the pupa that had formed.
Fight Period: April 14 - May 31 (Fig. 9).
200
130
180
170
160
150
140
130
120
110
100
30
80
70
60
50
40
30
20
10
early : April 14 late: May 31
Fig. 9: Flight phenogram for Euchloe olympia wagneri rangewide. Bars represent total cumulative
reported count for each date.
Apri L*
—
It:
1
. I |p|
.
Jim.
Habitat: Subspecies wagneri is endemic to lakeshore dune habitat around the south end of Lake
Michigan in the United States. The habitat consists of ancient dunes with extensive open, sandy areas,
with varied growth of coniferous and deciduous trees (Fig. 14).
Confirmed range and/or specimens examined: The following locations comprise the known
distribution:
25
Illinois: Cook Co.: Chicago (leg. O. Buchholz). Lake Co.: “northeastern Lake County” (leg. H. M.
Bower); Illinois Beach State Park (legs. W. J. Gerhard, I. Leeuw, R. Leuschner, E. Liljablad, J. Skrentny
[photos], P. Sweet [photos], D. Taron [photos]); Waukegan (legs. I. Leeuw, A. K. Wyatt); Waukegan
Dunes (leg. A. K. Wyatt). Indiana: Lake Co. (leg. W. H. Wagner); Gary “sand dunes” (G. Wren).
Porter Co. (leg. C. A. Shull); Dune Acres (leg. W. Henderson); Indiana Dunes National Lakeshore
(Bentley, 2010; Belth, 2013; Sass, 2015); Indiana Dunes State Park (M. Beatty [photo]). Michigan:
Berrien County: New Buffalo (leg. W. H. Wagner); Grand Mere Lakes Dunes (leg. M. Nielsen).
Wisconsin: Sheboygan Co.: Kohler Andrae State Park (leg. James Ebner). Milwaukee Co.: Texas Rock
(leg. not indicated).
Etymology: The subspecies is named in honor of Warren Herb Wagner, Jr., who described this
distinct taxon but declined to name it. I propose the common name “Lake Michigan Dune Marble”.
Holotype, allotype and paratypes:
Holotype: Male (Fig. 10), IL., Lake Co., I lli nois Beach State Park, May 15, 1951 (leg. R. Leuschner).
Specimen resides in the National Museum of Natural History, Washington, D.C. Allotype female (Fig.
11), Waukegan, Lake Co., IL., USA. 5/14/1988, leg. Irwin Leeuw. Specimen resides in the present
author’s collection.
-f - » ?
/ v -
f
Fig. 10. Holotype male, Euchloe olympia wagneri (Pavulaan, 2020). Dorsum (left), venter (right).
Photograph by permission of the Smithsonian National Museum of Natural History. [Note distinct
ventral chain-like marbling]
Fig. 11. Allotype female, Euchloe olympia wagneri (Pavulaan, 2020). Dorsum (left), venter (right).
Waukegan, Lake Co., IL., USA. 5/14/1988, leg. Irwin Leeuw. [Note distinct ventral chain-like
marbling]
26
Paratypes in the Field Museum, Chicago, IL:
IL., Cook Co., Chicago, May 13, 1917 (leg. A. K. Wyatt).
IL., Lake Co., “Beach” (Illinois Beach?), May 21, 1922 (leg. W. J. Gerhard, 5 specimens).
IL., Lake Co., Illinois Beach State Park, May 18, 1972 (G. Nielsen, 2 specimens).
IL., Lake Co., Waukegan, May 17, 1931 (leg. A. W. Herz, 2 specimens).
IL., Lake Co., Waukegan, May 15, 1932 (leg. A. W. Herz, 3 specimens).
IL., Lake Co., Waukegan, May 23, 1950 (leg. Banks).
IL., Lake Co., Waukegan, May 28, 1960 (leg. D. H. Kistner).
IN., Lake Co., East Gary, May 11, 1913.
IN., Lake Co., Hammond, May 10, 1903 (leg. Sala).
Paratypes in the McGuire Center for Lepidoptera and Biodiversity, Gainesville, FL:
IL., Lake Co., I lli nois Beach State Park, May 15, 1951 (2 specimens).
IL., Lake Co., Illinois Beach State Park, May 5, 1974.
IL., Lake Co., Illinois Beach State Park, May 19, 1974.
IL., Lake Co., Illinois Beach State Park, May 14, 1988 (leg. Irwin Leeuw).
IL., Lake Co., Waukegan, May 11, 1991 (leg. Irwin Leeuw).
IN., Lake Co., “Duros State Park”, May 7, 1959.
Paratypes in the Museum of Comparative Zoology, Cambridge, MA.:
IL., Lake Co., Waukegan, May 18, 1950 (leg. A. K. Wyatt), accession #MCZ-ENT 00134124.
IL., Lake Co., Waukegan, May 18, 1950 (leg. A. K. Wyatt), accession #MCZ-ENT 00134126.
Paratypes in the National Museum of Natural History, Washington, D.C.:
IL., Lake Co., May 17, 1931 (leg. H. M. Bower, 6 specimens).
IL., Lake Co., Illinois Beach State Park, May 20, 1930 (leg. E. Liljablad).
IL., Lake Co., Illinois Beach State Park, May 13, 1931.
IL., Lake Co., Illinois Beach State Park, May 15, 1951 (R. Leuschner, 2 specimens).
IL., Lake Co., Illinois Beach State Park, May 15, 1951 (leg. E. Liljablad, 2 specimens).
IL., Lake Co., Illinois Beach State Park, May 9, 1973 (I. Leeuw, 2 specimens).
IL., Lake Co., Illinois Beach State Park, May 20, 1973 (I. Leeuw).
IL., Lake Co., Waukegan, May 17, 1931 (leg. A. K. Wyatt).
IL., Lake Co., Waukegan, May 14, 1933 (leg. A. K. Wyatt).
IL., Lake Co., Waukegan Dunes, May 17, 1931 (leg. A. K. Wyatt, 3 specimens).
IL., Lake Co., Waukegan Dunes, May 15, 1932 (leg. A. K. Wyatt).
IL., Lake Co., Waukegan Dunes, May 14, 1933 (leg. A. K. Wyatt, 3 specimens).
IN., Lake Co., Gary, May 3, 1908 (leg. A. K. Wyatt).
IN., Lake Co., Miller (Miller Beach), May 13, 1917 (leg. A. K. Wyatt, 3 specimens).
IN., Lake Co., Miller (Miller Beach), April 27, 1919 (leg. A. K. Wyatt, 4 specimens).
IN., Lake Co., Miller (Miller Beach), May 11, 1922 (leg. E. Liljeblad, 3 specimens).
IN., Lake Co., Miller (Miller Beach), May 4, 1924 (leg. W. J. Gerhard, 10 specimens).
IN., Lake Co., Miller (Miller Beach), May 30, 1924 (leg. W. J. Gerhard, 3 specimens).
IN., Lake Co., Miller (Miller Beach), April 14, 1929 (leg. A. W. Herz).
Paratypes in the Peabody Museum of Natural History, New Haven, CT.:
IL., Lake Co., Illinois Beach State Park, (undated), accession #YPM ENT 433066.
IL., Lake Co., Illinois Beach State Park, May 16, 1971 (leg. I. Leeuw),
accession #YPM ENT 433067.
