Volume 10 Number 3 28 April 2022

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

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

A new species of Celastrina from the northwestern United States and

southwestern Canada with a lectotype designation of Lycaena

pseudargiolus var. nigrescens Fletcher (Lycaenidae: Polyommatinae)

Caitlin C. LaBar

Kelso, WA, USA

Email: northwestbutterflies@gmail.com

Jonathan P. Pelham

Burke Museum of Natural History and Culture

University of Washington

Box 353010, Seattle, WA 98195-3010, USA

Email: zapjammer@comcast.net

Norbert G. Kondla

47 Pensacola Close SE, Calgary, Alberta, T2A 2K5, Canada

Email: coliasOO7@gmail.com

ABSTRACT. A new western azure species, Celastrina asheri, is described from the northwestern United States and

southwestern Canada. A lectotype is designated for Lycaena pseudargiolus var. nigrescens Fletcher, 1903 to secure the identity

of that name. C. asheri broadly overlaps with the northeastern range of C. echo (W. H. Edwards, 1864) but is usually very

distinct from that species. C. /ucia (W. Kirby, 1837) is narrowly sympatric with C. asheri in British Columbia, the southwest

comer of Alberta and along the Rocky Mountains in Montana. As far as is known, C. asheri uses only red osier dogwood

(Cornus sericea) and possibly oceanspray (Holodiscus discolor) as larval foodplants, while C. echo and C. lucia use a wide

variety of larval foodplants. A lack of past research and confusion as a “form” of C. echo obscured the identity of C. asheri.

Additional key words: new species, sympatry, life history, larvae, Cornus sericea

ZooBank registration: urn:|sid:zoobank.org: pub: F7F2600A-6690-4277-AFE7-F8DC85E1178F

INTRODUCTION

The history of Ce/astrina has long been a complex and fascinating puzzle. Downey (1961) and

Langston (1975) treated all North American azures as Celastrina argiolus (Linnaeus, 1758). Miller &

Brown (1981) treated North American species as C. /adon (Cramer, 1780) and C. nigra (=ebenina)

(Clench, 1972). By 2008, nine species of azures were recognized and have persisted to this day (Pelham

2008 & 2022). This increasing taxonomic complexity and consequent confusion is notable in southwestern

Canada and northwestern United States. Layberry, et al. (1998) considered the taxa /ucia (W. Kirby, 1837),

echo (W. H. Edwards, 1864) and nigrescens (Fletcher, 1903) as subspecies of C. /adon in western Canada.

They stated that nigrescens may exhibit either the ‘lucia’ (dark patches on the VHW) or ‘violacea’ (no

dark patches) form. Guppy & Shepard (2001) elevated C. echo to the species level and considered the

1

taxon nigrescens aS Synonymous with the nominate subspecies, C. e. echo. They gave the range: Southern

BC to the tip of Baja California and south through the Rocky Mountains to northern Mexico. This is

accurate for at least the coastal portion of its range. Between 1997 and 2002, the third author began closely

investigating Ce/astrina in the vicinity of the nigrescens type locality in southeastern British Columbia. He

noticed both a phenotypic and flight phenology difference separate from C. echo, leading him to suspect

the ventrally darker, lucia-like phenotype was an undescribed subspecies of C. /ucia or an undescribed

species of Celastrina.

The confusion with these azures is illustrated by what Newcomer (1964) said about them: “Two

forms occur, one typical of ssp. echo and the other with the dark markings as in ssp. /ucia, and both may

be taken at the same place on the same day.” It is conceptually impossible for two subspecies to occur

sympatrically and synchronically. Time has revealed that Newcomer was right, but he did not correctly

identify the taxa concerned. It is not obvious. In 1999, David Nunnallee discovered the Cowiche Canyon

colony which came to be known as C. lucia Central WA segregate (Warren, et al. 2017). This butterfly is

accessible and regularly recorded at this locality. In 2001, Andrew Warren visited the collection at Burke

Museum of Natural History and Culture and identified these and other Washington specimens as C. lucia.

These small colonies exhibited heavy ventral markings consistent with Canadian C. /ucia, and the species

became a familiar butterfly in the Washington butterfly community, believed to only occur on the east

slope of the central Washington Cascades, far from its range across Canada and the northeastern United

States (Pyle, 2002; Warren, 2005; Pyle & LaBar, 2018). Warren (2005) discussed Ce/astrina populations

in northeastern Oregon and suggested there may be more than one species, noting that most males were

a paler blue above than typical C. echo, as well as the wing fringes being darker and usually strongly

checkered, and the ventral hindwing having highly variable maculation. Warren stated that C. /ucia should

not be applied to those variable forms in Oregon but was uncertain how they should be classified based

on data available at the time, suggesting either nigrescens, bakeri, or an as-yet undescribed species. These

less bold, lucia-like individuals found in eastern Washington and Oregon were considered part of the taxon

nigrescens but questions about it possibly being a distinct species persisted (Warren, 2005; Pyle & LaBar,

2018).

In May 2020, another clue emerged when Melanie Weiss photographed a “C. lucia” at lower

Reecer Canyon in Kittitas County, Washington. Variably marked individuals had long been known from

this frequently visited location, but specimens with heavy ventral maculation were uncommon, and the

pale blue dorsal of Melanie’s butterfly had not been noticed there before. In April 2021, the discovery of

a new colony of “C. lucia” by Dan Dunphy and Cathy Clark in central Washington provided new clues

about their habitat preferences and prompted us to investigate further. The results of this investigation are

reported here.

MATERIALS AND METHODS

There has been a concerted three-decade effort to accumulate butterfly distribution records in

Washington and the surrounding states and provinces. This has resulted in tens of thousands of individual

recordings. Most of these represent specimens that reside in either personal or institutional collections,

in particular the Burke Museum of Natural History and Culture (UWBM) and Oregon State Arthropod

Collection. The current research intends to reduce the confusion that prevailed when Celastrina records

were initially gathered and depends more directly on specimens at hand. We examined specimens from

the collections of UWBM, Caitlin C. LaBar (CCL) and Norbert G. Kondla (NGK), images of live adults

reported to the authors by members of the Washington Butterfly Association, and records from iNaturalist,

Butterflies and Moths of North America and Symbiota Collections of Arthropods Network. Larvae of

the new taxon were collected and reared from two locations in Washington: Ringer South Trail, Kittitas

County, and Sinlahekin Wildlife Area, Okanogan County. These were compared with life histories of

Celastrina reared by the second author and those reported in James & Nunnallee (2011).

RESULTS AND DISCUSSION

Assessment of phenotypes, phenology and habitat clearly show three Celastrina species in

northwestern United States and southwestern Canada: C. echo, C. lucia and an undescribed species.

In this region, C. /ucia occupies boreal forest habitats north of approximately latitude N50.7 in British

Columbia, extending further south in Alberta and rarely west of the Rockies in Montana. C. echo overlaps

with C. /ucia around the periphery of its range and is widespread and common from southern British

Columbia to southern California. The undescribed species occurs at lower elevations throughout much

of the northeastern range of C. echo. All three species co-occur in southwest Alberta where they are

distinguishable in appearance and exhibit slightly different flight periods. With the undescribed species

being more closely associated with C. echo, determining what name to apply to it involved the identity

of the taxa nigrescens and bakeri. Both have been intermittently attributed to a mix of C. echo and the

undescribed species.

The holotype of Lycaenopsis pseudargiolus bakeri Clench, 1944 (synonym of C. echo nigrescens)

has a dark blue dorsal (Fig. 9a), a pale, white-gray ventral with greenish blue coloration of the thorax and

the basal to postbasal region of the wings, faint spots and caps on the wing margins and small, rounded

discal maculation (Fig. 9b). Clench (1944) described the hindwing fringe of bakeri as white and only

occasionally with a faint, charcoal-colored line at the vein ends, which is seen on the holotype. All of

these features are typical of C. echo. While researching the taxon nigrescens, we discovered that it has no

defined holotype and the type series is primarily C. echo with a few specimens of the undescribed species.

