Celastrina asheri LaBar, Pelham & Kondla, 2022

LaBar, Caitlin C., Pelham, Jonathan P. & Kondla, Norbert G., 2022, 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), The Taxonomic Report of the International Lepidoptera Survey 10 (3), pp. 1-25 : 5-23

Celastrina asheri LaBar, Pelham & Kondla , new species

(Asher’s Blue)

( Figs. 10-41 )

ZooBank registration: urn:lsid:zoobank.org:act:6E70FCCC-505A-4AA4-80C2-4D52C63A8274

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. VFW 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 VFW 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 CuA1 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, 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.

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). Twentyfour male and 1 female paratypes, same data as holotype. Five additional paratypes from same locality: 2021-06-12, C. C. LaBar (1♂ and 1♀, ex larvae) ; 2021-06-21, C. C. LaBar (2♂, wild-caught), 2022- 03-15 (1♂, ex larva). Three paratypes from McCabe Pond fishing access area southeast of Ringer South Trail , Kittitas Co., WA, 2021-04-21 (2♂) and 2021-04-22 (1♀), 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 (5♂, NGK) ; South Castle River Valley, vic. N49.3081 W114.2938 elev. 1400-1460m, 1988-05-16, N. G. Kondla (1♂, NGK) ; same locality, 1982-05-24, N. G. Kondla (1♂, NGK) ; same locality, 1983-05-22, N. G. Kondla (1♂, NGK) ; BRITISH COLUMBIA: Beaver Creek at Columbia River , vic. N49.0658 W117.6103 elev. 410-450m, 1999-04- 11, N. G. Kondla (1♂, NGK) ; same locality, 2006-04-24, N. G. Kondla (1♂, NGK) ; Blackwater Rd near Clearwater, N51.573 W120.149 elev. 408m, 1994-05-22, N. G. Kondla (1♀, NGK) ; Brilliant, vic. N49.3177 W117.6439 elev. 420-430m, 2002-04-18, N. G. Kondla (1♂, NGK) ; same locality, 2002-04- 19, N. G. Kondla (3♂, NGK) ; same locality, 2002-05-01, N. G. Kondla (7♂, NGK) ; same locality, 2002-05-10, N. G. Kondla (4♂, 1♀, NGK) ; same locality, 2002-05-13, N. G. Kondla (1♂, NGK) ; same locality, 2002-06-03, N. G. Kondla (1♀, NGK) ; Charbonneau Creek, vic. N49.0470 W117.4753 elev. 560-870m, 1998-04-26, N. G. Kondla (5♂, NGK) ; same locality, 1998-04-29, N. G. Kondla (3♂, NGK) ; same locality, 2000-04-08, N. G. Kondla (1♂, NGK) ; same locality, 2000-05-19, N. G. Kondla (1♂, NGK) ; same locality, 2002-04-27, N. G. Kondla (12♂, NGK) ; same locality, 2002-05-11, N. G. Kondla (5♂, NGK) ; same locality, 2002-05-12, N. G. Kondla (27♂, 1♀, NGK) ; same locality, 2002- 05-19, N. G. Kondla (20♂, 1♀, NGK) ; same locality, 2002-05-31, N. G. Kondla (6♂, NGK) ; same locality, 2003-04-20, N. G. Kondla (3♂, NGK) ; same locality, 2003-04-21, N. G. Kondla (1♂, NGK) ; same locality, 2003-04-28, N. G. Kondla (112♂, NGK) ; same locality, 2004-04-25, N. G. Kondla (1♂, NGK) ; same locality, 2004-04-26, N. G. Kondla (35♂, NGK) ; same locality, 2006-04-30, N. G. Kondla (1♂, NGK) ; Ellison Ridge, vic. N50.1264 W119.4129 elev. 460-1040m, 2001-04-26, D. L. Threatful (7♂, 2♀, 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 (1♂, NGK) ; km 2-3 Handley Road, Pend-d’Oreille, vic. N49.0270 W117.5348 elev. 580-610m, 2002-06-02, N. G. Kondla (3♂, NGK) ; Hulme Creek Rd, N49.0885 W119.0237 elev. 1030m, 2009-06-01 or 03, D. L. Threatful (1♂, NGK) ; Kettle Valley Rail Trail near Carmi, vic. N49.4896 W119.1187 elev. 820-840m, 2009-05-22 to 24, D. L. Threatful (2♂, 5♀, NGK) ; Kid Creek Forest Service Road at Hwy 3, vic. N49.1626 W116.2853 elev. 760-770m, 2003- 05-24, N. G. Kondla (2♂, 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 (1♂, NGK) ; same locality, 2003-04-30, N. G. Kondla (2♂, NGK) ; same locality, 2006-04-27, N. G. Kondla (1♀, NGK) ; 1 km south of Ootischenia, vic. N49.2690 W117.6240 elev. 550-570m, 2002-05-16, N. G. Kondla (1♂, NGK) ; Rock Mountain Forest Service Road near Rock Creek, N49.0336 W119.053 elev. 970m, 