IL., Lake Co., Illinois Beach State Park, May 10, 1971 (leg. I. Leeuw),
accession #YPM ENT 433068.
IL., Lake Co., Zion, Illinois Beach State Park, (undated), accession #YPM ENT 569514.
IL., Lake Co., Zion, Illinois Beach State Park, (undated), accession #YPM ENT 569515.
IL., Lake Co., Zion, Illinois Beach State Park, (undated), accession #YPM ENT 569516.
IL., Lake Co., Zion, Illinois Beach State Park, (undated), accession #YPM ENT 570386.
IL., Lake Co., Zion, Illinois Beach State Park, (undated), accession #YPM ENT 570387.
27
IL., Lake Co., Illinois Beach State Park, May 16, 1971 (leg. I. Leeuw),
accession #YPM ENT 570388.
IL., Lake Co., Zion, Illinois Beach State Park, (undated), accession #YPM ENT 570389.
IL., Lake Co., Illinois Beach State Park, May 10, 1971 (leg. I. Leeuw),
accession #YPM ENT 570390.
IL., Lake Co., Illinois Beach State Park, May 15, 1971 (leg. I. Leeuw),
accession #YPM ENT 724214.
IL., Lake Co., Illinois Beach State Park, May 15, 1971 (leg. I. Leeuw),
accession #YPM ENT 724215.
IL., Lake Co., Illinois Beach State Park, May 15, 1971 (leg. I. Leeuw),
accession #YPM ENT 724219.
IL., Lake Co., Illinois Beach State Park, May 15, 1971 (leg. I. Leeuw),
accession #YPM ENT 724220.
Paratypes in the collection of Andrew D. Warren, Castle Rock, Colorado, USA:
IL., Lake Co., Illinois Beach State Park, May 11, 1991 (leg. I. Leeuw, 18 specimens).
IL., Lake Co., Illinois Beach State Park, May 8, 1992 (leg. I. Leeuw, 6 specimens).
IL., Lake Co., Illinois Beach State Park, May 31, 1992 (leg. I. Leeuw, 3 specimens).
IL., Lake Co., Illinois Beach State Park, May 7, 1993 (leg. I. Leeuw, 8 specimens).
IL., Lake Co., Illinois Beach State Park, April 30, 2002 (leg. I. Leeuw, 32 specimens).
IL., Lake Co., Illinois Beach State Park, May 4, 2002 (leg. I. Leeuw, 16 specimens).
Paratypes in the collection of the present author:
IL., Lake Co., Waukegan, May 17, 1931 (leg. I. Leeuw, 4 specimens).
IL., Lake Co., Illinois Beach State Park, April 17, 1976 (leg. I. Leeuw).
IL., Lake Co., Illinois Beach State Park, May 9, 1987 (leg. I. Leeuw).
IL., Lake Co., Illinois Beach State Park, May 7, 1988 (leg. I. Leeuw).
IL., Lake Co., Illinois Beach State Park, May 14, 1988 (leg. I. Leeuw, 4 specimens incl. allotype) .
IL., Lake Co., Illinois Beach State Park, May 15, 1988 (leg. I. Leeuw, 4 specimens).
IL., Lake Co., Illinois Beach State Park, May 11, 1991 (leg. I. Leeuw, 19 specimens).
IL., Lake Co., Illinois Beach State Park, May 8, 1992 (leg. I. Leeuw, 10 specimens).
Fig. 12. Euchloe olympia wagneri resting, Miller Beach, Fig. 13. Host Arabis lyrata. [Photo © by courtesy of
IN. (left). [Photo © by courtesy of Jeffrey Belth] Jeffrey Belth]
28
Fig. 14. Euchloe olympia wagneri habitat at Miller Beach, IN. [Photo © by courtesy of Jeffrey Belth]
Euchloe olympia huron - new subspecies
ZooBank registration: urn:lsid:zoobank.org:act:691D3ED0-CA09-4E7D-B7C4-71D4BCC8758B
Description: (Figs. 16, 17). Consistently smaller size in comparison to either subspecies olympia
and rosa. Male FW length 14-18 mm (15.9 mm average); Female FW length 14-19 mm (16.3 mm
average). Similar to subspecies olympia in extent of black markings at the dorsal forewing apex, but
dissimilar from either rosa or wagneri , in which both the black subapical bar is narrowed or reduced
(mainly in rosa) to two black subapical spots. The ventral hindwing is most distinct from all other
olympia subspecies, being characterized by an enlarged, smeared, and more complex pattern of yellowish
gray patterns (Figs. 16, 17, 18, 19). This pattern is differentiated from subspecies wagneri by being
ventrally paler overall.
Host: The only documented host for this subspecies (in Ontario) is Arabis lyrata (Wagner, 1977;
Hall, et al. , 2014; J. Cossey, photos; K. Thorne, pers. comm.).
Fight Period: April 14 - June 13. Notable maximum single day report of 200 on April 29, 1990
by the Toronto Entomologists’ Association, with cumulative maximum of 208 reported for April 29 (Fig.
15).
29
April max. 208
May
June
early: April 14 late: June 13
Fig. 15. Flight phenogram for Euchloe olympia Huron at Lambton County, Ontario. Bars represent total
cumulative reported count for each date.
Habitat: Subspecies Huron is endemic to lakeshore dune habitat around the south end of Lake
Huron in Canada. The habitat is confined to a very limited area and consists of ancient dunes with
extensive open, sandy areas, but overgrown with Junipers, Pines and Oaks (Fig. 20). The butterfly
appears to be reasonably more common in this area than in other portions of Euchloe olympiads range.
Confirmed range and/or specimens examined: Essentially ah known locations are within two
miles of Lake Huron in Lambton County, Ontario. The following locations comprise the known
distribution based on photos and specimens:
Ontario: Lambton Co.: Ausable River Cut Conservation Area (R. McLeod); Grand Bend (Q. F. Hess, D.
Murphy); Ipperwash (G. Clements, J. Cossey, B. Curry, Q. F. Hess, J. Holds worth, K. Hoo, B. Kulon, B.
Lamond, A. Rocchi, T. Sabo, J. Skevington, K. Stead, K. Thorne, J. Troubridge, G. Vogg, W. H. Wagner,
B. Yukich, K. Zufelt); Ipperwash Provincial Park (J. A. Brown, B. Curry, Q. F. Hess, B. Kulon, J. Larson,
S. T. Pike, J. E. Pilkington, T. Sabo, G. Sutton, W. H. Wagner, C. Wood, S. Wood, A. Wormington);
Karner Blue Sanctuary (P. Deacon); Kettle Point Indian Reserve 44 (J. Cossey, A. H. Rider); Lambton
Shores (anonymous, iNaturalist); Pinery (Q. F. Hess, B. Kulon, J. Skevington, T. Toyomi); Pinery
Provincial Park (P. Carter, G. Clements, J. Cossey, T. Crabe, S. Daniels, P. Deacon, W. Gregory, Q. F.
Hess, A. M. Holmes, B. Kulon, L. MacGregor, A. MacKenzie, S. McEwan, B. Ottaway, J. E. Pilkington,
M. van der Poorten, N. van der Poorten, R. E. Scott, D. Scovell, J. Skevington, G. Sutton, K. Thome, W.
H. Wagner, J. Walty, J. G. Woods, K. Zufelt); Port Franks (B. A. Mann, G. Clements, B. Curry, P.
Deacon, Q. F. Hess, B. Kulon, B. Lamond, J. Larson, B. Mann, S. T. Pike, T. Sabo, K. Sealy, K. Stead, K.