This requires the designation of a lectotype, establishing the identity of nigrescens as a subspecies of C.

echo and enabling the definition of the undescribed species.

Designation of a lectotype for Lycaena pseudargiolus var. nigrescens Fletcher, 1903

(Figs. 1-8)

In the original description of C. echo nigrescens, two specimens, a male and female, are

illustrated and 16 total (8 males and 8 females) were stated as the types (Fletcher, 1903).

Per I.C.Z.N. Article 74.7, in the absence of a single, designated holotype, all specimens are

thereby treated as syntypes. Eight of these specimens (4 males and 4 females) were located by

David M. Wright in 2006 at the National Museum of Natural History, Smithsonian Institution

(Figs. 1-8). Of the four male specimens, one has a slightly dusky VHW margin (Fig. 3b)

and two have heavier maculation on the VHW disc (Figs. Ib and 8b), but otherwise display

characteristics typical of C. echo. All four females exhibit light ventral maculation, a dark

blue DFW discal area bordered with a dark charcoal DFW margin extending almost to the

median, a prominent black bar in the DFW cell bar, very little blue on the DHW, and mostly

white fringes; all features typical of female C. echo, particularly on the eastern side of the

Cascades. Fletcher (1903) described C. echo nigrescens as being distinct from other members

of the species because of the “large amount of black on the upper surface of the females.” He

described the males as having deep violet-blue dorsals and variably marked ventrals, with the

VHW discal dark blotches being “most typical of the variety” according to J. W. Cockle, who

provided the type specimens to Fletcher. There is no description of their habitat or details of

exactly where each specimen was collected other than the vicinity of Kaslo, Kootenay Lake,

British Columbia. Considering all this, the description of nigrescens appears to be primarily

dependent on the females while the males are a mix of C. echo and an undescribed species. We

designate the female specimen numbered ‘4’ in the photo plate as the lectotype for Celastrina

echo nigrescens. Lectotype and paralectotype labels will be mailed to the curator of the National

Museum of Natural History, Smithsonian Institution to be placed on the specimens of the type

series.

Lectotype label is red, printed: / LECTOTYPE / Lycaena pseudargiolus / var. nigrescens

3

Fletcher, 1903 / designated by / LaBar, Pelham & Kondla 2022 /. Paralectotype labels are

yellow, printed: / PARALECTOTYPE / Lycaena pseudargiolus / var. nigrescens Fletcher,

1903 / designated by / LaBar, Pelham & Kondla 2022 /.

1) Kaslo, B.C., 2) Kaslo, B.C., 3) Kaslo, B.C., 4) Kaslo,

B.C., 5) Kaslo, B.C., 6) Kaslo, B.C., 7) Kaslo, B.C., 8) Kaslo,

no date, 9-V-1902, Apr 24-30, 9-V-1902, Apr 24-30, Apr 24-30, Apr 24-30, B.C., Apr 24,

J. W. Cockle, J. W. Cockle, J. W. Cockle, J. W. Cockle, J. W. Cockle, J. W. Cockle/ J. W. Cockle/ J. W. Cockle/

NMNH NMNH NMNH NMNH NMNH Barnes, Barnes, Barnes,

NMNH NMNH NMNH

Figs. 1-8: Dorsal (a) and ventral (b) images of four male (1, 3, 6, 8) and four female (2, 4, 5, 7) type specimens of Lycaena

pseudargiolus var. nigrescens Fletcher, 1903 located at the National Museum of Natural History, Smithsonian Institution

(NMNH) and photographed by David M. Wright in 2006.

é Lycacno sss. i

opseudarq posing bakert

8 = ety df a

ss é 7 ‘ a Baker, Ore.

5; , “ tw Eostys

<f ake ie : TG. Roker 9¢

Fig. 9: Dorsal (a), ventral (b) and specimen label (c) of Lycaenopsis pseudargiolus bakeri Clench, 1944 holotype at the National

Museum of Natural History, Smithsonian Institution, photographed by David M. Wright in 2006.

4

Celastrina asheri LaBar, Pelham & Kondla, new species

(Asher’s Blue)

(Figs. 10-41)

ZooBank registration: urn:|sid:zoobank.org:act:}6 E7OFCCC-S505A-4A A4-80C2-4D52C63A 8274

Description. MALE (Figs. 10-31, 41): mean forewing length =13.4 mm (10.7-15.1 mm, n=37,

holotype =13.3 mm). Dorsal is blue, varying in tint from ice blue to lavender, edged with thin

black border on all wings. Some individuals may exhibit white scaling in a broad postmedian

band across the DHW (Figs. 12a, 13a, 15a, 17a, 24a, 25a). DFW fringe is always dark gray to

black from apex to M2 or M3; remainder of DFW, all of DHW and ventral wing fringes range

from slightly checkered to mostly black. Ventral ground color ranges from pale gray to dusky

medium gray, often with a brownish tint that becomes stronger as specimens age (Fig. 41).

Ventral maculation is colored medium to dark gray and is highly variable in size and shape.

This dark gray maculation fades to brown in older specimens (Fig. 41). Spots may be round or

irregular. The VHW frequently exhibits a large charcoal patch which ranges in size from only

filling the discal cell (Fig. 11b) to covering the entire disc from the postmedian to postbasal

region (Fig. 21b). In live, fresh specimens, the ventral maculation often has a pearlescent sheen

in sunlight (Fig. 41a). VHW marginal pattern ranges from a series of small- to medium-sized

dark gray spots which may or may not be surrounded by pale to dark gray scales, to a solid,

dark gray marginal band. The VHW marginal spots or band are usually capped with prominent

dark gray crescents. VF W margin usually a slightly lighter version of the VHW margin. In

individuals with heavy maculation, these crescents often form thick, dark caps above the

marginal band and in rare cases merge into the solid discal patch in cells CuA1 and CuA2 (Fig.

21b). Ventral wings are frequently frosted with white outside of the dark maculation, especially

in the VHW postmedian band, where it often resembles fuzzy white arrowheads (Figs. 12b,

14b, 15b, 22b). No dorsal discal cell bars. Thin to somewhat blocky discal cell bar on VF W

equal to or slightly lighter than other maculation. When not incorporated into a discal patch,

the VHW discal cell bar is of similar width and shade as surrounding discal spots, or only

slightly lighter (Figs. 28b, 30b, 31b). Veins on ventral side frequently stand out as they tend to

be covered in scales slightly darker than the background color and lighter than the maculation.

Ventral thorax and basal area of ventral wings is gray-black or dark gray-blue, almost never

with greenish tint, and rarely reaching submedian area. Second brood phenotype tends to have

a brighter dorsal color, often with white in the DHW postmedian band, a paler ventral ground

color, and reduced VHW discal maculation (Figs. 24, 25, 31). Genitalia of North American

Celastrina have not proven useful taxonomically and are not considered herein.

FEMALE (Figs. 32-40): mean forewing length =12.7 mm (11.6-13.6 mm, n=7). Dorsal ranges

from ice blue to lavender, occasionally with a dark charcoal patch in CuA1 and CuA2 cells on

DFW and DHW (Fig. 33a). DFW dark gray or black margin is often narrow, but may extend

into the postmedian region, especially in northern populations (Figs. 36a, 39a, 40a, 97a). The

dark gray border widens at the DFW apex and may extend down the costal margin. Blue on

the DHW often extends to the border and may be mixed with dark gray (Fig. 38a) or white

scales (Fig. 35a). When the DHW exhibits a dark gray margin, it is usually wider along the

costal margin and flecked with blue scales around gray spots in the outer margin, giving it the

appearance of indistinct gray crescents riming the marginal band. DHW marginal spots usually

fuzzy or absent. Wing fringe as in males. Thin, dark gray cell bars sometimes occur on DFW

and are almost always absent from DHW. Ventral ground color, cell bars and other maculation

as in males. Females exhibiting dark gray patches in CuAl and CuA2 cells on the DFW and

DHW also tend to have heavier ventral maculation (Fig. 33). Similar to males, females of the

second brood phenotype tend to have more white on the DHW, a paler ventral ground color,

5)

and reduced VHW discal maculation (Fig. 35). The “summer form” female depicted in the

Celastrina echo account by Guppy & Shepard (2001) on page 227 is a second brood C. asheri.