2001-05-06, N. G. Kondla (1♂, 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 (4♂, NGK) ; same locality, 2002-05-12, N. G. Kondla (33♂, NGK) ; same locality, 2002-06-23, N. G. Kondla (2♂, NGK) ; km 2.7 west of Seven Mile Dam, N49.0221 W117.5338 elev. 480m, 2002-04-27, N. G. Kondla (18♂, NGK) ; same locality, 2002-05-11, N. G. Kondla (57♂, 2♀, NGK) ; same locality, 2002-05-12, N. G. Kondla (18♂, NGK) ; same locality, 2002-05-19, N. G. Kondla (5♂, NGK) ; same locality, 2002-05-31, N. G. Kondla (41♂, NGK) ; same locality, 2002-06-02, N. G. Kondla (20♂, 1♀, NGK) ; same locality, 2002-06-12, N. G. Kondla (7♂, NGK) ; same locality, 2003-04-20, N. G. Kondla (16♂, NGK) ; same locality, 2003-04-21, N. G. Kondla (35♂, NGK) ; same locality, 2003-04-28, N. G. Kondla (26♂, NGK) ; same locality, 2003-05-01, N. G. Kondla (19♂, NGK) ; same locality, 2004-04- 09, N. G. Kondla (3♂, NGK) ; same locality, 2004-04-25, N. G. Kondla (8♂, NGK) ; same locality, 2004- 04-26, N. G. Kondla (6♂, NGK) ; same locality, 2005-04-22, N. G. Kondla (3♂, NGK) ; same locality, 2006-04-20, N. G. Kondla (1♂, NGK) ; same locality, 2006-05-04, N. G. Kondla (1♂, 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 (9♂, NGK) ; same locality, 2003-05-23, N. G. Kondla (2♂, NGK) ; same locality, 2006-05-10, N. G. Kondla (1♂, NGK) ; km 3 east of Seven Mile Dam, N49.0446 W117.4823 elev. 585m, 2003-05-10, N. G. Kondla (1♀, NGK) ; km 3.5 east of Seven Mile Dam, N49.0469 W117.4762 elev. 600m, 2002-05-11, N. G. Kondla (1♂, NGK) ; same locality, 2002-05-19, N. G. Kondla (1♂, NGK) ; same locality, 2002- 05-31, N. G. Kondla (5♂, NGK) ; same locality, 2002-06-02, N. G. Kondla (3♂, NGK) ; same locality, 2002-06-12, N. G. Kondla (1♂, NGK) ; same locality, 2003-05-01, N. G. Kondla (3♂, NGK) ; same locality, 2003-05-28, N. G. Kondla (1♂, NGK) ; same locality, 2003-05-29, N. G. Kondla (1♂, NGK) ; km 5.1 east of Seven Mile Dam, N49.0486 W117.4596 elev. 585m, 1999-04-24, N. G. Kondla (1♂, NGK) ; same locality, 2002-04-20, N. G. Kondla (3♂, NGK) ; same locality, 2002-04-27, N. G. Kondla (6♂, NGK) ; same locality, 2002-05-11, N. G. Kondla (33♂, 1♀, NGK) ; same locality, 2002-05-12, N. G. Kondla (6♂, NGK) ; same locality, 2002-05-19, N. G. Kondla (6♂, NGK) ; same locality, 2002-05- 31, N. G. Kondla (12♂, NGK) ; same locality, 2002-06-02, N. G. Kondla (1♂, NGK) ; same locality, 2002-06-21, N. G. Kondla (2♂, NGK) ; same locality, 2003-04-21, N. G. Kondla (1♂, NGK) ; same locality, 2006-04-24, N. G. Kondla (2♂, NGK) ; same locality, 2004-03-29, N. G. Kondla (1♂, NGK) ; same locality, 2006-04-20, N. G. Kondla (3♂, 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 (2♂, NGK) ; km 6 east of Seven Mile Dam, N49.0483 W117.4483 elev. 590m, 2002-05-31, N. G. Kondla (14♂, 1♀, NGK) ; same locality, 2005-04-22, N. G. Kondla (1♀, NGK) ; km 6 to 7 east of Seven Mile Dam, vic. N49.0490 W117.4398 elev. 580-610m, 2002-05-12, N. G. Kondla (17♂, 2♀, NGK) ; km 7 east of Seven Mile Dam, N49.0490 W117.4347 elev. 580m, 2002-04-20, N. G. Kondla (2♂, NGK) ; same locality, 2002-04-27, N. G. Kondla (7♂, 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 (5♂, 3♀, NGK) ; same locality, 2003-04-21, N. G. Kondla (2♂, NGK) ; same locality, 2003-05-01, N. G. Kondla (2♂, NGK) ; same locality, 2003-05-23, N. G. Kondla (1♂, NGK) ; same locality, 2004-04- 25, N. G. Kondla (1♂, NGK) ; same locality, 2005-04-10, N. G. Kondla (1♂, NGK) ; same locality, 2006-04-20, N. G. Kondla (2♂, NGK) ; same locality, 2006-04-30, N. G. Kondla (4♂, NGK) ; Shadey Creek Road, 2005-04-30, N. G. Kondla (1♂, NGK) ; Silver Star Mountain, vic. N50.3884 W119.0941 elev. 1150m, 2002-06-04, D. L. Threatful (1♀, 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 (7♂, NGK) ; same locality, 2007-04-30, N. G. Kondla (18♂, 1♀, NGK) ; same locality, 2006-04-30, N. G. Kondla (8♂, NGK) ; same locality, 