Thome, A. Wormington, K. Zufelt).
Etymology: The subspecies is named to reflect its association with Lake Huron. I propose the
common name “Lake Huron Dune Marble”.
30
Holotype, allotype and paratypes:
Holotype: Male (Fig. 16). ON., Lambton Co., Ipperwash, May 12, 1983 (leg. K. Thorne). Specimen
resides in the National Museum of Natural History, Washington, D.C. Allotype female (Fig. 17),
Ipperwash, Lambton Co., ON., Canada, 5/7/2015, leg. K. Thorne. Specimen resides in the present
author’s collection.
fP
, <Sk
w *
> . t <
- /V .. *
H
E? ,-f K
i
Pieridae V-
12-1981
Euchloe olympia rosa M
Ipperwash, Lambton Co,
ONT, Canada. K. Thome
Fig. 16. Holotype male, Euchloe olympia Huron (Pavulaan, 2020). Dorsum (left), venter (right).
Specimen is identified as “Euchloe olympia rosa ” on the original label. Photograph by permission
of the Smithsonian National Museum of Natural History. [Note distinct heavy ventral marbling]
Fig. 17. Allotype female, Euchloe olympia Huron (Pavulaan, 2020). Dorsum (left), venter (right).
Ipperwash, Lambton Co., ON., Canada. 5/7/2015, leg. K. Thorne. [Note distinct heavy ventral
marbling]
Paratypes in the Lyman Museum, Hilo, HI:
ON., Lambton Co., Ipperwash, April 22, 1986 (leg. W. H. Wagner).
Paratypes in the McGuire Center for Lepidoptera and Biodiversity, Gainesville, FL:
ON., Lambton Co., Ipperwash, May 7, 2015 (leg. K. Thome, 2 specimens).
Paratypes in the National Museum of Natural History, Washington, D.C.:
ON., Lambton Co., Ipperwash, May 10, 1980 (leg. K. Thorne, 3 specimens).
ON., Lambton Co., Ipperwash, May 21, 1980 (leg. K. Thorne).
ON., Lambton Co., Ipperwash, May 12, 1983 (leg. K. Thorne, 3 specimens).
31
Paratypes in the collection of Ken Thorne:
ON., Lambton Co., Ipperwash, May 2, 1978 (leg. K. Thome, 2 specimens).
ON., Lambton Co., Ipperwash, May 19, 1979 (leg. K. Thorne, 2 specimens).
ON., Lambton Co., Ipperwash, May 20, 1979 (leg. K. Thorne).
ON., Lambton Co., Ipperwash, May 22, 1979 (leg. K. Thorne).
ON., Lambton Co., Ipperwash, May 23, 1979 (leg. K. Thorne, 2 specimens).
ON., Lambton Co., Ipperwash, May 23, 1984 (leg. K. Thorne, 2 specimens).
ON., Lambton Co., Ipperwash, May 8, 1999 (leg. K. Thome, 2 specimens).
ON., Lambton Co., Ipperwash, May 7, 2015 (leg. K. Thome, 19 specimens).
ON., Lambton Co., Ipperwash, April 27, 2017 (leg. K. Thorne, 30 specimens).
ON., Lambton Co., Pinery Provincial Park, May 3, 1970 (leg. K. Thorne, 10 specimens).
ON., Lambton Co., Pinery Provincial Park, May 7, 1971 (leg. K. Thorne, 5 specimens).
Paratypes in the collection of the present author:
ON., Lambton Co., Ipperwash, May 11, 1980 (leg. J. Troubridge, 6 specimens).
ON., Lambton Co., Ipperwash, May 7, 2015 (leg. K. Thome, 14 specimens incl. allotype) .
ON., Lambton Co., Ipperwash, April 27, 2017 (leg. K. Thorne, 8 specimens).
Fig. 18. Euchloe olympia Huron nectaring on hostplant Arabis lyrata, Ipperwash Beach, ON. (left).
Euchloe olympia Huron chrysalis, same location (right). [Photos © by courtesy of Jay Cossey]
Fig. 19. Euchloe olympia Huron larva on hostplant Arabis lyrata, Pinery Provincial Park, ON. (left). Euchloe
olympia Huron ventral hindwing scales magnified, Ipperwash Beach, ON (right). [Photos © by courtesy of
Jay Cossey]
32
Fig. 20. Euchloe olympia Huron habitat at Ipperwash, Ontario. [Photo © by courtesy of Ken Thorne]
CONCLUSION
Appalachian and Great Plains populations of Euchloe olympia are here separated as nominotypical
subspecies olympia and as subspecies rosa , respectively. Great Lakes regional (“inland”) populations are
intermediate, displaying a broad range of variation between Appalachian and Great Plains populations,
with populations east of Lake Michigan tending more toward nominotypical olympia , and populations
west of Lake Michigan tending more toward subspecies rosa. The eastern Missouri populations are
similarly considered intermediates but appear closer to nominotypical olympia. Great Lakes dune
populations are here treated as new subspecies wagneri (lower Lake Michigan) and Huron (lower Lake
Huron).
Wagner (1977) left us with the following comment: “As to whether the Dune Form merits
designation as a subspecies, only the future can tell. I am hesitant to affix a new name on populations that
are still so little known or understood”. The present paper fulfills this role. Torre-Bueno (1930) stated
this well: “A description has one primary purpose: to make a category known to those whom it is
unknown ; to ticket it for future reference.” Based on the present study, the following subspecific
arrangement is hereby proposed:
• Euchloe olympia olympia (W. H. Edwards, 1871)
• Euchloe olympia rosa (W. H. Edwards, 1882)
=anniha Ebner, 1970
• Euchloe olympia wagneri Pavulaan, 2019
• Euchloe olympia huron Pavulaan, 2019
33
In NatureServe (2017), under Conservation Status, it is stated: “If they were treated separately the
Appalachian populations would be at least globally uncommon if not imperiled”. Hopefully this paper, if
anything, will help provide some level of framework to help facilitate future conservation strategies for
the Appalachian and Great Lakes Dune-associated populations.
ACKNOWLEDGMENTS
I wish to sincerely thank both David M. Wright who reviewed early drafts of this manuscript and
offered several helpful suggestions, and Norbert Kondla who provided final critical review and offered
many suggestions. Irwin Leeuw was the source of specimens of the lower Lake Michigan dune
population. Jim Troubridge and Ken Thorne were the source of specimens of the lower Lake Huron dune
population. Steve Spomer provided records of rosa from several Great Plains states. John Calhoun
provided photographs of the original Edwards (1884, 1897) figures. Jeffrey Belth provided specimen,
habitat and host images from Miller Beach, IN. Jay Cossey provided living specimen images from
Lambton, ON. Ken Thorne provided the habitat image from Lambton, ON. Photographs examined in
critical detail on many varied website galleries are attributed to: Annette Allor, James Barron, Tom
Bentley, Andrew Birch, Andy Birkey, Bill Bouton, Jim Brighton, Brett Budach, Aaron Carlson, Steve
Collins, Will Cook, Jay Cossey, Steven Daniel, Patrick Deacon, Melinda Fawver, Tim Fenske, Karen
Finch, Jeff Fischer, Carolyn Gritzmaker, Aaron Gunnar, Jerry Hatfield, Frode Jacobsen, David Kinney,
Blake Mann, Robin McLeod, Steven Mlodinow, Dwayne Murphy, Colin Murray, Larry Neily, Tommie
Rogers, Steve Sass, Kurt Schaefer, Greg Sievert, Doug Taron, Ralph Thorpe, Rob Vanepps, Shirley
Wilkerson; and too many to mention via Flickr.com, Butterfliesandmoths.org and iNaturalist.org who
listed themselves with abbreviated names, anonymous monikers or pseudonyms. Permission to reproduce
the figures from Brown (1973) were granted by Dr. JoVonn Hill (American Entomological Society).