It was collected by J. and S. Shepard on 1981-08-04 at the north end of Osoyoos Lake, BC.

lcm

Figs. 10-25: Celastrina asheri sp. n. male type specimens. 10: holotype male dorsal (a) and ventral (b). 11-25: selection of male

paratypes. Specimens collected from Ringer South Trail, Kittitas County, WA on 2021-04-21 (Figs. 10-23, first brood) and

2021-06-21 (Figs. 24-25, second brood). Photos by C. C. LaBar.

6

lcm

Figs. 26-31: Celastrina asheri sp. n. males (not paratypes). 26: from lower Reecer Canyon, Kittitas County, WA on 2005-04-26.

27-31: from upper Sinlahekin Creek, Sinlahekin Wildlife Area, Okanogan County, WA on 2013-05-11 (27-30) and 2020-07-08

(31, ex larva collected 2021-06-18). Photos by C. C. LaBar.

lcm

Figs. 32-35: Celastrina asheri sp. n. female type specimens. 32: allotype female dorsal (a) and ventral (b). 33-35: selection of

female paratypes. Specimens collected from Ringer South Trail on 2021-04-21 (32, 34) and 2021-06-12 (35, second brood, ex

larva collected 2021-05-28), and McCabe Pond fishing access area on 2021-04-22 (33), Kittitas County, WA. Photos by C. C.

LaBar.

Figs. 36-40: Celastrina asheri sp. n. females, all from Kettle Valley Rail Trail near Carmi, BC, leg. D. L. Threatful, 2009-05-22

to 24. Photos by N. G. Kondla.

i i ei Bie / : :% d

1 ph Cn

Fig. 41: Comparison of same Celastrina asheri sp. n. individual from upper Sinlahekin Creek, Sinlahekin Wildlife Area,

Okanogan County, WA. A) live male on 2013-05-11 exhibiting pearlescent sheen and blue-gray coloration. B) same specimen

photographed 2021-05-08 showing ventral coloration faded to brownish gray (=Fig. 27). Photos by C. C. LaBar.

Specimens examined. Holotype male (Fig. 10) with the following labels: white, printed: / WA: Kittitas

Co. 2021-Apr-21 / Ringer South Trail, Ringer Loop Rd. / short walking trail at the Yakima River /

N46.92757 W120.51949 elev. 1430’ / coll. Caitlin C. LaBar / ; red, printed: / HOLOTYPE / Celastrina

asheri / LaBar, Pelham & Kondla 2022 /. Allotype female (Fig. 32) with same data. The holotype,

allotype and a selection of male paratypes will be deposited at the McGuire Center for Lepidoptera &

Biodiversity, Florida Museum of Natural History, University of Florida, Gainesville (MGCL). Twenty-

four male and | female paratypes, same data as holotype. Five additional paratypes from same locality:

2021-06-12, C. C. LaBar (1¢ and 19, ex larvae); 2021-06-21, C. C. LaBar (26, wild-caught), 2022-

03-15 (16, ex larva). Three paratypes from McCabe Pond fishing access area southeast of Ringer South

Trail, Kittitas Co., WA, 2021-04-21 (24) and 2021-04-22 (19), C. C. LaBar. Data for additional

specimens (not paratypes) examined are provided only for specimens in CCL collection, UWBM and a

selection of specimens from NGK collection. CANADA: ALBERTA: Crowsnest River near Hillcrest,

N49.5504 W114.3480 elev. 1230m, 2009-05-29, N. G. Kondla (54, NGK); South Castle River Valley,

vic. N49.3081 W114.2938 elev. 1400-1460m, 1988-05-16, N. G. Kondla (14, NGK); same locality,

1982-05-24, N. G. Kondla (14, NGK); same locality, 1983-05-22, N. G. Kondla (146, NGK); BRITISH

COLUMBIA: Beaver Creek at Columbia River, vic. N49.0658 W117.6103 elev. 410-450m, 1999-04-

11, N. G. Kondla (146, NGK); same locality, 2006-04-24, N. G. Kondla (14, NGK); Blackwater Rd

near Clearwater, N51.573 W120.149 elev. 408m, 1994-05-22, N. G. Kondla (12, NGK); Brilliant, vic.

N49.3177 W117.6439 elev. 420-430m, 2002-04-18, N. G. Kondla (14, NGK); same locality, 2002-04-

19, N. G. Kondla (34', NGK); same locality, 2002-05-01, N. G. Kondla (74, NGK); same locality,

2002-05-10, N. G. Kondla (4¢, 12, NGK); same locality, 2002-05-13, N. G. Kondla (14, NGK); same

locality, 2002-06-03, N. G. Kondla (12, NGK); Charbonneau Creek, vic. N49.0470 W117.4753 elev.

560-870m, 1998-04-26, N. G. Kondla (54, NGK); same locality, 1998-04-29, N. G. Kondla (3¢,

NGK); same locality, 2000-04-08, N. G. Kondla (13, NGK); same locality, 2000-05-19, N. G. Kondla

(13, NGK); same locality, 2002-04-27, N. G. Kondla (124, NGK); same locality, 2002-05-11, N. G.

Kondla (53, NGK); same locality, 2002-05-12, N. G. Kondla (274, 12, NGK); same locality, 2002-

05-19, N. G. Kondla (203, 12, NGK); same locality, 2002-05-31, N. G. Kondla (64, NGK); same

locality, 2003-04-20, N. G. Kondla (34, NGK); same locality, 2003-04-21, N. G. Kondla (14, NGK):

same locality, 2003-04-28, N. G. Kondla (1124, NGK); same locality, 2004-04-25, N. G. Kondla (12,

NGK); same locality, 2004-04-26, N. G. Kondla (354, NGK); same locality, 2006-04-30, N. G. Kondla

(13, NGK); Ellison Ridge, vic. N50.1264 W119.4129 elev. 460-1040m, 2001-04-26, D. L. Threatful

(73, 22, NGK); Fort Shepherd flats, N49.0025 W117.62 elev. 411m, 2006-05-05, N. G. Kondla (2¢,

NGK); same locality, 2006-05-14, N. G. Kondla (16, NGK); km 2-3 Handley Road, Pend-d’ Oreille,

vic. N49.0270 W117.5348 elev. 580-610m, 2002-06-02, N. G. Kondla (34, NGK); Hulme Creek Rd,

N49.0885 W119.0237 elev. 1030m, 2009-06-01 or 03, D. L. Threatful (14, NGK); Kettle Valley Rail

Trail near Carmi, vic. N49.4896 W119.1187 elev. 820-840m, 2009-05-22 to 24, D. L. Threatful (23,

52, NGK); Kid Creek Forest Service Road at Hwy 3, vic. N49.1626 W116.2853 elev. 760-770m, 2003-

05-24, N. G. Kondla (24, NGK); Marsden Road west of Nelson, vic. N49.4957 W117.3818 elev. 530-

550m, 2002-04-29, N. G. Kondla (9¢', NGK); Ootischenia, vic. N49.2787 W117.6326 elev. 430-550m,

2002-06-25, N. G. Kondla (134', NGK); same locality, 2003-04-30, N. G. Kondla (2, NGK); same

locality, 2006-04-27, N. G. Kondla (12, NGK); 1 km south of Ootischenia, vic. N49.2690 W117.6240

8

elev. 550-570m, 2002-05-16, N. G. Kondla (14, NGK); Rock Mountain Forest Service Road near Rock

Creek, N49.0336 W119.053 elev. 970m, 2001-05-06, N. G. Kondla (14, NGK): km 2 west of Salmo

River, Pend-d’ Oreille valley, vic. N49.1912 W117.3077 elev. 690-700m, 2002-04-27, N. G. Kondla

(43, NGK); same locality, 2002-05-12, N. G. Kondla (334, NGK); same locality, 2002-06-23, N. G.

