Moselia zonata Stark & Harrison, 2016

Stark, Bill P. & Harrison, Audrey B., 2016, The Banded-Wing Moselia (Plecoptera: Leuctridae) Revisited, Illiesia 12 (9), pp. 42-58 : 44-51

Moselia zonata Stark & Harrison , sp. n.

Banded-wing Needlefly

( Figs. 2 , 3-8 , 9-40, 42 , 52 )

http://lsid.speciesfile.org/urn:lsid: Plecoptera .speciesfile.org: TaxonName:492220

Moselia infuscata: Gill et al., 2015: 593 (in part), “Banded- wing phenotype”, population 1

Material examined. Holotype ♂, OREGON, Jackson Co., Sheep Creek, Wagner Gap Road, NFD 22 Rd , 10.5 miles SW Talent, 42° 06.442’N, 122° 47.921’W, 22 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C.J. Verdone ( USNM) . Paratypes: CALIFORNIA: Butte Co., Butte Creek, Humboldt Rd , above Colby Creek , 40.11138° N, 121.49526° W, 25 May 2014, B.C. Kondratieff, C.J. Verdone, 15♂, 9♀ ( CSUC), 1♂ SEM . Del Norte Co., small falls into Patrick Creek, 41° 54.321’ N, 123° 51.357’ W, 24 May 2014, B. Stark, A.B. Harrison, 1♀ ( BPSC). Humboldt Co. , tributary of Willow Creek , 2.7 miles E Berry Summit, Hwy 299, 19 May 1998, S.W. Szczytko, B. Stark, C. R . Nelson, I. Sivec, 5♂, 14♀ ( BYUC). 2♀ SEM . tributary of Willow Creek , 2.5 miles E of Berry Summit, 19 May 1998, B. Stark, I. Sivec, C. R . Nelson, S.W. Szczytko, 2♂, 1♀ ( BPSC), 4♂, 5♀ ( BYUC). tributary of Willow Creek, 0.5 miles E of Cedar Creek, Hwy 299, 25 April 1987, R . W. Baumann, B. Stark, C. R . Nelson, S. Wells, 8♂, 5♀ ( BPSC), 15♂, 15♀ ( BYUC), 3♂, 1♀ SEM . tributary of Ruby Creek , first stream E of highway crossing, 22 April 1993, G. R . Fiala, 41♂, 25♀ ( BYUC), 1♂ SEM . small creek N of Fish Lake , 24 April 1987, R . W. Baumann, B. Stark, C. R . Nelson, S. Wells, 15♂, 10♀ ( BYUC), 1♂ SEM . small stream, 0.5 miles E Cedar Creek, Hwy 299, 25 April 1987, B. Stark, R . W. Baumann, C. R . Nelson, S. Wells, 8♂, 5♀ ( BPSC), 2♂ SEM . Prairie Creek, Prairie Creek Redwoods State Park, Drury Parkway at trailhead, 41° 24.459’ N, 124° 01.827’ W, 25 May 2014, B. Stark, A.B. Harrison, 1♂ ( BPSC), 1♂ SEM . tributary Prairie Creek , 10.6 miles N Orick, 22 May 1982, B. Stark, D. Ziegler, 1♂ ( BPSC). Siskiyou Co., small stream, 41.90335° N, 123.90345° W, 19 April 2013, C. Kerst, 3♂, 1♀ ( CSUC), 1♂ SEM . Tehama Co., South Fork Calf Creek, Hwy 32 at FR 27N06, 26 April 2010, B.C. Kondratieff, R . W. Baumann, 8♂, 2♀ ( CSUC) 1♂ SEM . Trinity Co., Bidden Creek , W of Cedar Flat, Hwy 299, 25 April 1987, R . W. Baumann, B. Stark, C. R . Nelson, S. Wells, 4♂, 8♀ ( BYUC), 1♂ SEM . unknown stream, Hwy 36, Mile 12.12, South Fork Mountain , approximate co-ordinates 40.39366° N, 123.41517° W, 2 May 2014, J.J. Lee, 5♂, 6♀ ( JJLC), 1♂, 1♀ SEM . OREGON: Curry Co., Fly Catcher Spring, 12 miles E Hwy 101, Hunter Creek Rd , 42° 22’ N, 124° 18’ W, 9 June 2005, R .W. Baumann, B.C.