Permission to reproduce the figures from Ebner (1970) was granted by Sara Podejko (Registrar,
Milwaukee Public Museum). Permission to publish the images of the anniha neotype was granted by
Floyd Shockley (Collections Manager, Dept, of Entomology, Smithsonian National Museum of Natural
History), and helpful comments.
LITERATURE CITED
Acorn, J. 1993. Butterflies of Alberta. Lone Pine Publishing. Edmonton, Alberta: 143 pp.
Acorn, J. & I. Sheldon. 2016. Butterflies of Ontario & Eastern Canada. Partners Publishing & Lone Pine
Media Productions, Edmonton, Alberta: 320 pp.
Allen, T. J. 1997. The Butterflies of West Virginia and Their Caterpillars. University of Pittsburgh
Press, Pittsburgh, Pennsylvania: xi + 388 pp.
Arnhold, F. R. 1952. Notes on collecting Anthocaris midea and Euchloe olympia. Lepidoptera News, 6:
99-100.
Balogh, G. 1980. Wisconsin’s Lupine feeding butterflies and their pine barrens habitat. Newsletter of
the Wisconsin Entomological Society 8(2): 4-8.
Barnes, W. & J. McDunnough. 1917. Check List of the Lepidoptera of Boreal America. Herald Press,
Decatur, Illinois: viv + 392 pp.
Belth, J. E. 2013. Butterflies of Indiana: A Field Guide. Indiana University Press, Bloomington,
Indiana: 329 pp.
Bess, J. 1983. 1982 Season Summary report of Euchloe olympia on Arabis drummondii and Arabis
lyrata in Michigan. News of the Lepidopterists’ Society, 25(2): 28.
Betros, B. 2008. A Photographic Field Guide to the Butterflies in the Kansas City Region. Kansas City
Star Books, Kansas City, Missouri: vi + 407 pp.
Beutenmuller, W. (ed.). 1897. Proceedings of the New York Entomological Society. Journal of the New
York Entomological Society, 5(4): 205-208.
Beutenmuller, W. 1898. Revision of the species of Euchloe inhabiting America, north of Mexico.
Bulletin of the American Museum of Natural History, 10: 235-248.
34
Bird, C. D., G. J. Hilchie, N. G. Kondla, E. M. Pike & F. A. H. Sperling. 1995. Alberta Butterflies. The
Provincial Museum of Alberta, Edmonton, Alberta: viii + 349 pp.
Blatchley, W. S. 1891. A catalogue of the butterflies known to occur in Indiana. Annual Report of the
Indiana State Geologist, 17: 365-408.
Bouseman, J. K. & J. G. Sternburg. 2001. Field Guide to the Butterflies of Illinois. Illinois Natural
History Survey, Champaign, Illinois. Manual 9: xii + 264 pp.
Bridges, C. A. 1988. Catalogue of Papilionidae & Pieridae (Lepidoptera: Rhopalocera). Published by
author. Urbana, Illinois: vii + 727 pp.
Brock, J. P. & K. Kaufman. 2003. Butterflies of North America. Kaufman Focus Guides. Houghton
Mifflin Company, New York, New York: 384 pp.
Brown, F. M. 1973. The types of the Pierid butterflies named by William Henry Edwards. Transactions
of the American Entomological Society, 99(1): 29-118.
Brown, F. M., D. Eff & B. Rotger. 1957. Colorado Butterflies. Denver Museum of Natural History,
Denver, Colorado: viii + 368 pp.
Butler, A. G. 1899. Notes on the American forms of Euchloe (Hlibn.). Canadian Entomologist, 31: 19.
Chu, J. R. 2009. Inventories of Butterflies in Boulder County. Published by author. Boulder, Colorado:
39 pp.
Chu, J. R. & S. R. Jones. 2011. Butterflies of the Colorado Front Range. Boulder County Nature
Association, Boulder, Colorado: 112 pp.
Clark, A. H. 1940. Butterflies of Virginia. Smithsonian Explorations, 1939: 63-66.
Clark, A. H. & L. F. Clark. 1939. Butterflies from Virginia. Proceedings of the Biological Society of
Washington, 52: 177-184.
Clark, A. H. & L. F. Clark. 1951. The Butterflies of Virginia. Smithsonian Miscellaneous Collections,
116(7): vii + 239 pp.
Clarke, J. F. G. 1962. Butterflies. Golden Press. New York, New York: 68 pp.
Clench, H. K. & P. A. Opler. 1983. Studies on Nearctic Euchloe. Part 8. Distribution, ecology, and
variation of Euchloe olympia (Pieridae) populations. Annals of Carnegie Museum, 52(2): 41-54.
Comstock, J. H. & A. B. Comstock. 1904. A Manual of the Butterflies of the Eastern United States. D.
Appleton and Company, New York, N.Y.: xii + 311 pp.
Coveil, C. V., Jr. 1999. The Butterflies and Moths (Lepidoptera) of Kentucky: an annotated checklist.
Kentucky State Nature Preserves Commission. Scientific and Technical Series No. 6: xiv +220
pp.
Cross, F. C. 1937. Butterflies of Colorado. Proceedings of the Colorado Museum of Natural History,
16(1): 1-28.
D’Abrera, B. 1990. Butterflies of the Holarctic Region. Part I: Papilionidae, Pieridae, Danaidae &
Satyridae (Partim). Hill House, Victoria, Australia: 185 pp.
Daniel, S. 2018. Olympia Marble ( Euchloe olympia ) larva on Boechera grahamii. iNaturalist, accessed
December 1, 2018.
Daniels, J. C. 2004. Butterflies of Ohio. Adventure Publications, Inc., Cambridge, Minnesota: 344 pp.
Daniels, J. C. 2005. Butterflies of Michigan. Adventure Publications, Inc., Cambridge, Minnesota: 376
pp.
Denton, S. F. 1900. Moths and Butterflies of the United States, East of the Rocky Mountains. Part II.
The Butterflies. Bradlee Whidden, Boston, Massachusetts: xv + 164-361.
Dole, J. M., W. B. Gerard & J. M. Nelson. 2004. Butterflies of Oklahoma, Kansas & North Texas.
University of Oklahoma Press, Norman, Oklahoma: xiv + 282 pp.
dos Passos, C. F. 1964. A Synonymic List of the Nearctic Rhopalocera. The Lepidopterists’ Society.
Memoir No. 1. v + 145 pp.
Douglas, M. M. & J. M. Douglas. 2005. Butterflies of the Great Lakes Region. The University of
Michigan Press, Ann Arbor, Michigan: 345 pp.