Kondla (24, NGK); km 2.7 west of Seven Mile Dam, N49.0221 W117.5338 elev. 480m, 2002-04-27,

N. G. Kondla (184', NGK); same locality, 2002-05-11, N. G. Kondla (574, 22, NGK); same locality,

2002-05-12, N. G. Kondla (184, NGK); same locality, 2002-05-19, N. G. Kondla (54, NGK); same

locality, 2002-05-31, N. G. Kondla (414, NGK); same locality, 2002-06-02, N. G. Kondla (20¢, 19,

NGK); same locality, 2002-06-12, N. G. Kondla (7, NGK); same locality, 2003-04-20, N. G. Kondla

(164, NGK); same locality, 2003-04-21, N. G. Kondla (35, NGK); same locality, 2003-04-28, N. G.

Kondla (264, NGK); same locality, 2003-05-01, N. G. Kondla (194, NGK); same locality, 2004-04-

09, N. G. Kondla (34, NGK);same locality, 2004-04-25, N. G. Kondla (8), NGK):same locality, 2004-

04-26, N. G. Kondla (64', NGK); same locality, 2005-04-22, N. G. Kondla (34, NGK); same locality,

2006-04-20, N. G. Kondla (134', NGK); same locality, 2006-05-04, N. G. Kondla (14, NGK); same

locality, 2006-05-04, N. G. Kondla (4¢', NGK); km 4 west of Seven Mile Dam, N49.0160 W117.5430

elev. 480m, 2002-05-11, N. G. Kondla (3, NGK); same locality, 2002-05-12, N. G. Kondla (903,

NGK); same locality, 2003-05-23, N. G. Kondla (2, NGK); same locality, 2006-05-10, N. G. Kondla

(13, NGK); km 3 east of Seven Mile Dam, N49.0446 W117.4823 elev. 585m, 2003-05-10, N. G.

Kondla (12, NGK); km 3.5 east of Seven Mile Dam, N49.0469 W117.4762 elev. 600m, 2002-05-11,

N. G. Kondla (14, NGK); same locality, 2002-05-19, N. G. Kondla (14, NGK); same locality, 2002-

05-31, N. G. Kondla (54, NGK); same locality, 2002-06-02, N. G. Kondla (34, NGK); same locality,

2002-06-12, N. G. Kondla (13', NGK); same locality, 2003-05-01, N. G. Kondla (3, NGK); same

locality, 2003-05-28, N. G. Kondla (14, NGK); same locality, 2003-05-29, N. G. Kondla (14, NGK):

km 5.1 east of Seven Mile Dam, N49.0486 W117.4596 elev. 585m, 1999-04-24, N. G. Kondla (16,

NGK); same locality, 2002-04-20, N. G. Kondla (30, NGK); same locality, 2002-04-27, N. G. Kondla

(63, NGK); same locality, 2002-05-11, N. G. Kondla (334, 12, NGK); same locality, 2002-05-12, N.

G. Kondla (64, NGK); same locality, 2002-05-19, N. G. Kondla (64, NGK); same locality, 2002-05-

31, N. G. Kondla (12¢', NGK); same locality, 2002-06-02, N. G. Kondla (14, NGK); same locality,

2002-06-21, N. G. Kondla (23', NGK); same locality, 2003-04-21, N. G. Kondla (14, NGK); same

locality, 2006-04-24, N. G. Kondla (24, NGK); same locality, 2004-03-29, N. G. Kondla (14, NGK):

same locality, 2006-04-20, N. G. Kondla (34, NGK); same locality, 2006-04-24, N. G. Kondla (2¢,

NGK); km 5.2 east of Seven Mile Dam, N49.0486 W117.4576 elev. 585m, 2002-06-23, N. G. Kondla

(23, NGK); km 6 east of Seven Mile Dam, N49.0483 W117.4483 elev. 590m, 2002-05-31, N. G.

Kondla (144, 12, NGK): same locality, 2005-04-22, N. G. Kondla (12, NGK); km 6 to 7 east of Seven

Mile Dam, vic. N49.0490 W117.4398 elev. 580-610m, 2002-05-12, N. G. Kondla (174', 29, NGK); km

7 east of Seven Mile Dam, N49.0490 W117.4347 elev. 580m, 2002-04-20, N. G. Kondla (23', NGK);

same locality, 2002-04-27, N. G. Kondla (75, NGK); same locality, 2002-05-31, N. G. Kondla (3¢,

NGK); same locality, 2002-06-23, N. G. Kondla (1¢', NGK); same locality, 2003-04-19, N. G. Kondla

(53, 32, NGK):; same locality, 2003-04-21, N. G. Kondla (24, NGK); same locality, 2003-05-01, N.

G. Kondla (24, NGK); same locality, 2003-05-23, N. G. Kondla (14, NGK); same locality, 2004-04-

25, N. G. Kondla (14, NGK); same locality, 2005-04-10, N. G. Kondla (14, NGK); same locality,

2006-04-20, N. G. Kondla (24', NGK): same locality, 2006-04-30, N. G. Kondla (4<, NGK); Shadey

Creek Road, 2005-04-30, N. G. Kondla (14, NGK); Silver Star Mountain, vic. N50.3884 W119.0941

elev. 1150m, 2002-06-04, D. L. Threatful (12, NGK); Tillicum Creek/Meadow, Pend-d’ Oreille valley,

vic. N49.0387 W117.4297 elev. 580-980m, 2002-06-23, N. G. Kondla (3, NGK); same locality, 2003-

05-01, N. G. Kondla (54, NGK); same locality, 2002-05-12, N. G. Kondla (74, NGK); same locality,

2007-04-30, N. G. Kondla (184, 19, NGK); same locality, 2006-04-30, N. G. Kondla (84, NGK):

same locality, 2006-05-04, N. G. Kondla (154, NGK):; mountainside above Waneta Dam, vic. N49.007

W117.611 elev. 450-518m, 2006-04-30, N. G. Kondla (1d, NGK); USA: MONTANA: Lincoln Co.:

Bad Medicine campground, Bull Lake, N48.22 W115.858 elev 2370ft (722m), 2006-05-19, N. G.

Kondla (12, NGK); Dorr Skeels campground, Bull Lake, N48.268 W115.855 elev 2350ft (716m),

2006-05-19, N. G. Kondla (14, NGK); Missoula Co.: Miller Creek, vic. N46.765 W113.942 elev.

3730-4000ft (1137-1219m), 1976-05-12, S. Kohler (14, NGK); Sanders Co.: Forest Road 1022

(McKay Creek Rd) near Noxon, N47.968 W115.696 elev. 2450-2850ft (747-869m), 2006-05-19, N. G.

