Soyedina carolinensis ( Claassen, 1923 )

Grubbs, Scott A. & Baumann, Richard W., 2019, Soyedina Ricker, 1952 (Plecoptera: Nemouridae) in the eastern Nearctic: review of species concepts, proposed morphology-based species groups, and description of a new species from North Carolina, Zootaxa 4658 (2), pp. 223-250 : 231-233

Soyedina carolinensis ( Claassen, 1923)

Carolina Forestfly

http://lsid.speciesfile.org/urn:lsid: Plecoptera .speciesfile.org:TaxonName:606 5

( Figs. 25–34 , 80 )

Nemoura carolinensis Claassen 1923:287 . Holotype ♂ (MCZC), Morganton , (Burke Co.), North Carolina, USA

Nemoura carolinensis Needham & Claassen, 1925:215

Nemoura (Soyedina) washingtoni carolinensis Ricker, 1952:53

Soyedina carolinensis Illies, 1966:249

Soyedina carolinensis Stark, 2017:187

Distribution. USA: DE, MD, NC, PA, TN, VA, WV ( DeWalt et al. 2019).

Material examined. USA, Maryland, Frederick Co., spring seep Raven Rock Run , 17 March 1990, R.M. Duffield, 1 pharate male, 10 larvae ( CHNC) ; Fishing Creek , 15 March 1991, R.M. Duffield, 1 male, 1 female ( CHNC) ; tributary to Steep Creek , 39.56006, -77.48271, 9 March 1997, S.A. Grubbs, 1 male, 2 females ( WKUC) . North Carolina, Buncombe Co., stream, Tanbark ( Ridge ) Tunnel , Blue Ridge Parkway , 5 March 1991, R.W. Baumann & S.M. Clark, 2 males ( BYU) ; spring into Ox Creek, Blue Ridge Parkway , 35.67078, -82.47066, 18 April 2009, S.A. Grubbs, 1 male ( WKUC) ; Graham Co., spring seeps, FS 81, Nantahala National Forest , 35.34784, -83.99531, 1 March 2008, A.L. Sheldon, 1 male ( WKUC) ; seep into Sand Creek, FS 81F, Nantahala National Forest , 35.36208, -83.99576, 2 March 2008, A.L. Sheldon, 1 male ( WKUC) ; Haywood Co., Haywood Co., GRSM ATBI Plot , Purchase Knob , (malaise trap), 35.59004, -83.06024, 7–27 March 2002, P.F. Super & J. Lowe, 3 males, 9 females ( BYU) ; Big East Fork, Pigeon River , 14 March 2006, J. Robinson, 1 male, 2 females ( BYU) ; tributary to Laurel Creek, Pisgah National Forest , 35.75263, -83.09060, 1 March 2008, S.A. Grubbs, 7 males, 3 females ( WKUC) ; tributary to Laurel Creek, Pisgah National Forest , 35.75255, -83.08755, 1 March 2008, S.A. Grubbs, 4 males, 4 females ( WKUC) ; tributary to Flat Branch, Pisgah National Forest , 35.74612, -83.07928, 1 March 2008, S.A. Grubbs, 1 male, 1 female ( WKUC) ; Cold Springs Creek, FR 148, Pisgah National Forest , 35.77850, -82.95590, 2 March 2008, 1 male ( WKUC) ; same but 18 April 2009, S.A. Grubbs, 1 male, 1 female ( WKUC) ; tributary to Hemphill Creek , 35.57690, -83.07230, 2 March 2008, S.A. Grubbs, 14 males, 6 females ( WKUC) ; Jackson Co., 3 mi. SW Cashiers, Whiteside Mountain , 9 May 1997, C.R. Nelson, 1 male ( BYU) ; Macon Co., Rattlesnake Springs, Nantahala National Forest , 35.17060, -83.57741, 18 March 1982, J. Weaver, R. Holzenthal, 2 males ( BPSC) ; same but 1–18 May 1983, J. Weaver & R. Holzenthal, 31 males, 93 females ( BYU) ; same but 23 March 2002, S.A. Grubbs, 6 males, 8 females ( WKUC) ; tributary to Jarrett Creek, Nantahala National Forest , 35.15903, -83.62873, 23 March 2002, S.A. Grubbs, 1 male, 5 females ( WKUC) ; same but 8 March 2004, S.A. Grubbs, 1 male, 1 female ( WKUC) ; Robin Branch, Nantahala National Forest , 35.16832, -83.58465, B.P. Stark, 19 May 1983, 1 male ( BYU) ; same but K.W. Stewart , 1 March 1986, 1 male, 1 female ( BYU) ; same but 28 February 2008, A.L. Sheldon, 1 male, 1 female ( WKUC) ; same but 15 March 2009, B.P. Stark, 3 males, 3 pharate males, 1 female ( BPSC) ; upper Ball Creek, Coweeta Hydrologic Laboratory , 12 March 1983, A.D. Huryn, 1 male ( ADHC) ; same but 7–20 April 1984, A.D. Huryn, 1 male ( BYU 1824 ) ; Kit Springs Branch, FS 711, Nantahala National Forest , 35.20191, -83.58354, 29 