2006-05-04, N. G. Kondla (15♂, NGK) ; mountainside above Waneta Dam, vic. N49.007 W117.611 elev. 450-518m, 2006-04-30, N. G. Kondla (1♂, NGK) ; USA: MONTANA: Lincoln Co.: Bad Medicine campground, Bull Lake , N48.22 W115.858 elev 2370ft (722m), 2006-05-19, N. G. Kondla (1♀, NGK) ; Dorr Skeels campground, Bull Lake, N48.268 W115.855 elev 2350ft (716m), 2006-05-19, N. G. Kondla (1♂, NGK) ; Missoula Co.: Miller Creek , vic. N46.765 W113.942 elev. 3730-4000ft (1137-1219m), 1976-05-12, S. Kohler (1♂, 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. Kondla (11♂, 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 (11♂, 4♀, UWBM) ; same locality, 1975-05-21, J. P. Pelham & T. S. Pelham (6♂, 5♀, 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 (5♀, UWBM) ; Reecer Creek Road parking loop, N47.1703 W120.5923 elev. 3400-3500ft (1036-1067m), 1978-07-01, J. P. Pelham & R. Massey (2♂, 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 (1♂, UWBM) ; same locality, 2005-04- 26, C. C. LaBar (1♂, 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♂, 3♀, CCL) ; Naneum Basin/Naneum Road at pullout along creek, N47.17965 W120.44689 elev. 2980ft (908m), 2021-05-29, C. C. LaBar (3♂, 2♀, 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 (670-731m), 1967-05-15, R. Cheyne (1♀, UWBM) ; Wilson Creek (Canyon), [N47.1542 W120.5077] elev. 2800-3000ft (853-914m), 1966-07-15, R. Cheyne (1♂, UWBM) ; Wilson Creek Road at Bar 14 Road, N47.0734 W120.4961 elev. 2050-2060ft (625-628m), 1966-07-18, R. Cheyne (2♀, UWBM) ; Yakima River at US Hwy 90, N47.1858 W121.0432 elev. 2010-2025ft (613-617m), 1958-04-05, D. Carney (1♀, UWBM) ; same locality, 1958-04-11, D. Carney (5♂, UWBM) ; same locality, 1958-04-28, D. Carney (2♂, UWBM) ; same locality, 1958-05-28, D. Carney (1♀, UWBM) ; same locality, 1972-05-28, D. Carney (1♀, UWBM) ; Klickitat Co.: Satus Creek at US Hwy 97, N46.0268 W120.6244 elev. 2139ft (652m), 1955-05-07, G. Schenk (10♂, 4♀, UWBM) ; Okanogan Co.: Chiliwist Rd , N48.3085 W119.8479 elev. 2640ft (805m), 1978-04-28, R. M. Pyle (2♂, 3♀, UWBM) ; Chiwiliken Valley Road, N48.5813 W119.2525 elev. 4415-4425ft (1346- 1349m), 2006-06-17, D. & J. Nunnallee (1♂, UWBM) ; Sinlahekin Wildlife Area, west canyon, N48.68861 W119.74018 elev. 2765ft (843m), 2013-05-11 (3♂, CCL) ; same locality, 2020-07-08, C. C. LaBar (1♂, ex larva collected 2021-06-18, CCL) ; same locality, 2022-03-03, C. C. LaBar (1♂, 1♀, ex larvae collected 2021-06-18, CCL) ; same locality, 2022-03-13, C. C. LaBar (1♂, 1♀, ex larvae collected 2021-06-18, CCL) ; same locality, 2022-03-14, C. C. LaBar (1♀, ex larva collected 2021-06-18, CCL) ; same locality, 2022-03-15, C. C. LaBar (1♂, 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 (1♀, UWBM) ; State Hwy 31 at Threemile Creek, N48.8938 W117.3280 elev. 2550ft (777m), 1982-06-16, R. M. Pyle (1♂, UWBM) ; East Fork Leclerc Creek Rd, N48.561 W117.2724 elev. 2500ft (762m), 1982-06-18, R. M. Pyle (1♀, UWBM) ; Horseshoe Lake Road, N48.09033 W117.4276 elev. 2225ft (678m), 2018-05- 26, C. C. LaBar (1♂, 1♀, CCL) ; Yakima Co.: Bear Canyon , N46.7096 W120.9043 elev. 2000-2500ft (700-762m), 1968-04-09, J. P. Pelham, L. Obata & H. Wiles (1♂, UWBM) ; same locality, 1968-04-28, J. P. Pelham & H. Wiles (1♂, UWBM) ; same locality, 1989-05-06, J. P. Pelham, K. Hiruma, L. Crabo (5♂, 1♀, UWBM) ; Bear Canyon, N46.6997 W120.9383 elev. 2985-3040ft (910-927m), 1973-04-07, J. P. Pelham (1♂, 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♂, 2♀, UWBM) ; Cowiche Creek Canyon, N46.625019 W120.650809 elev. 1430ft (436m), 2003-04-12, J. P. Pelham, Z. A. Pelham, D. Nunnallee & K. Hiruma (6♀, 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. (1♂, UWBM) ; Oak Creek, N46.7303 W120.8395 elev. 1900-2100ft (579-640m), 1995-05-07, J. F. Rial (1♂, 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 (1♀, UWBM) .