9

Kondla (114, NGK); WASHINGTON: Columbia Co.: Hompegg Falls, North Fork Touchet River,

N46.1656 W117.8126 elev. 3300ft (1006m), 1971-05-22, J. P. Pelham, R. E. Miller & F. Van Buskirk

(113, 42, UWBM); same locality, 1975-05-21, J. P. Pelham & T. S. Pelham (60, 52, UWBM); Wolf

Fork Touchet River at Whitney Creek, N46.18109 W117.8624 elev. 2720-2760ft (830-840m), 1989-04-

09, L. G. Crabo & J. P. Pelham (1¢, UWBM); Kittitas Co.: State Hwy 10, Yakima River, N47.1342

W120.7863 elev. 1845ft (562m), 1961-05-04, R. Cheyne (58, UWBM); Reecer Creek Road parking

loop, N47.1703 W120.5923 elev. 3400-3500ft (1036-1067m), 1978-07-01, J. P. Pelham & R. Massey

(23, UWBM); “azure corner” at the mouth of Reecer Canyon, N47.17526 W120.58819 elev. 3560ft

(1085m), 1979-05-24, J. P. Pelham, T. S. Pelham & S. F. Jones (14, UWBM); same locality, 2005-04-

26, C. C. LaBar (13, CCL); same locality, 2009-05-22, C. C. LaBar (2¢', CCL); Naneum Basin/Naneum

Road at bottom of talus slope in Cornus sericea thicket, N47.17788 W120.44789 elev. 2980ft (908m),

2021-05-29, C. C. LaBar (6¢, 32, CCL); Naneum Basin/Naneum Road at pullout along creek,

N47.17965 W120.44689 elev. 2980ft (908m), 2021-05-29, C. C. LaBar (3, 22, CCL); Dawson Road

seep, N47.12705 W120.40138 elev. 3220ft (981m), 2021-05-30, C. C. LaBar (1¢', CCL); Robinson

Canyon, [N47.0142 W120.7107] elev. 2200-2400ft (670-73 1m), 1967-05-15, R. Cheyne (12, UWBM);

Wilson Creek (Canyon), [N47.1542 W120.5077] elev. 2800-3000ft (853-914m), 1966-07-15, R.

Cheyne (14, UWBM); Wilson Creek Road at Bar 14 Road, N47.0734 W120.4961 elev. 2050-2060ft

(625-628m), 1966-07-18, R. Cheyne (22, UWBM); Yakima River at US Hwy 90, N47.1858 W121.0432

elev. 2010-2025ft (613-617m), 1958-04-05, D. Carney (12, UWBM); same locality, 1958-04-11, D.

Carney (53, UWBM); same locality, 1958-04-28, D. Carney (2¢', UWBM);: same locality, 1958-05-28,

D. Carney (12, UWBM); same locality, 1972-05-28, D. Carney (12, UWBM); Klickitat Co.: Satus

Creek at US Hwy 97, N46.0268 W120.6244 elev. 2139ft (652m), 1955-05-07, G. Schenk (106, 49,

UWBM); Okanogan Co.: Chiliwist Rd, N48.3085 W119.8479 elev. 2640ft (805m), 1978-04-28, R. M.

Pyle (23, 32, UWBM); Chiwiliken Valley Road, N48.5813 W119.2525 elev. 4415-4425ft (1346-

1349m), 2006-06-17, D. & J. Nunnallee (14, UWBM); Sinlahekin Wildlife Area, west canyon,

N48.68861 W119.74018 elev. 2765ft (843m), 2013-05-11 (34, CCL); same locality, 2020-07-08, C. C.

LaBar (13, ex larva collected 2021-06-18, CCL); same locality, 2022-03-03, C. C. LaBar (14, 19, ex

larvae collected 2021-06-18, CCL); same locality, 2022-03-13, C. C. LaBar (13, 19, ex larvae collected

2021-06-18, CCL); same locality, 2022-03-14, C. C. LaBar (19, ex larva collected 2021-06-18, CCL);

same locality, 2022-03-15, C. C. LaBar (16, ex larva collected 2021-06-18, CCL); Pend Oreille Co.:

Sullivan Lake Rd, N48.8605 W117.3285 elev. 2400ft (732m), 1982-06-16, R. M. Pyle (12, UWBM);

State Hwy 31 at Threemile Creek, N48.8938 W117.3280 elev. 2550ft (777m), 1982-06-16, R. M. Pyle

(13, UWBM); East Fork Leclerc Creek Rd, N48.561 W117.2724 elev. 2500ft (762m), 1982-06-18, R.

M. Pyle (12, UWBM); Horseshoe Lake Road, N48.09033 W117.4276 elev. 2225ft (678m), 2018-05-

26, C. C. LaBar (13, 19, CCL); Yakima Co.: Bear Canyon, N46.7096 W120.9043 elev. 2000-2500ft

(700-762m), 1968-04-09, J. P. Pelham, L. Obata & H. Wiles (14, UWBM); same locality, 1968-04-28,

J. P. Pelham & H. Wiles (14, UWBM)); same locality, 1989-05-06, J. P. Pelham, K. Hiruma, L. Crabo

(53, 12, UWBM);: Bear Canyon, N46.6997 W 120.9383 elev. 2985-304 0ft (910-927m), 1973-04-07, J.

P. Pelham (13, UWBM); Cowiche Canyon, N46.6281 W120.6605 elev. 1465-1470ft (447-448m),

2003-04-03, J. P. Pelham, K. Hiruma, D. Nunnallee & S. Wechsler (2¢', 29, UWBM); Cowiche Creek

Canyon, N46.625019 W120.650809 elev. 1430ft (436m), 2003-04-12, J. P. Pelham, Z. A. Pelham, D.

Nunnallee & K. Hiruma (62, UWBM); Cowiche Mill Rd, N46.6488 W120.8479 elev. 2330-2550ft

(710-777m), 2004-04-24, J. P. Pelham, K. Hiruma, M. W. Robinson, et al. (14, UWBM); Oak Creek,

N46.7303 W120.8395 elev. 1900-2100ft (579-640m), 1995-05-07, J. F. Rial (16, UWBM); Oak Creek

Rd, N46.7338 W120.9292 elev. 2800ft (853m), 1993-05-18, J. F. Rial (1¢, UWBM); Signal Peak

Road, N46.2954 W120.7592 elev. 1800ft (549m), 1959-04-25, D. Carney (12, UWBM).

10

Type locality. Ringer South Trail, five miles south of Ellensburg in Kittitas County, Washington.

This is a Bureau of Land Management river access site next

to the Yakima River at the south end of Ringer Loop with a

short loop trail through a thicket of red osier dogwood (Cornus

sericea), Wild rose (Rosa spp.) and willow (Salix spp.) shaded

by black cottonwood (Populus trichocarpa) and scattered

ponderosa pine (Pinus ponderosa) (Fig. 42). It is subject to

occasional flooding by the Yakima River.

Etymology. Celastrina asheri is named in honor of 8-year-old

Asher (Fig. 43). Surname is not given in accordance with family

wishes. Asher suffers from an extremely rare neurological

disease called 4H (or POLR3B-related) Leukodystrophy. His

perseverance and kind heart inspire everyone around him,

bringing rays of sunshine into the daily struggle of living with

a disease that has no cure. Asher loves blue, which is also the

representative color of Leukodystrophy, and he loves spending

time outdoors, especially in the mountains. In naming this

butterfly Celastrina asheri, Asher’s Blue, it is our wish to bring

joy to Asher and his family and help promote awareness of this

disease with the hope that one day a cure will be found.

jis

Ls

F

ig. 43: Asher, our butt

Le ey A

erfly’s namesake.

(xs Columbia River

Watershed

Type Locality

8 C. asheri

© C. echo nigrescens

Celastrina Records

© C. asheri

© C lucia

C. echo

——

St

Ae Boat

> - |

es ~GEBCO,,NOAA

a Ame 28

Fig. 44: Map of Celastrina asheri sp. n., C. lucia and C. echo records in northwestern United States and southwestern Canada.

Some C. echo records may turn out to be C. asheri but unless we were able to confirm this via images or specimens, we chose

to accept the data as they are currently recorded.

Distribution and phenology. The range of this species encompasses most of the Columbia

River Basin from southeast British Columbia, through Washington east of the Cascade crest,

south to the Blue Mountains in northeast and central Oregon and east through Idaho into

western Montana and the southwest corner of Alberta (Fig. 44). There is one record from

the base of Steens Mountain in Harney County, Oregon. A described “aberration” of C. echo

from Lake County, Oregon may in fact be C. asheri, see Diagnosis and discussion section

below. Currently, the extent of the southeastern edge of the range is uncertain, in part due to

the taxonomic uncertainty over the past several decades of record keeping. Examination of

specimens from this region and additional field work is needed to clarify the entire distribution

of C. asheri.