February 2008, A.L. Sheldon, 1 male ( WKUC) ; Sassafras Branch, FS 711, Nantahala National Forest , 35.20230, -83.59406, 29 February 2008, A.L. Sheldon, 1 male ( WKUC) ; seep into Split Whiteoak Branch , FS 711, 35.20458, - 83.58046, 29 February 2008, A.L. Sheldon, 1 male ( WKUC) ; tributary to Split Whiteoak Branch, FS 711, Nantahala National Forest , 35.20640, -83.59059, 29 February 2008, A.L. Sheldon, 3 males, 2 females ( WKUC) ; Blue Valley Overlook, Hwy 106, 12 March 1995, B.P. Stark, R. Simmons & R. Frederick, 3 males ( BYU) ; same but 14 March 2006, B.P. Stark, 4 males ( BPSC) ; Swain Co., tributary to Connelly Creek, Nantahala National Forest , 35.35226, -83.35175, 23 March 2002, S.A. Grubbs, 1 male, 5 females ( WKUC) ; Yancey Co., Big Crabtree Creek, FR 5511, Pisgah National Forest , 35.79264, -82.17388, 24 March 2002, S.A. Grubbs, 2 males, 7 females ( WKUC) . Pennsylvania, Chester Co., East Branch, White Clay Creek , 0.6 mi. W London Grove, 24 February 1980, D.H. Funk, 2 males, 1 female ( BYU) ; same but March 6 1980, D.H. Funk, 7 males, 2 females ( BYU) ; tributary, White Clay Creek, Stroud Laboratory , 26 April 1964, J.K. Gelhaus & D P. Koenig, 11 males, 22 females ( BYU) ; [Philadelphia Co.], Philadelphia , 24 April 1910, G.M. Greene, 1 male (removed from pin) ( USNM) . Tennessee, Carter Co., Twin Springs, Cherokee National Forest , 36.12128, -82.08654, 10 April 2002, S.A. Grubbs, 1 male, 1 female ( WKUC) ; Sevier Co., Middle Prong Little Pigeon River, Great Smoky Mountains National Park , 22–23 April 1978, C.H. Nelson, 2 males, 3 females ( CHNC) ; Ramsay Prong Little Pigeon River, Great Smoky Mountains National Park , 22–23 April 1978, C.H. Nelson, 2 males, 2 females ( CHNC) ; Gatlinburg, Great Smoky Mountains National Park , 28 April 1978, F.F. Sherberger, 1 male, 1 female ( BYU) ; Cades Cove, 3.5 mi. E Cabin Mill, Great Smoky Mountains National Park , 25 March 1980, B.P. Stark, K.W. Stewart & J. Feminella, 4 males, 4 females ( BYU) ; Rocky Spur Branch, Great Smoky Mountains National Park , 35.68027, -83.46450, 25 March 1999, B.C. Kondratieff, 3 males, 3 females ( WKUC) ; Road Prong, Great Smoky Mountains National Park , 35.60975, -83.44918, 15 April 2014, S.A. Grubbs, 1 male, 1 female ( WKUC) . Virginia, Bedford Co., Hemp Mill Branch, Sheep Creek , 37.40222, -79.65056, 25 February 1980, C.F. Burgoon & C.E. Dunn, 2 males ( BYU) ; Floyd Co., Tory Creek, Hwy 58, west of Meadows of Dan , 36.72798, -80.47042, 11 March 1991, R.W. Baumann & R.F. Kirchner, 6 males ( BYU, WKUC) ; Grayson Co., spring on top of Straight Mountain , Rt. 603, 22 April 1979, B.C. Kondratieff, 1 male, 1 female ( BYU 1941 ) ; unnamed spring, Rt. 603, near Lewis Fork Bridge , 18 February 1990, R.F. Kirchner & K.W. Stewart, 8 males, 4 females ( BYU) ; seep into Lewis Fork, Jefferson National Forest , 36.66883, -81.53128, 13 April 2008, A.L. Sheldon, 5 males, 2 females ( WKUC) ; same but 36.67208, -81.52467, 13 April 2008, A.L. Sheldon, 2 males, 2 females ( WKUC) ; same but 36.66790, -81.53016, 13 April 2008, A.L. Sheldon, 2 males ( WKUC) ; Madison Co., Rapidan River, Criglersville , 27 March 1938, B.D. Burks, 2 males, ( INHS Plecoptera 7755) ; Rapidan River, Rapidan Camp, R.M. Duffield , 30 April 1988, 1 male ( CHNC) ; spring into Hogcamp Brook, Dark Hollow Falls Trail, Shenandoah National Park , 38.51867, -78.42349, 15 May 1995, S.A. Grubbs, 1 male, 4 females ( WKUC) ; Washington Co., spring, FR 90, Mount Rogers National Recreation Area , 36.65629, -81.70547, 22 March 2002, S.A. Grubbs, 2 males, 2 females ( WKUC) . West Virginia, Randolph Co., Marsh Fork of Buckhannon River, Route 47 near Pickens , 38.66300, -80.23180, 24 March 1992, S.M. Clark, 13 males, 23 females ( BYU) .