Kondratieff , 10♂, 13♀ ( BYUC), 2♂ SEM . Jackson Co., Ashland Ski Road seeps, 2.5 miles W Interstate Highway 5, 9 June 2004, B. Stark, R . W. Baumann , 8♂, 17♀ ( BPSC), 1♂, 1♀ SEM . Jim Creek , Wagner Gap Road, NFD 22 Rd, 11.1 miles SW Talent, 42.10609 N, 122.7874 W, 22 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C.J. Verdone, 1♂, 1♀ ( BPSC), 4♂, 2♀ ( CSUC), 1♂ SEM . Sheep Creek , Wagner Gap Road, NFD 22 Rd, 10.5 miles SW Talent, 42.11254 N, 122.7858 W, 22 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C.J. Verdone, 4♂, 2♀ ( BPSC), 4♀ ( CSUC), 1♂ SEM . Split Rock Creek , Wagner Creek Road, 12.4 miles SW Talent, 42.17018 N, 122.784174 W, 22 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C.J. Verdone, 2♂, 1♀ ( BPSC), 6♀ ( CSUC), 1♂ SEM . Wagner Creek , Wagner Gap Road, 5.9 miles SW Talent, 42.17018 N, 122.784174 W, 22 May 2014, B.C. Kondratieff, B. Stark, J.C. Sandberg, C.J. Verdone, 9♂, 5♀ ( CSUC) . Josephine Co., Lake Creek and seeps along NF-070, 42.09274 N, 123.37444 W, 21 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C.J. Verdone, 1♂ ( BPSC) .

