Sierraperla tolowa Stark & Kondratieff & Sandberg & Gill & Verdone & Harrison 2015, sp. n.

Sierraperla tolowa Stark & Kondratieff, sp. n.

(Figs. 17-32, 34)

Peltoperla cora: Needham & Claassen, 1925:172. In part, male description (not Needham & Smith, 1916) Sierraperla cora: Stark & Stewart 1981:294. Egg and larval description (not Needham & Smith, 1916) Sierraperla cora: Stewart & Stark, 1988:263. Larval description (not Needham & Smith, 1916)

Sierraperla cora: Stewart & Stark, 2002:295. Larval description (not Needham & Smith, 1916)

Published Records (as S. cora but presumptively S. tolowa): California: Needham & Claassen (1925) report 10 males, 4 females from Shasta Springs (Siskiyou Co.), collected 8-29 June [year not given], by C.L. Fox and E.P. Van Duzee. Jewett (1960) includes records from Mt. Shasta and Shasta Springs in Siskiyou Co. Nelson & Stark (1987) include a larval record from Bidden Creek, Hwy 299, W of Cedar Flat in Trinity Co. Oregon: Jackson Co., Wrangle Camp, 8 July 1979, B. Stark, K.W. Stewart (collection data listed below larval habitus, Fig. 12.3 in Stewart & Stark 1988, 2002).

Material Examined. Holotype ♂ (reared), California, Del Norte Co., small falls into Patrick Creek, Patrick Creek Rd, 41° 53.180’N, 123° 51.124’W, 24 May 2014, B. Stark, A.B. Harrison (USNM). Paratypes: California: Del Norte Co., same data as holotype, 4♂, 1♀ (3♂, 1♀ reared) (BPSC). Humboldt Co., Boise Creek, Hwy 299, W Willow Creek, 24 May 2006, J.J. Lee, 1♂, 2♀ (BYUC). Unnamed stream, Hwy 299, mile 31.48, 11 July 2008, J.J. Lee, 1♂ (JJLC). Small waterfall, Hwy 299, W Boise Creek Campground, 40° 55.985’N, 123° 40.550’W, 25 May 2014, B. Stark, A.B. Harrison, 1♂ (BPSC). Siskiyou Co., Big Spring, Mt. Shasta City Park, 7 July 1979, B. Stark, K.W. Stewart, 3♂, 6♀, 14 larvae (BPSC). Same site, 15 May 1982, D. Zeigler, B. Stark, 9♂, 7♀ (BPSC). Same site, 20 May 1998, B. Stark, C.R. Nelson, S.W. Szczytko, I. Sivec, 1♂ (BPSC). Same site, 19 May 2001, B. Stark, K.W. Stewart, 1♀ (BPSC). Same site, 8 June 2004, B. Stark, R.W. Baumann, 1♂, 1♀ (BPSC). Same site, 28 June 2009, B. Stark, A. Harrison, K. Nye, 2♀ (BPSC). Same site, 23 May 2014, B.C. Kondratieff, C. Verdone, J.B. Sandberg, B. Stark, 5♂, 4♀, reared (CSUC). Mt. Shasta, Sacramento River, 5 June 1965, S.G. Jewett, Jr., 1♂, 1♀ (BYUC). Trinity Co., Hennessy Creek abv. Burnt Ranch Campground, Hwy 299, 22 June 1985, R.W. Baumann, C.R. Nelson, M. Whiting, 2♂, 4♀ (BYUC). Oregon: Jackson Co., 1 mile N Wrangle Camp, Rogue River National Forest, 8 July 1979, B. Stark, K.W. Stewart, 2♀, 14 larvae (BPSC). Pearsony Falls, Prospect Natural Trail, Mill Creek, 10 June 2004, B. Stark, R.W. Baumann, 1♀ (BPSC). Split Rock Creek, Wagner Gap Rd, 42.09479°N, 122.774°W, 22 May 2014, B.C. Kondratieff, C. Verdone, J.B. Sandberg, B. Stark, 5♂, 4♀, reared (CSUC).