35
Dyar, H. G. 1902. A List of North American Lepidoptera and Key to the Literature of this Order of
Insects. Bulletin of the United States National Museum No. 52: xix + 723 pp.
Ebner, J. A. 1970. Butterflies of Wisconsin. Milwaukee Public Museum. Popular Science Handbook
No. 12. 205 pp.
Edwards, W. H. 1871. Descriptions of new species of North American Butterflies. Transactions of the
American Entomological Society, 3(3/4): 266-277.
Edwards, W. H. 1882. Description of new species of Butterflies found in the United States. Papilio,
2(3): 45-49.
Edwards, W. H. 1884. The Butterflies of North America. Second Series. Houghton, Mifflin and
Company, Boston, Massachusetts: 463 pp.
Edwards, W. H. 1897. The Butterflies of North America. Third Series. Houghton, Mifflin and
Company, Boston, Massachusetts: 623 pp.
Ehrlich, P. R. & A. H. Ehrlich. 1961. How to Know the Butterflies. Wm. C. Brown Co. Publishers,
Dubuque, Iowa: 262 pp.
Elrod, M. J. 1906. The Butterflies of Montana - With Keys for Determination of Species. University of
Montana, Missoula, Montana. Bulletin of the University of Montana Biological Series (Bulletin
No. 30), 2(10): xvi + 175 pp.
Ely, C. A., M. D. Schwilling & M. E. Rolfs. 1986. An Annotated List of the Butterflies of Kansas. Fort
Hayes Studies; Third Series, 7: 224 pp.
Ferge, L. 2011. Checklist of Wisconsin Butterflies [revised]. Wisconsin Entomological Society Special
Publication No. 5: 10 pp.
Ferris, C. D. 1971. An Annotated Checklist of the Rhopalocera [Butterflies] of Wyoming. Science
Monograph 23. University of Wyoming, Laramie, Wyoming: 75 pp.
Ferris, C. D. & F. M. Brown. 1980. Butterflies of the Rocky Mountain States. University of Oklahoma
Press, Norman, Oklahoma: xviii + 442 pp.
Field, W. D. 1938. Studies in Kansas Insects: A Manual of the Butterflies and Skippers of Kansas.
Bulletin of the University of Kansas, Biological Series, Vol. 39(10): 328 pp.
Fisher, M. S. 2012. The Butterflies of Colorado- Part 5- Pieridae and Papilionidae. Lepidoptera of
North America series. C.P. Gillette Museum of Arthropod Diversity, Fort Collins, Colorado: iv +
193 pp.
Forbes, W. T. M. 1960. Lepidoptera of New York and Neighboring States, Agaristidae through
Nymphalidae including Butterflies, Part IV. Memoir 371. Cornell University Agricultural
Experiment Station, New York State College of Agriculture, Ithaca, New York: 188 pp.
Freeman, H. A. 1959. Butterfly collecting in Texas and New Mexico. Journal of the Lepidopterists’
Society, 13(2): 89-93.
French, G. H. 1886. The Butterflies of the Eastern United States. For the Use of Classes in Zoology and
Private Students. J. B. Lippincott Company. Philadelphia, Pennsylvania: 402 pp.
French, G. H. 1914. The Butterflies of the Eastern United States. For the use of Classes in Zoology and
Private Students. 4 th edition. J. B. Lippincott Company. Philadelphia, Pennsylvania: 429 pp.
Glassberg, J. 1999. Butterflies Through Binoculars, the East. A Field Guide to the Butterflies of Eastern
North America. Oxford University Press, Oxford, New York: x + 242 pp.
Glassberg, J. 2001. Butterflies Through Binoculars, the West. A Field Guide to the Butterflies of
Western North America. Oxford University Press, Oxford, New York: x + 374 pp.
Glassberg, J. 2012. A Swift Guide to Butterflies of North America. Sunstreak Books, Inc., U.S.A.: 416
pp.
Hall, P. W., C. D. Jones, A. Guidotti & B. Hubley. 2014. Butterflies of Ontario. Royal Ontario Museum.
Toronto, Ontario: 488 pp.
Handfield, L. 2011. Les Papillons du Quebec. Broquet, Inc., Saint-Constant, Quebec: 672 pp. + 166
plates.
Hardesty, R. L. 2015. The Butterflies of the Laramie Mountains. Papillon Press. Hungry Horse,
Montana: 49 pp.
36
Harp, C. 1986. 1985 Season Summary report of Euchloe olympia on Thelypodium wrightii in Oklahoma.
News of the Lepidopterists’ Society, 28(2): 29.
Heitzman, R. J. & J. E. Heitzman. 1987. Butterflies and Moths of Missouri. Missouri Department of
Conservation, Jefferson City, Missouri: viii + 385 pp.
Hodges, R. W., et al. (eds.). 1983. Check List of the Lepidoptera of America North of Mexico. E. W.
Classey, London. 284 pp.
Holland, W. J. 1899, 1900, 1902, 1904. The Butterfly Book - A Popular Guide to a Knowledge of the
Butterflies of North America. Doubleday, Page & Company, Garden City, New York: xx +
382 pp.
Holland, W. J. 1916a The Butterfly Book - A Popular Guide to a Knowledge of the Butterflies of North
America. The New Nature Library. Doubleday & Company, Inc., Garden City, New York: xx +
382 pp. + 77 plates.
Holland, W. J. 1916b. The Butterfly Guide - A Popular Manual for the Ready Identification of the
Commoner Species Found in the United States and Canada. Doubleday, Page & Company,
Garden City, New York: 237 pp.
Holland, W. J. 1931, 1951. The Butterfly Book - A Popular and Scientific Manual, Describing and
Depicting all the Butterflies of the United States and Canada. Doubleday & Company, Inc.,
Garden City, New York: xii + 424 pp. + 77 plates.
Holmes, A. M., Q. F. Hess, R. R. Tasker & A. J. Hanks. 1991. The Ontario Butterfly Atlas. Toronto
Entomologists’ Association, Toronto, Ontario: viii + 167 pp.
Hooper, R. R. 1969. A preliminary list of the Butterflies of Saskatchewan. Mid-Continent Lepidoptera
Series 5: 9 pp.
Hooper, R. R. 1973. Butterflies of Saskatchewan - A Field Guide. Saskatchewan Department of Natural
Resources, Regina, Saskatchewan: 216 pp.
Howe, W. H. 1975. The Butterflies of North America. Doubleday & Company, Inc., Garden City, New
York: xii + 633 pp.
Iftner, D. C., J. A. Shuey & J. V. Calhoun. 1992. Butterflies and Skippers of Ohio. Bulletin of the Ohio
Biological Survey, New Series, 9(1): xii + 212 pp.
Irwin, R. R. & J. C. Downey. 1973. Annotated checklist of the Butterflies of Illinois. Illinois Natural
History Survey. Biological Notes, No. 81. 60 pp.
Jeffords, M. R., S. L. Post & J. R. Wiker. 2014. Butterflies of Illinois - a Field Guide. Illinois Natural
History Survey. Manual 14. Champaign, Illinois: xvii + 440 pp.
Johnson, K. 1972. The Butterflies of Nebraska. Journal of Research on the Lepidoptera, 11(1): 1-64.