Additional specimens: CALIFORNIA: Alpine Co., Willow Creek, 6 miles NW Woodfords, 38.827053 N, 119.9033331 W, 27 July 1995, R.L. Bottorff, 1♂ (RLBC) , 1♂ SEM. Butte Co., spring tributary Big Chico Creek, Hwy 32, 40° 08’ N, 121° 40’ W, 26 April 2010, R.W. Baumann, B.C. Kondratieff, 12♂, 7♀ ( BYUC) , 2♂ SEM. El Dorado Co., Bendorf Spring, NE Grizzly Flat, 3 May 1995, R.L. Bottorff, 7♂, 2♀ (RLBC) , 1♂ SEM. tributary North Cosumnes River, 11 miles E Grizzly Flat, 38.64018 N, 120.31283 W, 30 June 1986, R.L. Bottorff, 1♂ (RLBC) , 1♂ SEM. North Fork Cosumnes River, Sciaroni Crossing, 3 miles N Grizzly Flat, 38.669092 N, 120.533568 W, 20 April 1981, R.L. Bottorff, 1 ♂ (RLBC) . Glenn Co., spring fed creek, 3 miles above Salt Creek, 15 March 1984, J. Stanger, 3♂, 1♀ ( BYUC) , 2♂ SEM. Mendocino Co., Caspar Creek, Jackson Demonstration State Forest, 18 May 1998, B. Stark, C. R. Nelson, S.W. Szczytko, I. Sivec, 1♂, 1♀ ( BPSC) , damaged paraprocts. Irish Gulch, 1 mile above Hwy 1, 39° 01.37’ N, 123° 59.57’ W, 7 April 2006, L.D. Bottorff, 1♂, 3♀ (RLBC) , 1♂ SEM. Mono Co., west source spring on E North Canyon Creek, ca. 7.6 miles ESE Mono Mills, 37.8591 N, 118.8216 W, 1 August 1998, M. Myers, 1♂ ( BYUC) . Nevada Co., spring on Sagehen Creek, Sagehen Creek Biological Station, 39.43° N, 120.25° W, 20 June 2009, R.W. Baumann, B.C. Kondratieff, 9♂, 13♀, 2♂ SEM. Plumas Co., Domingo Springs, Feather River Rd/Old Red Bluff Rd, 40° 21.672’ N, 121° 20.842’ W, 26 May 2014, B. Stark, A.B. Harrison, 14♂, 6♀ ( BPSC) , 1♂ SEM. same site, 8 June 2004, B. Stark, R.W. Baumann, 8♂, 16♀ ( BPSC) . same site, 25 May 2007, B.C. Kondratieff, R.W. Baumann, 12♂, 13♀ ( CSUC) . Shasta Co., East Fork Hat Creek, Hwy 89, Lassen Volcanic National Park, 8 June 2004, B. Stark, R.W. Baumann, 1♂ ( BPSC) , 1♂ SEM. Old Boundary Spring, Lassen Volcanic National Park, 8 June 2004, B. Stark, R.W. Baumann, 31♂, 8♀ ( BPSC) , 1♂ SEM. Siskiyou Co., Big Springs, Mt. Shasta City Park, 41° 19.692’ N, 122° 19.605’ W, 8 June 2004, B. Stark, R.W. Baumann, 2♂, 2♀ ( BPSC) , 1♂ SEM. Black Butte Springs, 41° 23.667’ N, 122° 21.626’ W, 23 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C.J. Verdone, 3♂, 1♀ ( BPSC) , 1♂ SEM. Tehama Co., Bluff Falls, Hwy 89, 8 June 2004, B. Stark, R.W. Baumann, 1♂ ( BPSC) , 1♂ SEM, damaged paraprocts. Trinity Co., East Fork Trinity Creek at campground, 40.77229° N, 122.92150° W, 17 April 2014, C. Kerst, 2♂, 2♀ ( CSUC) . NEVADA: Douglas Co., Corsser Creek, Hwy 207, Kingsbury Glade, 38.94806 N, 119.85004 W, 16 May 2013, R.L. Bottorff, 2♂ (RLBC) , 2♂ SEM. Edgewood Creek, Stateline, 38.96315 N, 119.92562 W, 6 June 2014, R.L. Bottorff, 1♂ (RLBC) . Elko Co., Jarbidge Mountains, springbrook, 10 July 1999, A.L. Sheldon, 5♂, 7♀ ( BYUC) , 1♂ SEM. Humboldt Co., Road Creek, E side Santa Rosa Range, 13 miles N Paradise Valley, 41.67881 N, 117.33959 W, 13 June 2013, R.L. Bottorff, 2♂ (RLBC) , 2♂ SEM. Left tributary, Rebel Creek, Santa Rosa Range, 28 June 1999, A.L. Sheldon, 18♂, 9♀ ( BYUC) , 2 ♂ SEM. Singas Creek, Singas Creek Campground, W Paradise Valley, Santa Rosa Range, 41° 41’ N, 117° 64’ W, 15 June 2009, R.W. Baumann, B.C. Kondratieff, 12♂, 8♀ ( BYUC) , 2♂ SEM. Summer Camp Creek, Summit Lake Mountain, 1 July 1999, A.L. Sheldon, 3♂, 4♀ ( BYUC) , 2♂ SEM. Washoe Co., small stream from Marlette Lake, Hwy 28, 24 June 2009, B. Stark, C. R. Nelson, A.B. Harrison, K.C. Nye, 3♂, 8♀ ( BPSC) , 1♂ SEM. Ophir Creek, Mt. Rose Hwy, Tahoe Meadows, 39.30118 N, 119.92132 W, 29 July 2013, L.D. Bottorff, R.L. Bottorff, 1♂ (RLBC) , 1♂ SEM. Rock Creek, Granite Range, 2 July 1999, A.L. Sheldon, 5♂, 8♀ ( BYUC) , 2♂ SEM. Whites Creek, Whites Creek Trailhead, Rd 047, 39.38° N, 119.85° W, 16 June 2009, R.W. Baumann, B.C. Kondratieff, 14♂, 5♀ ( BYUC) , 2♂ SEM. OREGON: Jackson Co., McDonald Creek, NFD 22 Rd, 22 May 2014, B.C. Kondratieff, C.J. Verdone, 1♂ ( CSUC) .