Celastrina asheri is primarily univoltine, flying from late March to mid-June depending on

elevation and latitude; however, in some areas a small second flight appears approximately

two months after the first, peaking in mid-July (Fig. 45). C. lucia from Montana, southern

British Columbia and southern Alberta exhibit a similar pattern but the flight is shifted one to

two weeks after C. asheri (Fig. 45). In contrast, the flight period of C. echo varies widely: the

overall peak flight east of the Cascades occurs mid to late May, but a smaller peak occurring

in late April at lower elevations and warmer habitats and another small peak occurring in mid-

|

June are likely a combination of high elevation first brood and low elevation second brood

emergence records (Fig. 45). Where C. asheri flies in sympatry with C. echo, which species

emerges first seems to vary by location based on the personal observations of the authors. In

central Washington, the first and second authors usually observe C. asheri emerging prior to

C. echo, while in the Pend-d’ Oreille River valley in south-central British Columbia, the third

author has observed C. echo emerging prior to C. asheri (Fig. 46).

The second brood of C. asheri is interesting both in phenotype and comparative phenology with

C. echo. While C. echo frequently has a partial second brood in coastal areas, and possibly even

a third brood in some places as evidenced by late records into November, multiple emergences

of C. echo are rarer east of the Cascades. For example, iNaturalist records dated between

June and August show 5 C. echo and 25 C. asheri records east of the Cascades and over 300

records of C. echo west of the Cascades. James & Nunnallee (2011) reported around 30% of

their lab-reared C. asheri (“C. lucia’) emerged as a second brood. A similar ratio was observed

by the first author from C. asheri larvae collected at the type locality: 2 males and 1 female

160

150

ah Wc. asheri

120 MC. echo

, 110

+. 100 MC. lucia

S 90

x g0

Ss 70

© 60

50

40

30

20

: es '

0 = rae =_— = = —

> oD PB O D OP SH PPL Ly a PD PD OD YW

& = = s = ~~ ~~ ~ a Ss s Ss = OR: & Ry ey S

ee ee ee FS wr ey Wy ws

Month (Week)

Fig. 45: Flight period of Ce/astrina asheri sp. n. compared with C. /ucia and C. echo from within the same geographical area

east of the Cascade crest. Records are grouped into weeks.

Comparison of adult numbers over a six week period in 2002

120

100

= mC.echo WC. asheri

= 80

2)

x

° 60

rs

=

sua

20 ,

O ~ | al a | |

x 4

OS PP SF Oe rere eo re tt

No ss vy \v vy ne a no ave ay a\e Ao

Fig. 46: Comparison of adult Celastrina asheri sp. n. and C. echo collected (n=866) over a six week period in 2002 from

multiple locations in the vicinity of the Pend-d’ Oreille River valley near the Columbia River in southeast British Columbia by

Norbert Kondla. Specimens collected on a “next up” basis with no bias for phenotype.

13

eclosed from 12 larvae reared to pupae in June 2021. The first author also collected 2 males and

observed at least one other male flying at the type locality on 2021-06-21 and Melanie Weiss

observed more adults over the following two weeks at the same location in numbers roughly

equaling 20-25% of the first brood. In contrast, from 14 larvae collected by the first author at

the Sinlahekin Wildlife Area in Okanogan County, Washington, only | male eclosed in the

same year. Additionally, the first and third authors have visited northern Okanogan County,

Washington and the Pend-d’Oreille Valley, British Columbia, respectively, numerous times in

June and July over several years. During these visits, no Ce/astrina were ever recorded by the

first author and only a very small number were recorded by the third author.

Biology. C. asheri eggs were photographed at Naneum Basin, Kittitas County, Washington on

2021-05-29 (Figs. 48, 49), where five female C. asheri were observed ovipositing on Cornus

sericea. Eggs were tucked between the flower buds. Two female C. echo were also captured

flying around the dogwood where Ceanothus velutinus was abundant on the opposite slope

across the road. Around 20 third and fourth instar larvae were found (12 collected) at the

Ringer South Trail type locality on 2021-05-28. All were medium green, one had extensive

dorsal and lateral rose-brown markings (Fig. 58), at least three had a rose-brown patch on

the prothoracic shield (Fig. 63), three had a pale green ventrolateral line (Figs. 53, 57), one

had faint whitish dorsal chevrons (Fig. 58), and most had a slightly darker green dorsal stripe

(Figs. 62, 63). Around 24 larvae were found (14 collected) at the Sinlahekin Wildlife Area,

Okanogan County, Washington on 2021-06-18. The second instars were pale yellow-green

(Figs. 50, 51). All third and fourth instar larvae were medium green with a slightly darker green

dorsal stripe (Fig. 65), one had some white dorsal markings and white ventrolateral line in the

third instar (Fig. 55) which persisted through the fourth instar. At least four had a rose-brown

patch on the prothoracic shield (Figs. 59, 64), all others were unmarked (Figs. 60, 61, 65).

Figs. 47-54: Immature stages of Celastrina echo (47) and C. asheri sp. n. (48-54). 47: C. echo third instar larva found on

Ceanothus velutinus at Swakane Canyon, Chelan County, WA, 2009-06-28. 48-49: C. asheri unhatched (48) and hatched (49)

eges on Cornus sericea at Naneum Basin, Kittitas County, WA, 2021-05-29. 50-51: C. asheri second instar larvae on Co.

sericea at the Sinlahekin Wildlife Area, Okanogan County, WA, 2021-06-18. 52-54: C. asheri third instar larvae on Co. sericea

at Ringer South Trail, Kittitas County, WA, 2021-05-28.

14

Figs. 55-71: Immature stages of Celastrina asheri sp. n. 55: C. asheri third instar larva on Co. sericea at the Sinlahekin

Wildlife Area, Okanogan County, WA, 2021-06-23. 56-58: C. asheri late third or early fourth instar larvae on Co. sericea at

Ringer South Trail, Kittitas County, WA, 2021-05-28. Photos by C. C. LaBar. 59-61 & 64: fourth instar larvae on Co. sericea

at the Sinlahekin Wildlife Area, Okanogan County, WA, 2021-06-18, all being tended by Formica sp. ants. 62-63: fourth instar

larvae on Co. sericea at Ringer South Trail, Kittitas County, WA, 2021-05-28. 65: fourth instar larva on Co. sericea at the

Sinlahekin Wildlife Area, Okanogan County, WA, 2021-06-17. 66: early pre-pupal larva that has recently stopped feeding, from

the Sinlahekin Wildlife Area, Okanogan County, WA, 2021-06-17. 67: late pre-pupal larva from Ringer South Trail, Kittitas

County, WA, 2021-06-04. 68-71: newly formed (68-69) and few-day-old (70-71) pupae from Ringer South Trail, Kittitas

County, WA, 2021-06-02 (68, 69, 71) and 2021-06-04 (70). Photos by C. C. LaBar.

1)

* Be Ga ae

Figs. 72-77: Immature stages of Celastrina echo (72-75) and C. asheri sp. n. (76-77). 72-73: C. echo third instar larvae on

Ceanothus velutinus at Reecer Canyon, Kittitas County, WA, 2009-06-06. 74-75: C. echo fourth instar larva on Ceanothus

velutinus at Reecer Canyon, Kittitas County, WA, 2009-06-10. 76-77: C. asheri fourth instar larva on Cornus sericea at Black

Canyon, Okanogan County, WA, 2009-05-16. Photos by D. Nunnallee.

Some of the fourth instar larvae were tended by Formica sp. ants (Figs. 59-61, 64). The rose-

brown coloration on some fourth instar larvae appears to offer excellent camouflage among

the Cornus sericea seedpods which are tipped with the same color (Fig. 64). It is unknown if

this pattern occurs widely in C. asheri larvae. All larvae from both locations lost all markings

and turned rose-brown after leaving the food plant and searching for a spot to pupate (Fig. 66).