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 POLR 3B-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.

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 midJune 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 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 1 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 Celastrina 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 ). 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 1 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. lucia ” 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 ).

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. lucia and even fewer images are available. Schmidt & Layberry (2016) illustrate a range of variation in C. lucia 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.

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. 32 a-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. 36 a-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. 81 A, 82 A ) in contrast to C. echo females in which the dorsal cell bar is almost always visible on both fore and hindwings ( Figs. 87 A, 88 A ) and C. lucia 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 , 82 B ) similar to female C. lucia ( 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 , 81 B , 97a ). The area between the ventral marginal spots and submarginal crescents of both sexes usually contains varying amounts of dark gray scales ( Figs. 85 D ) compared to C. echo in which this band never contains dark gray scales ( Figs. 91 D, 92 D ). 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. 81-83 C ) to solid gray or black ( Fig. 84 C ), 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. 87-90 C ), 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. 86 E ), the appearance of fuzzy, white postmedian arrowheads on some C. asheri ( Fig. 85 F ) or the frequent duskiness in the VHW marginal band ( Figs. 28b , 39b, 40b , 85 D , 94 ), compared to the pale ventral and thin VHW discal cell bar on C. echo ( Figs. 91 E, 92 E , 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. 91 G, 92 G ), C. asheri may have a very small amount of greenish-blue (less often) or dark gray (more often) scales in this area ( Figs. 10 b-40b, 85 G, 86 G, 96b, 97b), while C. lucia 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. lucia 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 .

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_i. htm and http://butterfliesofamerica.com/ L /ih/celastrina0003_i.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 .

Figures

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.

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.

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.

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

Fig. 42: Ringer South Trail, type locality of Celastrina asheri sp. n.

Fig. 43: Asher, our butterfly’s namesake.

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.

Fig. 45: Flight period of Celastrina asheri sp. n. compared with C. lucia and C. echo from within the same geographical area east of the Cascade crest. Records are grouped into weeks.

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.

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) eggs 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.

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.

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.

Fig. 78: Celastrina asheri sp. n. puddle party in the Pend- d’Oreille River valley, BC, 2003-04-28. Photo by N. G. Kondla.

Fig. 79: Left to right: Celastrina asheri sp. n., C. echo, asheri, echo in the Pend-d’Oreille River valley, BC, 2007-04-23. Photo by N. G. Kondla.

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.

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

Fig. 94: A sample of Celastrina asheri sp. n. from the Pend-d’Oreille River valley, BC. Photos by N. G. Kondla.

Fig. 96: Comparison of Celastrina asheri sp. n. (rows a & b) and C. lucia (rows c & d) males. Photos by N. G. Kondla.

Figs. 81-92: Comparison of Celastrina 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.

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.

Fig. 95: Celastrina lucia near Nazko, BC, 2004-05-09. Photo by N. G. Kondla.