Adult habitus. Male forewing length 5.0- 5.5 mm (n = 17), female 6.5 mm (n = 9). General body color dark brown, dorsum of head brown except for a transverse pale band extending behind ocelli between compound eyes. Pronotal disc mostly pale but with dark brown sutures; anterior and posterior sutures extend about 2/3 of pronotal width; on either side of median suture an irregular pale-brown, gnome, or somewhat C-shaped rugosity occurs ( Figs. 3, 6 ). Wings brown, but with conspicuous pale humeral markings, and many specimens with an irregular broad pale band extending across wings from stigma to stigma ( Fig. 2 ). Tibiae mostly pale along posterior margin but with dark bands at the distal and proximal margins.

Male genitalia (n=41). Typical of the genus in most respects, but paraprocts are relatively short and the fin is relatively tall ( Figs. 4, 7 , 9-32 , 39-40, 42 ). Paraproct body extending from base to fin apex 335- 395 μm; greatest paraproct height across fin apex 120-148 μm; posterior margin of fin and much of paraproct body pale, anterior and basal margin of paraproct body sclerotized, dark brown. Apical digit of the paraprocts 125-142 μm in length.

Female subgenital plate (n = 6). Median lobe parabolic, partially projecting over tips of heavily sclerotized, blade-like, smooth lateral lobes. Sternum 8 with dark pigment interrupted on midline; sternum 9 sclerotized around lateral and posterior margins, but membranous in the area posterior to the subgenital plate ( Figs. 5, 8 , 33-36 ).

Egg. Elongate oval, without collar. Micropyles not detected. Chorionic surface covered with large, hexagonal follicle cell impressions comprised of thin outer walls and a depressed floor bearing clusters of smaller hexagonal units ( Figs. 37-38 ).

Larva. Unknown.

Etymology. The species name refers to the conspicuous, transverse pale band near the apex of the wings in many specimens. The proposed common name is “Banded- wing Needlefly” ( Stark et al., 2012).

Diagnosis. Male and female adult specimens can often be distinguished from those of M. infuscata by the apical pale band across the wings ( Fig. 2 ), however, many specimens apparently lack this marking and it is more conspicuous in fresh specimens. The banded-wing character appears to be restricted to Moselia populations in the Humboldt/Trinity county area of northern California and the Jackson/Josephine/Curry county area of southern Oregon. Other California and Nevada specimens examined in this study did not display the banded-wing character. In addition, males have a broad, subapical, wing-like expansion (= fin) on the anterior paraproct margin ( Figs. 9-32 ), and the paraproct body, extending from the paraproct base to the base of the apical digit, is approximately 335-395 μm (419-500 μm for M. infuscata ). The paraproct fin is smoothly curved along the posterior margin from its junction near the hook through the point of greatest fin height; the greatest height of this structure ranges from approximately 120-148 μm (92-127 μm for M. infuscata ). Fin shape varies among the specimens examined from an almost perpendicular ( Figs. 22, 23, 26 ) to a more slanted profile of the apical fin margin relative to the longitudinal axis of the paraprocts ( Figs. 9-11 , 27-32 ), however, fin shape is relatively uniform at a given location. These data appear to be consistent with the DNA data reported by Gill et al. (2015) for these populations. The distribution of the species encompasses northern California, northern and northwestern Nevada, and extreme southwestern Oregon at the California border ( Fig. 52 ).

Gill et al. (2015) reported 3% mean sequence divergence for the CO1 gene between specimens we determine as this species from Split Rock Creek, Jackson Co., Oregon, and those from Big Springs, Mt. Shasta City Park, Siskiyou Co., California. They reported the same level of differentiation between the Big Springs, Mt. Shasta City Park population and specimens from Butte Co., California. Specimens from these three populations each have relatively high paraproct fins ( Figs. 33-35 ) and relatively short paraprocts. In addition, the mean sequence divergence between these three populations and those from Big Spring, Sierra Co., California that we determined as M. infuscata , is 6% ( Gill et al. 2015). Unfortunately, no genetic data are available for the Oregon and Washington populations of M. infuscata , or for the Nevada populations of M. zonata .