Johnson, K. & J. M. Malick. 1972. An annotated list of central Wisconsin butterflies. Reports on the
Fauna and Flora of Wisconsin No. 7. The Museum of Natural History, University of Wisconsin,
Stevens Point, Wisconsin: 6 pp.
Kirby, W. F. 1877. A Synonymic Catalogue of Diurnal Lepidoptera. Supplement, March, 1871-June,
1877. London, United Kingdom: 192 pp.
Klassen, P., A. R. Westwood, W. B. Preston & W. B. McKillop. 1989. The Butterflies of Manitoba.
Manitoba Museum of Man and Nature, Winnipeg, Manitoba: vi + 290 pp.
Klots, A. B. 1951. A Field Guide to the Butterflies of Eastern North America. Peterson Field Guide
Series. Houghton Mifflin Company, Boston, Massachusetts: xvi + 349 pp.
Kohler, S. 1980. Checklist of Montana Butterflies (Rhopalocera). Journal of the Lepidopterists’ Society
34(1): 1-19.
Kondla, N. G. 2004. Conservation overview of butterflies in the Southern Headwaters at Risk Project
(SHARP) area. Alberta Sustainable Resource Development, Fish and Wildlife Division, Alberta
Species at Risk Report No. 8. Edmonton, Alberta: 35 pp.
Layberry, R. A. 1985. 1984 Season Summary report of Euchloe olympia on Arabis divaricarpa in
Quebec. News of the Lepidopterists’ Society, 27(2): 26.
Layberry, R. A. 1986. 1985 Season Summary report of Euchloe olympia on Arabis divaricarpa in
Ontario. News of the Lepidopterists’ Society, 28(2): 30.
37
Layberry, R. A., J. D. Lafontaine & P. W. Hall. 1982. Butterflies of the Ottawa District. Trail &
Landscape, 16(1): 3-59.
Layberry, R. A., P. W. Hall & J. D. Lafontaine. 1998. The Butterflies of Canada. University of Toronto
Press, Toronto, Ontario: vii + 280 pp.
Leboeuf, M. & S. Le Tirant. 2012. Papillons et Chenilles du Quebec et des Maritimes. Editions Michel
Quintin, Waterloo, Quebec: 391 pp.
LeGrand, Jr., H. E. & T. E. Howard, Jr. 2019. Butterflies of North Carolina. Twenty-sixth
Approximation. North Carolina Division of Parks and Recreation. Available online:
http ://www.dpr.ncparks. gov/nbnc/
Leussler, R. A. 1938. An annotated list of the butterflies of Nebraska, with the description of a new
species. Entomological News, 49: 3-9, 76-80, 213-218, 275-280.
Lindsay, S. A. 1971. A host plant for northern populations of Euchloe olympia (Pieridae). Journal of the
Lepidopterists’ Society, 25(1): 64.
Macy, R. W. & H. H. Shepard. 1941. Butterflies: A Handbook of the Butterflies of the United States,
Complete for the Region North of the Potomac and Ohio Rivers and East of the Dakotas. The
University of Minnesota Press, Minneapolis, Minnesota: vii + 247 pp.
Marrone, G. M. 2002. Field Guide to Butterflies of South Dakota. South Dakota Department of Game,
Fish and Parks, Pierre, South Dakota: 478 pp.
Masters, J. H. 1972. Critical Comments on James A. Ebner’s “Butterflies of Wisconsin”. Mid-Continent
Lepidoptera Series, 4(59): 5-11.
Masters, J. H. 1973. Butterfly records for three northwest Wisconsin counties. Journal of Research on
the Lepidoptera 1972(1973) 11(3): 175-182.
McDunnough, J. 1938. Check List of the Lepidoptera of Canada and the United States of America - Part
1, Macrolepidoptera. Memoirs of the Southern California Academy of Sciences, Vol. 1: 275 pp.
Meiners, E. P. 1939. The life history of Euchloe olympia Edwards, with some notes on its habits.
Proceedings of the Missouri Academy of Science 4(6): 154-156.
Meiners, E. P. 1956. Lepidoptera Collecting at Ranken, Missouri. Lepidopterists’News 10(5): 163-168.
Miller, L. D. & F. M. Brown. 1981. A Catalogue/Checklist of the Butterflies of America North of
Mexico. The Lepidopterists’ Society, Memoir No. 2. vii + 280 pp.
Moore, S. 1922. A list of northern Michigan lepidoptera. Occasional Papers of the Museum of Zoology
114:5-28.
Murray, C. 2018. Euchloe olympia entry posted 5/14/2018 in iNaturalist.
Neck, R. W. 1996. A Field Guide to Butterflies of Texas. Gulf Publishing Company, Houston, Texas:
xvii + 323 pp.
Nielsen, M. C. 1966. 1965 Season Summary report of Euchloe olympia on Arabis drummondii in
Michigan. News of the Lepidopterists’ Society, 8(3): 11.
Nielsen, M. C. 1967. 1966 Season Summary report of Euchloe olympia on Arabis drummondii in
Michigan. News of the Lepidopterists’ Society, 9(3): 11.
Nielsen, M. C. 1979. 1978 Season Summary report of Euchloe olympia on Arabis drummondii in
Michigan. News of the Lepidopterists’ Society, 21(2): 11.
Nielsen, M. C. 1999. Michigan Butterflies & Skippers: A Field Guide and Reference. MSU Extension,
Michigan State University, East Lansing, Michigan: 248 pp.
Opler, P. A. 1966. Studies on Nearctic Euchloe. Part 3. Complete Synonymical Treatment. Part 4.
Type Data and Type Locality Restrictions. Journal of Research on the Lepidoptera, 5(3): 185-
195.
Opler, P. A. 1974. Studies on Nearctic Euchloe. Part 7. Comparative Life Histories, Hosts and the
Morphology of Immature Stages. Journal of Research on the Lepidoptera, 13(1): 1-20.
Opler, P. A. 1985. Invertebrates - Chapter 2 in: Genoways, H. H. & F. J. Brenner (eds.), Species of
Special Concern in Pennsylvania. Special publication of Carnegie Museum of Natural History,
Pittsburg, Pennsylvania 11: 81-165.
38
Opler, P. A. & G. O. Krizek. 1984. Butterflies East of the Great Plains. The Johns Hopkins University
Press, Baltimore, Maryland: xvii + 294 pp.
Opler, P. A. & V. Malikul. 1992. A Field Guide to Eastern Butterflies. Peterson Field Guide Series.
Houghton Mifflin Company, New York, New York: xvii + 396 pp.
Opler, P. A. & A. D. Warren. 2003. Butterflies of North America. 2. Scientific Names List for Butterfly
Species of North America, north of Mexico. C. P. Gillette Museum of Arthropod Diversity,
Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort
Collins, Colorado. 79 pp.
Opler, P. A. & A. B. Wright. 1999. A Field Guide to Western Butterflies. Peterson Field Guide Series.
Houghton Mifflin Company, New York, New York: xiv + 540 pp.
Orwig, T. T. 1990. Loess Hills Prairies as Butterfly Survivia: Opportunities and Challenges.
Proceedings of the Twelfth North American Prairie Conference 1990: 131-135.
Parshall, D. K. 2002. Conservation Assessment for Olympia Marble Butterfly ( Euchloe olympia).
USDA Forest Service, Eastern Region: 16 pp.