Pre-pupal larvae are translucent turquoise and beige (Fig. 67). Newly formed pupae are light

pinkish brown with a dark brown dorsal stripe and numerous dark brown speckles (Figs. 68,

69), darkening to purplish or reddish brown and black within several days (Figs. 70, 71).

The only other known documentation of C. asheri immature stages is by James & Nunnallee

(2011). In their “C. echo nigrescens” account (pp. 192-193), all images are of C. asheri reared

from eggs gathered on April 10 and May | from Black Canyon, Okanogan County, Washington

(also see Figs. 76, 77) on Cornus sericea. A female collected on May 29 from Bear Canyon,

Yakima County, Washington, which laid eggs on Ceanothus sanguineus, was most likely C.

echo. Larvae from that female did not reach the final instar. In their “C. /ucia” account (pp. 194-

195), all images and descriptions are of C. asheri on Cornus sericea from Cowiche Canyon,

Yakima County, Washington. In both accounts, some adults emerged within two weeks of

pupating and either exhibited a smaller VHW discal patch (Cowiche Canyon “lucia”) or no

discal patch but heavy charcoal marginal markings (Black Canyon “nigrescens”’). These second

brood phenotypes are also seen on some UWBM specimens and iNaturalist observations of

C. asheri dated between June and July, as well as the individuals collected by the first author

(Figs. 24, 25, 31, 35).

16

Eggs and first and second instar larvae of C. asheri are identical to C. echo and C. lucia.

Larvae of all three species begin developing markings in the third instar and these usually

become stronger in the fourth (final) instar. C. echo larvae are known to be extremely variable

in coloration and markings throughout their range (James & Nunnallee, 2011; Stout, 2022; and

personal observations of the second author, J. P. Pelham). When C. echo larvae are closest in

appearance to C. asheri, there are a few identifiable differences. Pale green C. echo larvae with

only faint markings still have a pair of relatively dark green squares sometimes forming a bar

or “saddle” on the third segment behind the prothoracic shield and three dark green or brown

spots near the posterior (Fig. 47). These markings are even darker on fourth instar C. echo

that are also strongly marked with white dorsal chevrons (Figs. 74, 75) but may not always

be visible on third instar larvae (Figs. 72, 73). When visible on C. asheri, these spots are less

pronounced relative to the background color (Figs. 55, 56, 58, 62, 76). Also, when C. echo

exhibit dorsal white chevrons or lateral lines, they appear to be a brighter white (Figs. 72-75)

than the yellowish or greenish white seen on some C. asheri (Figs. 53, 55-57, 76, 77). Further

rearing studies are needed to determine the full range of variation in C. asheri larvae.

Very little is published about the immature stages of C. /ucia and even fewer images are

available. Schmidt & Layberry (2016) illustrate a range of variation in C. /ucia larvae from

Ottawa, Ontario, of which a consistent pattern appears to be a dark “saddle” on the third segment

behind the prothoracic shield and a dark dorsal stripe. These markings are both very similar

to C. echo and some C. asheri, but the dorsal stripe appears to stand out even on the palest C.

lucia larvae, unlike most C. echo and C. asheri. C. lucia are known to use a variety of larval

foodplants, including Cornus spp., Prunus spp., Rhododendron |Ledum| spp., Vaccinium spp..,

and Viburnum spp. (Bird, et al., 1995; Guppy & Shepard, 2001; Schmidt & Layberry, 2016).

A photo by Larry Everson (Pyle, 2002, p. 231) depicts an apparent C. asheri female ovipositing

on Holodiscus discolor flower buds in Jefferson County, Oregon. Warren (2005) reported

flushing several females and some males from H. discolor bushes along the Metolius River

near Camp Sherman, Jefferson County, Oregon. He confirms that these individuals were of the

lucia form (A. Warren, pers. comm.). Considering that his observations were of more females

than males, and most were worn, while typical C. echo males were seen at mud and flying

nearby, this seems to align with C. asheri usually having a peak flight slightly before C. echo,

further pointing to the strong probability of C. asheri using H. discolor as a larval food plant in

addition to Cornus sericea. Further research is needed to confirm if these are indeed C. asheri

using H. discolor as a larval food plant in Oregon and possibly elsewhere.

Fig. 78: Celastrina asheri sp. n. puddle party in the Pend- Fig. 79: Left to right: Ce/astrina asheri sp. n., C. echo, asheri,

d Oreille River valley, BC, 2003-04-28. Photo by N. G. echo in the Pend-d’Oreille River valley, BC, 2007-04-23.

Kondla. Photo by N. G. Kondla.

17

tt ~~, J - a

’ a“ 2

Fig. 80: Celastrina asheri sp. n. and C. echo puddle party in the Pend-d’ Oreille River valley, BC, 2007-04-30.

Photo by N. G. Kondla.

C. asheri appears to be highly adapted to xeric valleys and lowlands primarily within the

Columbia River watershed east of the Cascade crest. It is frequently associated with riparian

corridor habitats in at least the Washington and Montana parts of its range. These riparian

habitats are often devoid of flowers and many of the shrubs are only beginning to leaf out when

C. asheri begins its flight period, which historically led to the species being overlooked in these

areas by many lepidopterists early in its flight period. C. asheri males frequently visit Salix

spp. and were observed by the first author feeding on the sap around the catkins. Males are

also frequently observed at mud throughout their range, often mixed with C. echo (Figs. 78-80,

93, 94). Adults nectar on Crataegus spp., Ribes viscosissimum, Astragalus spp., Erythronium

grandiflorum, Linum lewisii, Lomatium spp., Maianthemum stellatum, Myosotis scorpioides,

Tanacetum vulgare, and likely other flowering shrubs and forbs.

Diagnosis and discussion. The dorsal color of male C. asheri is variable but almost never

as silvery blue as C. lucia (Fig. 96a vs. 96c) and rarely as violet as many C. echo (Figs. 89,

90). The dorsal blue coloration of female C. asheri tends to be more silvery and reaches the

DHW margin more often in central Washington populations (Figs. 32a-35a, 81, 82), similar

to C. lucia (Fig. 97c), while the northern populations of C. asheri tend to be more lavender

with broader dark gray margins (Figs. 36a-40a, 97a) similar to female C. echo (Figs. 2a, 4a,

5a, 7a, 87, 88). Thin, dark gray cell bars sometimes occur on the DFW of female C. asheri

and are almost always absent from the DHW (Figs. 81A, 82A) in contrast to C. echo females

in which the dorsal cell bar is almost always visible on both fore and hindwings (Figs. 87A,

88A) and C. /ucia females in which these cell bars are almost always absent (Fig. 97c). The

DHW margin of female C. asheri ranges from having small, fuzzy gray spots on a mostly blue

wing (Fig. 32-34a, 36a, 37a, 82B) similar to female C. /ucia (Fig. 97c), to having indistinct

submarginal gray crescents separating the blue discal area from a partially blue marginal band

around the spots (Figs. 38a, 40a, 81B, 97a). The area between the ventral marginal spots and

18

submarginal crescents of both sexes usually contains varying amounts of dark gray scales

(Figs. 85D) compared to C. echo in which this band never contains dark gray scales (Figs. 91D,

92D). The dorsal wing fringe of C. asheri varies from lightly checkered, usually with light gray

mixed into the white scales between the dark gray fringe at the tips of each wing vein (Figs.

81-83C) to solid gray or black (Fig. 84C), compared to C. echo in which the fringe is solid

white or white with a thin black line at the tips of each wing vein (Figs. 87-90C), and C. lucia

in which the fringe is usually heavily checkered with very little variation (Figs. 96c, 97c). C.

echo never exhibit the large, dark gray VHW discal patch seen in some C. asheri and C. lucia.