Figures

Figs. 1-2. Adult Moselia habitus. 1. M. infuscata, Oregon, Clackamas Co., Still Creek Campground. 2. M. zonata, Oregon, Jackson Co., Sheep Creek, Wagner Gap Road.

Figs. 3-8. Moselia zonata structures, 3-5, California, Humboldt Co., 0.5 miles E Cedar Creek, Hwy 299. 3. Adult pronotum. 4. Right male paraproct, lateral aspect (d = digit, f = fin, h = hook). 5. Female abdominal sterna 8-9. 6-8, California, Plumas Co., Domingo Springs. 6. Adult pronotum. 7. Right male paraproct, lateral aspect. 8. Female abdominal sterna 8-9.

Figs. 9-14. Moselia zonata right paraproct, lateral aspect. 9. Big Springs, Mt. Shasta, California. 10. Big Springs, Mt. Shasta, Siskiyou Co., California. 11. Black Butte Spring, Siskiyou Co., California. 12. Domingo Springs, Plumas Co., California. 13. Old Boundary Spring, Shasta Co., California. 14. Tributary Big Chico Creek, Butte Co., California.

Figs. 15-20. Moselia zonata right paraproct, lateral aspect. 15. Oregon, Jackson Co., Split Rock Creek, Wagner Creek Rd. 16. Oregon, Jackson Co., Split Rock Creek, Wagner Creek Rd. 17. Oregon, Jackson Co., Jim Creek, Wagner Gap Rd. 18. Oregon, Jackson Co., Ashland Ski Rd seeps. 19. Oregon, Jackson Co., Sheep Creek, Wagner Gap Rd. 20. Oregon, Curry Co., Fly Catcher Spring.

Figs. 21-26. Moselia zonata right paraproct, lateral aspect. 21. California, Humboldt Co., 0.5 miles E Cedar Creek, Hwy 299. 22. California, Humboldt Co., 0.5 miles E Cedar Creek, Hwy 299. 23. California, Humboldt Co., 2.7 miles E Berry Summit, Hwy 299. 24. California, Humboldt Co., Tributary Ruby Creek, Hwy 299. 25. California, Humboldt Co., small creek N Fish Lake. 26. California, Trinity Co., Bidden Creek, Hwy 299.

Figs. 27-32. Moselia zonata right paraproct, lateral aspect. 27. Nevada, Washoe Co., Whites Creek. 28. Nevada, Washoe Co., Whites Creek. 29. Nevada, Washoe Co., Ophir Creek. 30. Nevada, Douglas Co., Corsser Creek. 31. Nevada, Elko Co., Jarbidge Mountains. 32. Nevada, Humboldt Co., Summer Camp Creek.

Figs. 33-38. Moselia zonata female subgenital plate and eggs. 33. Abdominal terminalia, ventral, Ashland Ski Road, Jackson Co., Oregon. 34. Subgenital plate detail, Ashland Ski Road, Jackson Co., Oregon. 35. Subgenital plate detail, 2.7 miles E Berry Summit, Humboldt Co., California. 36. Subgenital plate detail, Domingo Springs, Plumas Co., California. 37. Egg, Domingo Springs, Plumas Co., California. 38. Chorionic detail, Domingo Springs, Plumas Co., California.

Figs. 39-42. Moselia right paraprocts from four populations studied by Gill et al. (2015). The same group designations used by Gill et al. are used here. 39. Group 1, Split Rock Creek, Jackson Co., Oregon, specimens with banded wings. 40. Group 2, Big Springs, Mt. Shasta City Park, Siskiyou Co., California, specimens without banded wings. 41. Group 3, Big Springs, Hwy 49, Sierra Co., California, specimens without banded wings. 42. Group 4, Butte Creek, Butte Co., California, specimens without banded wings. Group 1, group 2 and group 4 specimens each have large fins and a mean genetic distance of 0.03 from the other two groups, and each of these three groups have a mean genetic distance of 0.06 from small fin specimens of group 3. Specimens in groups 1, 2 and 4 are placed as M. zonata sp. n. and those in group 3 are placed as M. infuscata.

Fig. 52. Distribution of Moselia species in the Pacific Northwest. Published records of M. infuscata from British Columbia were not included because no specimens were available for this study.