Patterson, J. 2011. The Butterflies of Minnesota: A ‘fliers Manual. Trafford Publishing, U.S.A.: x +
410 pp.
Pelham, J. P. 2008. A Catalogue of the Butterflies of the United States and Canada - with a complete
bibliography of the descriptive and systematic literature. Journal of Research on the Lepidoptera,
40: xiii + 652 pp.
Pohl, G. R., G. G. Anweiler, B. C. Schmidt & N. G. Kondla. 2010. An annotated list of the Lepidoptera
of Alberta, Canada. ZooKeys 38: 1-549.
Pohl, G. R., J. Landry, C. Schmidt, J. D. Lafontaine, J. T. Troubridge, A. D. Macaulay, E. J. Van
Nieukerken, J. R. Dewaard, J. J. Dombroskie, J. Klymko, V. Nazari & K. Stead. 2018. Annotated
Checklist of the Moths and Butterflies (Lepidoptera) of Canada and Alaska. Pensoft Publishers.
Sofia, Bulgaria. 580 pp.
Pyle, R. M. 1981. National Audubon Society Field Guide to North American Butterflies. Chanticleer
Press, Inc., New York: 924 pp.
Rouse, E. P. 1969. The Pieridae of Arkansas. Arkansas Academy of Science Proceedings 23: 94-104.
Royer, R. A. 1988. Butterflies of North Dakota: an Atlas and Guide. Minot State University, Science
Monograph No. 1: x + 192 pp.
Royer, R. A. 2003. Butterflies of North Dakota. Minot State University, Science Monograph No. 2: xi +
192 pp.
Schlicht, D. W., J. C. Downey & J. C. Nekola. 2007. The Butterflies of Iowa. University of Iowa Press,
Iowa City, Iowa: xii + 233 pp.
Scott, J. A. 1986a. Larval hostplant records for butterflies and skippers (mainly from western U. S.),
with notes on their natural history. Papilio New Series, 4: 1-37.
Scott, J. A. 1986b. The Butterflies of North America, a Natural History and Field Guide. Stanford
University Press, Stanford, California: xiii + 583 pp.
Scott, J. A. 1992. Hostplant records for butterflies and skippers (mostly from Colorado) 1959-1992,
with new life histories and notes on oviposition, immatures, and ecology. Papilio New Series, 6:
1-185.
Scott, J. A. 2006. Butterfly hostplant records, 1992-2005, with a treatise on the evolution of Erynnis ,
and a note on new terminology for mate-locating behavior. Papilio New Series, 14: 1-74.
Scott, J. A. 2008. Biological Catalogue of North American Butterflies. Papilio New Series, 20: 1-51.
Scott, J. A. 2020. Butterflies of the southern Rocky Mountains area, and their natural history and
behavior. Papilio New Series, 27: 1-390.
Scudder, S. H. 1889. The Butterflies of the Eastern United States with Special Reference to New
England, Volume III. Published by the author, Cambridge, Massachusetts: vi + 1775-1958 + 89
plates + 3 maps.
Sedman, Y. 1983. 1982 Season Summary report of Euchloe olympia habitat in Illinois. News of the
Lepidopterists’ Society, 25(2): 26.
39
Sedman, Y. & D. Hess. 1985. The Butterflies of West Central Illinois. Western Illinois University.
Series in the Biological Sciences, No. 11: 120 pp.
Seitz, A. 1924. The Macrolepidoptera of the World. Vol. 5. The American Rhopalocera. Text. Alfred
Kernen Verlag, Stuttgart, Germany: viii + 1139 pp.
Seitz, A. 1924. The Macrolepidoptera of the World. Vol. 5. The American Rhopalocera. Plates. Alfred
Kernen Verlag, Stuttgart, Germany: vi + 194 plates.
Shull, C. A. 1907. Life history and habits of Anthocharis (Synchloe) olympia Edw. Entomological
News, 18:73-82.
Shull, E. M. 1987. The Butterflies of Indiana. Indiana Academy of Sciences, Indianapolis, Indiana: viii
+ 262 pp.
Skinner, H. 1896. Impressions received from a study of our North American Rhopalocera. Journal of
the New York Entomological Society, 4(3): 107-118.
Skinner, H. 1898. Synonymic Catalogue of the North American Rhopalocera. American Entomological
Society special publication. Philadelphia, Pennsylvania: xv + 100 pp. + index i-xiv.
Spencer, L. A. 2014. Arkansas Butterflies and Moths. Ozark Society Foundation, Little Rock, Arkansas:
xiv +314 pp.
Strecker, H. 1872. Lepidoptera, Rhopaloceres and Heteroceres, Indigenous and Exotic. B. F. Owen.
Reading, Pennsylvania: 143 pp.
Strecker, H. 1878. Butterflies and Moths of North America. B. F. Owen, Reading, Pennsylvania: ii +
281 pp.
Swengel, A. & S. R. Swengel. 1997. Co-occurrence of prairie and barrens butterflies: Applications to
ecosystem conservation. Journal of Insect Conservation 1(2): 131-144.
Tilden, J. W. 1975. An analysis of the W. G. Wright butterfly and skipper plesiotypes in the collection of
the California Academy of Sciences. Occasional Papers of the California Academy of Sciences,
118:44 pp.
Tilden, J. W. & A. C. Smith. 1986. A Field Guide to Western Butterflies. Peterson Field Guide Series.
Houghton Mifflin Company, Boston, Massachusetts: xii + 370 pp.
Torre-Bueno, J. R. 1930. The function of a description. Bull. Brooklyn Ent. Soc. 25(1): 54-55.
Venable, R. 2014. Butterflies of Tennessee. Maywood Publishing, Franklin, Tennessee: 391pp.
Wagner, W. H. 1977. A distinctive dune form of the Marbled White butterfly, Euchloe olympia
(Lepidoptera: Pieridae) in the Great Lakes area. Great Lakes Entomologist, 10(3): 107-112.
Wagner, W. H. & M. K. Hansen. 1980. Size reduction southward in Michigan’s Mustard White
butterfly, Pieris napi (Lepidoptera: Pieridae). Great Lakes Entomologist 13(2): 77- 80.
Wauer, R. H. 2006. Finding Butterflies in Texas: a guide to the best sites. Johnson Books, Boulder,
Colorado: viii + 327 pp.
Weed, C. M. 1917a, 1923. Butterflies Worth Knowing. Little Nature Library. Doubleday, Page &
Company, Garden City, New York: xiii + 286 pp.
Weed, C. M. 1917b. Canadian Butterflies Worth Knowing. Little Nature Library. The Musson Book
Co., Toronto, Ontario: xiii + 286 pp.
Wolfe, J., J. Harry & T. Stout. 2010. Tips on collecting and rearing immatures of 375 butterfly and
skipper taxa. The Taxonomic Report 7(3): 39 pp.
Wright, W. G. 1906. Butterflies of the West Coast of the United States. Second edition. Published by
the author, San Bernardino, California: vii + 257 pp.
40
ADDITIONAL INTERNET RESOURCES REFERENCED
Blackwater Birds. Nature blog hosted by Andrew Birch. 2010 Olympia Marble reference:
http://blackwaterbirder.blogspot.com/2010/04/butterflies-are-flying.html (last accessed June 28,
2015).