When C. asheri exhibits a lighter ventral phenotype resembling C. echo, the two species may

be differentiated by the bolder markings on C. asheri, particularly of the VHW discal cell bar

and median spots (Fig. 86E), the appearance of fuzzy, white postmedian arrowheads on some

C. asheri (Fig. 85F) or the frequent duskiness in the VHW marginal band (Figs. 28b, 39b, 40b,

85D, 94), compared to the pale ventral and thin VHW discal cell bar on C. echo (Figs. 91E,

92E, 93). A less consistent but somewhat useful identifier is the coloration of the basal area of

the ventral wings: C. echo tends to have a dusting of greenish-blue scales over this area (Figs.

91G, 92G), C. asheri may have a very small amount of greenish-blue (less often) or dark gray

(more often) scales in this area (Figs. 10b-40b, 85G, 86G, 96b, 97b), while C. /ucia has dark

gray to black scales in this area (Figs. 95, 96d, 97d). C. asheri and C. lucia are sympatric in

only a few places in British Columbia, Alberta and Montana and may be differentiated by

a combination of features. In addition to the dorsal color tint and other differences already

described, C. asheri average slightly larger than C. /ucia and the ventral of sympatric C. lucia

populations tends to be more evenly mixed with dark gray and light gray scales giving it a

“dirty” appearance overall (Fig. 95) compared to the lighter gray of C. asheri. C. lucia from

Saskatchewan and eastward exhibit “cleaner” ventrals very similar to some C. asheri.

C. asheri

C. echo

Figs. 81-92: Comparison of Ce/astrina asheri sp. n. (females 81-82, males 83-86) and C. echo (females 87-88, males 89-92),

all specimens from Washington. Letters indicate key features: A) presence or absence of dorsal cell bar on females, B) presence

or absence of marginal spots and surrounding blue and charcoal shading on DHW of females, C) dorsal wing fringe color, D)

clarity and color of ventral marginal band pattern, E) size of VHW median spots and discal cell bar, F) postmedian pattern of

white “arrowheads” standing out from ground color, and G) presence or absence of basal greenish-blue scales. Photos by C.

C. LaBar.

19

Fig. 93: A sample of Ce/astrina echo from the Pend-d’ Oreille River valley, BC. Photos by N. G. Kondla.

21

Fig. 97: Comparison of Celastrina asheri sp. n. (rows a & b) and C. lucia (rows c & d) females. Photos by N. G. Kondla.

22

The type specimen of Cyaniris ladon ab. nunenmacheri Strand, 1915 exhibits a discal patch

on the VHW and brownish coloration on the VFW typical of C. asheri, but has a mostly-white

dorsal wing fringe typical of C. echo (http://butterfliesofamerica.com/L/ih/celastrina0004_1.

htm and _ http://butterfliesofamerica.com/L/th/celastrina0003_1.htm). It is described as an

aberration, therefore the name nunenmacheri is unavailable to use for a new taxon. Also, it

is from Lake County, Oregon, where no other records of C. asheri have been located. At this

time, we consider it likely to be a normal C. asheri and not an aberration of C. echo, but until

further evidence is gathered indicating the occurrence of C. asheri in Lake County, it will

remain a synonym of C. echo echo.

CONCLUSION

The presence of lucia-like phenotypes in Washington noted by Newcomer in 1964 has not been

resolved until now. From that time forward, individual azures exhibiting lucia-like features were recorded

irregularly in a largely non-geographic pattern. When purported C. /ucia was identified as a second azure

species in Washington, it was speculated that introgression between it and C. echo explained the occurrence

of these phenotypes among otherwise “normal” C. echo populations. We demonstrate that C. asheri is

widely sympatric with C. echo, narrowly sympatric with C. /ucia and all three species are separated by

subtle but consistent phenotypic features accompanied by habitat, phenology, foodplant and life history

differences.

ACKNOWLEDGMENTS

David James, Steve Kohler, Harry Pavulaan, Paul Severns, Jon Shepard, Todd Stout and David

Wright were generous with information and perspective as well as critical review. Dave Nunnallee

contributed volumes of information concerning immature stages and the general life history of Ce/astrina

echo and C. asheri in Washington and provided important photographic images of larvae for use in this

paper. Andrew Warren was instrumental in providing knowledge of Ce/astrina in Oregon, both via his

2005 book and personal communication. John and Robin LaBar assisted the senior author in collecting the

adults and larvae for the type series from Ringer Loop, as well as a number of adults from Naneum Basin

and Reecer Canyon. Nick Grishin provided valuable insight into Celastrina genomics. Members of the

Pacific Northwest butterfly community supplied many records and photographs analyzed for this paper.

They are many and we apologize for inevitable oversight: Jeannette Barreca, John Baumann, Chery Bellin,

Gary Bernard, Keith Brady, Cathy Clark, Jeanne Dammarell, Dan Dunphy, Ben Mous, Jeff Pippen, Dale

Swedberg, Jeff Thompson, Melanie Weiss, Bill Yake. We are grateful to have such a vibrant community!

LITERATURE CITED

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, AB. 349 pp.

Clench, H. K. 1944. Two new subspecies of Lycaenopsis pseudargiolus Bdy. & Lec. (Lepidoptera,

Lycaenidae). Journal of the New York entomological Society 52(3): 273-276.

Clench, H. K. 1972. Celastrina ebenina, a new species of Lycaenidae (Lepidoptera) from the eastern

United States. Annals of the Carnegie Museum 44(4): 33-44.

Downey, J. C. 1961. Tribe Plebejini. (pp. 199-242) In P. Ehrlich & A. Ehrlich, How to Know the

Butterflies. W. C. Brown Co., Dubuque, IA.

Fletcher, J. 1903. Descriptions of some new species and varieties of Canadian butterflies. Proceedings

and Transactions of the Royal Society of Canada (2nd series) 9(Sect. IV): 207-216.

Guppy, C. S. & J. H. Shepard. 2001. Butterflies of British Columbia. Including western Alberta, southern

2

Yukon, the Alaska Panhandle, Washington, Northern Oregon, northern Idaho, northwestern

Montana. Royal British Columbia Museum, University of British Columbia Press, Vancouver-

Toronto, 414 pp.

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

Press, Corvallis, OR, 447 pp.

Langston, R. L. 1975. Genus Celastrina Tutt. (pp. 335 337) In W. Howe (Ed.), The Butterflies of North

America. Doubleday and Co., Garden City, NY, 633 pp.

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

Press, Toronto, Buffalo & London, 280 pp.

Miller, L. D. & F. M. Brown 1981 A catalogue/checklist of the butterflies of America North of Mexico.

Memoir. The Lepidopterists’ Society 2: vil + 280 pp.

Newcomer, E. J. 1964. Butterflies of Yakima County, Washington. Journal of the Lepidopterists’ Society

18(4): 217-228.

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

AQ: 658 pp.

Pelham, J. P. 2022. A catalogue of the butterflies of the United States and Canada. http://www.

butterfliesofamerica.com/US Can Cat.htm

Pyle, R. M. 2002. The Butterflies of Cascadia. A field guide to all the species of Washington, Oregon,

and surrounding territories. Seattle Audubon Society, Seattle, WA, 420pp.

Pyle, R. M & C. C. LaBar. 2018. Butterflies of the Pacific Northwest. Timber Press, Portland, OR, 461

PP.

Schmidt, B. C. & R. A. Layberry. 2016. What Azure blues occur in Canada? A re-assessment of

Celastrina Tutt species (Lepidoptera, Lycaenidae). ZooKeys 584: 135-164.

Stout, T. 2022. Raising Butterflies: Celastrina ladon echo immatures Photos. http://www.

raisingbutterflies.org/celastrina-ladon-echo-immature/

Warren, A. D. 2005. Butterflies of Oregon: their taxonomy, distribution, and biology. Contributions of

the C.P. Gillette Museum of Arthropod Diversity, Colorado State University, Fort Collins, 405

PP.

Warren, A. D., K. J. Davis, E. M. Stangeland, J. P. Pelham, K. R. Willmott and N. V. Grishin.

2017. Illustrated Lists of American Butterflies (North and South America). http://www.

butterfliesofamerica.com/L/Neotropical.htm

24

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