Remarks. The adult male was described and illustrated in Claassen (1923) and subsequently in Needham & Claassen (1925, with the same image of the male in lateral view; their plate 38, fig. 7). Ricker (1952) demoted S. carolinensis as a subspecies of S. washingtoni (both as subgenera of Nemoura ), noting that he found the two taxa as nearly indistinguishable. Full species status, however, was re-elevated by Illies (1966). The combination of the asymmetric dorsal ( Figs. 25–27, 30 ) and ventral sclerites ( Figs. 25–26 ), plus the inner epiproct member that is flanged and directed laterally with the flange lip approximately same width apically ( Figs. 25–27 ), aligns this species closely with S. merritti and S. washingtoni . We view S. carolinensis as a valid, albeit variable, species that is distinct from S. washingtoni . Although the outer paraproct lobe of S. carolinensis is variable in form (see below), for S. washingtoni it is consistently quadrate or subquadrate across its range ( Figs. 49–50 ).

The outer paraproct lobe of S. carolinensis varies in form. For example, early-emergent populations (i.e. late winter–early spring) from east Tennessee and western North Carolina have an outer lobe that has a narrow profile in apical ½ with anterior and posterior margins ca. parallel and without a basoposterior expansion ( Fig. 31 ). This is a very common phenotype. A slight variation in this theme is present on populations from central West Virginia ( Fig. 32 ). In contrast, populations from higher-elevation streams and seeps in southwestern Virginia and east Tennessee have an outer paraproct lobe that is more sickle-shaped and with a distinct anteriorly-directed spur compared to other phenotypes ( Fig. 33 ). Populations further northward along the Blue Ridge and upper Piedmont provinces into central Maryland and southeastern Pennsylvania have an outer lobe that is different along the anterior and posterior margins, and has a more bulbous basoposterior portion ( Fig. 34 ). Because of this, we contend that morphology alone cannot effectively address the question regarding of if there are evolutionarily independent units of S. carolinensis that represent undescribed species. Although the larva is undescribed, Stark (2017) noted one cercal feature that is distinct from that of S. vallicularia .

Soyedina carolinensis is known from the southern Appalachian Mountain region northeastward to the Piedmont Uplands of southeastern Pennsylvania and northern Delaware ( Fig. 80 ).

Figures

FIGURE 80. Distribution maps of eastern Nearctic Soyedina based on material examined during this study plus valid literature data. 80, S. carolinensis, S. merritti, and S. washingtoni.

FIGURES 25–30. Soyedina carolinensis. 25: USA, Virginia, seep into Lewis Fork; 26, 28: USA, West Virginia, Marsh Fork of Buckhannon River; 27, 30: USA, North Carolina, Rattlesnake Springs; 29: USA, North Carolina, Robin Branch; 25, epiproct, ventral view, 170X; 26, epiproct, ventral view, 370X; 27, epiproct apex, ventral view, 650X; 28, epiproct apex, dorsal view, 1200X; 29, epiproct, lateral view, 285X; 30, epiproct, caudal view, 230X. ds = dorsal sclerite, vs = ventral sclerite, im = inner member.

FIGURES 31–34. Soyedina carolinensis. 31: USA, North Carolina, Rattlesnake Springs, outer paraproct lobe, lateral view, 250X; 32: USA, West Virginia, Marsh Fork of Buckhannon River, outer paraproct lobe, lateral view, 190X; 33: USA, Virginia, seep into Lewis Fork, outer paraproct lobe, lateral view, 230X; 34, USA, Maryland, Fishing Creek, outer paraproct lobe, lateral view, 270X.

FIGURES 43–50. Soyedina washingtoni. 43–46, 49: USA, Maine, Thorndike; 47: USA, Maryland, spring into Elk Lick Run; 48: USA, Pennsylvania, Fourmile Creek; 50: USA, Maryland, Mill Run. 43, epiproct, ventral view, 105X; 43, epiproct, ventral view, 247X; 45, epiproct apex, ventral view, 637X; 46, epiproct apex, dorsal view, 488X; 47, epiproct, lateral view, 370X; 48, epiproct, caudal view, 250X; 49, outer paraproct lobe, lateral view, 166X; 50, outer paraproct lobe, lateral view, 300X; ds = dorsal sclerite, vs = ventral sclerite, im = inner member.