Butterflies and Moths of North America. Olympia Marble ( Euchloe olympia) search:
https://www.butterfliesandmoths.org/species/Euchloe-
olympia?region=All&sort by=field sightingdate value&sort order=DESC&=Submit (last
accessed April 5, 2020).
Butterflies of Ontario website. Olympia Marble ( Euchloe olympia) page:
http://www.ontariobutterflies.ca/families/pieridae/olympia-marble (last accessed December 1,
2018).
Canadian Biodiversity Information Facility (CBIF). Euchloe olympia search:
https://www.cbif.gc.ca/acp/eng/species-access/browse?scientific=Euchloe+olympia&reset=true
(last accessed April 20, 2020).
Carolina Nature website. Olympia Marble {Euchloe olympia) page:
http://www.carolinanature.com/butterflies/olympiamarble.html (last accessed April 5, 2020).
The Dallas County Lepidopterists’ Society website. Olympia Marble {Euchloe olympia) page:
http://www.dallasbutterflies.com/Butterflies/html/olympia.html (last accessed April 5, 2020).
e-Butterfly website. Olympia Marble {Euchloe olympia) search:
http://www.e-butterfly.Org/ebutterflyapp/#/observations/lists (last accessed April 6, 2020).
Flickr website. Olympia Marble {Euchloe olympia) search:
https://www.flickr.com/search/?text=01vmpia+marble+butterfly (last accessed December 1,
2018)
Harvard University, Museum of Comparative Zoology, Zoological Collections Database. Specimen
search page: http://mczbase.mcz.harvard.edu/SpecimenSearch.cfm (last accessed April 5, 2020).
iNaturalist website. Olympia Marble {Euchloe olympia) search:
https://www.inaturalist.org/observations7place id=l&taxon id=58527 (last accessed April 5,
2020 ).
Minnesota Butterfly Walks. Nature blog hosted by “Minnesotastan” (anonymous):
http://butterflywalks.blogspot.com/ (last accessed May 5, 2015).
NatureServe. 2017. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.1.
NatureServe, Arlington, Virginia. Available http://explorer.natureserye.org (last accessed:
February 28, 2018).
Toronto Entomologists’ Association. Ontario Butterfly Atlas Online.
http://www.ontarioinsects.org/atlas online.htm (last accessed April 20, 2020).
USDA Plants Database: https://plants.usda.gov/iava/ (last accessed December 30, 2019).
Wisconsin Butterflies website. Olympia Marble {Euchloe olympia) search:
htt ps://wisconsinbutterflies.org/ (April 5, 2020).
Yale Peabody Museum of Natural History, collections database, Euchloe olympia online search:
http://collections.peabody.yale.edu/search/Search/Results?lookfor=Euchloe+olvmpia (last
accessed December 1, 2018).
Submitted for review April 7, 2020, accepted for publication April 21, 2020.
APPENDIX A (SUMMARY OF HOSTPLANTS)
HOSTPLANT
STATE/PROVINCE
SUBSPECIES
Arabidopsis thaliana
MD
olympia
Arabis alpina ( =albida)
ON
intermediate population
Arabis divaricarpa
NY
intermediate population
ON
intermediate population
QC
intermediate population
Arabis drummondii
CO
rosa
MI
intermediate 'inland' population
ON
intermediate population
Arabis fendleri
CO
rosa
Arabis glabra
QC
intermediate population
ON
intermediate population
Arabis lyrata
MN, ON, WI
intermediate population
IN, MI, WI
wagneri
ON
huron
Arabis missouriensis ( -viridis )
MO
intermediate population
Arabis serotina
WV
olympia
Boechera canadensis
TX
rosa
Boechera grahamii
NY
intermediate population
Boechera laevigata
NC, VA, WV
olympia
Boechera (sp.)
ON
intermediate population
Cardamine hirsuta
WV
olympia
Cardamine pensylvanica
MD
olympia
Chorispora tenella
SD
rosa
Descurainia incana (-richardsonii )
CO
rosa
Descurainia pinnata
CO, TX
rosa
Descurainia sophia
CO
rosa
Isatis tinctoria
unspecified (lab host)
rosa
Lepidium campestre
CO
rosa
Lepidium virginicum
CO, IL
rosa
Sisymbrium altissimum
CO
rosa
Sisymbrium officinale
KS, SK, TX
rosa
Thelypodium wrightii
OK
rosa
42
DISTRIBUTION OF EUCHLOE OLYMPIA SUBSPECIES
IN THE UNITED STATES OF AMERICA
Map 1. Key: Gray counties = published records of Euchloe olympia for which no specimens or images
were available for evaluation. Red counties = records of only ssp. olympia phenotype based on examined
specimens or images. Yellow counties = records of only ssp. rosa phenotype based on examined
specimens or images. Orange counties = records of intermediate phenotypes or presence of BOTH
olympia and rosa phenotypes based on examined specimens or images. Blue counties = ssp. wagneri.
43
DISTRIBUTION OF EUCHLOE OLYMPIA SUBSPECIES
IN CANADA
were available for evaluation. Red counties = records of only ssp. olympia phenotype based on examined
specimens or images. Yellow counties = records of only ssp. rosa phenotype based on examined
specimens or images. Orange counties = records of intermediate phenotypes or presence of BOTH
olympia and rosa phenotypes based on examined specimens or images. Blue county = ssp. Huron. [Note:
For the purpose of this atlas, the western provinces of Alberta, Saskatchewan and Manitoba are
represented by their census divisions (see Google Earth for exact boundaries), because counties are not
established as they are in the eastern provinces of Ontario and Quebec. This is partly to maintain a visual
consistency with the mapping scheme applied to the eastern provinces and to the map of the United
States.
44
The Taxonomic Report
is a publication of
The International Lepidoptera Survey (TILS)
The International Lepidoptera Survey is registered as a Limited Liability Company (LLC) in the state of
Virginia, U.S.A. The Taxonomic Report (TTR) is published for the purpose of providing a public and
permanent scientific record. It appears in digital, open-access form, is regularly disseminated in hardcopy
form to select institutional repositories and is also available as printed copy upon request at the discretion
of authors and/or the editor. Printing and postage costs may apply. Contents are peer-reviewed but not
necessarily through the anonymous review and comment process preferred by some publishers of serial
literature.
All issues of The Taxonomic Report are uploaded to, and available via the following digital repositories
and websites:
Biodiversity Heritage Library https://www.biodiversitvlibrary.org
Internet Archive https://archive.org/
Zobodat https://www.zobodat.at/publikation series.php?id=21284
TILS Purpose
TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific discovery,
determination, and documentation, without which there can be no preservation.
TILS Motto
“As a world community, we cannot protect that which we do not know”
Articles for publication are sought
They may deal with any area of research on Lepidoptera, including faunal surveys, conservation topics,
methods, etc. Taxonomic papers are especially welcome. There are no page charges for authors. Before
sending a manuscript, simply write to TTR editor, Harry Pavulaan, 606 Hunton Place NE, Leesburg,
VA, 20176 to initiate discussion on how to best handle your material for publication, and to discuss peer
review options; or email to intlepsurvev@gmail.com .
Visit The International Lepidoptera Survey on the World Wide Web at:
http://lepsurvey.carolinanature.com
&
Join the discussion at our list serves on Yahoo! Groups at:
http://groups.yahoo.com/group/TILS-leps-talk/
http://groups.yahoo.com/group/TILS-moth-rah/
45