Additional Specimens. California: Del Norte Co., Same data as holotype, 4 larvae, 3 in 95% EtOH (BPSC). Small falls above Shelly Creek into Patrick Creek, Patrick Creek Rd, 41° 54.320’N, 123° 51.370’W, 24 May 2014, B. Stark, A.B. Harrison, 6 larvae (95% EtOH) (BPSC). Humboldt Co., Ruby Creek, Hwy 299, 40° 54.470’N, 123° 43.093’W, 25 May 2014, B. Stark, A.B. Harrison, 3 larvae (95% EtOH) (BPSC). Boise Creek, Hwy 299, abv. Boise Creek Campground, 40.94155°N, 123.65765°W, 20 April 2009, J.B. Sandberg, 1 larva (JBSC). Small waterfall, Hwy 299, abv. Boise Creek Campground, 40° 55.985’N, 123° 40.550’W, 25 May 2014, B. Stark, A.B. Harrison, 5 larvae (95% EtOH) (BPSC). Siskiyou Co., Big Spring, Mt. Shasta City Park, 41° 19.692’N, 122° 19.605’W, 23 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C. Verdone, 3 larvae (95% EtOH) (BPSC). Oregon: Curry Co., Elk River Canyon, seep 1 mi below fish hatchery, 3 June 1991, B. Stark, R.W. Baumann, C. Henderson, 5 larvae (BPSC). Jackson Co., 1 mi N Wrangle Camp, Rogue River National Forest, 8 July 1979, B. Stark, K.W. Stewart, 14 larvae (BPSC). 15 mi SW Talent, Rogue River National Forest, 8 July 1979, B. Stark, K.W. Stewart, 13 larvae (BPSC). Sheep Creek, Wagner Gap Rd, 42° 06.442’N, 122° 47.921’W, 22 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C. Verdone, 1 larva (95% EtOH) (BPSC). Split Rock Creek, Wagner Gap Rd, 42° 05.676’N, 122° 46.493’W, 22 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C. Verdone, 5 larvae (95% EtOH), 2 larvae (95% EtOH) (JBSC). Wagner Creek, Wagner Creek Rd crossing, 42° 10.222’N, 122° 47.045’W, 22 May 2014, B. Stark, B.C. Kondratieff, J.B. Sandberg, C. Verdone, 1 larva (95% EtOH) (BPSC), 4 larvae (95% EtOH) (JBSC). Josephine Co., Lake Creek, below Oregon Caves National Monument, 9 July 1979, B. Stark, K.W. Stewart, 4 larvae (BPSC).

Adult Habitus. General color black patterned with yellow-brown (Figs. 34). Head mostly yelloworange but with extensive dark pigment in area surrounding ocelli; interocellar area mostly pale but with a narrow median band separating pale areas around ocelli (Fig. 17); antennal segments 1-7 pale on posterior margin and black on anterior margin, additional segments uniformly black. Pronotum mostly dark except for pale curved band extending from posteromedian line to midlateral margin; an additional pale diamond shaped area located near anteromedian margin, and an intricate pattern of small pale spots and lines occurs adjacent to midline and extends laterally onto discs. Femora brown, darker along dorsum; tibiae dark brown but bearing a narrow pale line on ventral margin. Wing membrane dusky, veins black.

Male. Forewing length ca. 15-18 mm. Abdominal tergum 10 yellow over most of surface, but bearing a transverse dark, anterior band which constricts near midlength; epiproct sclerite almost rectangular, but wider near anterior margin and often strongly excavated with a pale V-shaped notch on anteromedian margin (Fig. 18). Abdominal sternum 9 with a wide, oval hammer. Ventral aspect of aedeagus a wide membranous bag without sclerites and with sparse thin setalspines on lateral lobes; lateral lobes near midlength bifurcate; smaller pair of lobes located between bases of lateral pair; apex broadly and shallowly notched, each lobe consisting of a pair of closely appressed small lobes (Fig. 19).

Female. Forewing length ca. 18-21 mm. Subgenital plate extends over ca. half of abdominal sternum 9; lateral margins subparallel for much of plate length; apical margin bearing a slight median notch (Fig. 20).

Larva. Described by Stewart & Stark (1988, 2002) as S. cora.

Egg. Hemispherical in lateral aspect, polar aspects circular (Figs. 21, 24, 25). Diameter (n = 16) ca. 376- 420 μm. Collar a flattened disc covered with shallow pits and reticulations; collar surrounded by a narrow groove (Fig. 21). Anchor flattened, membranous, and bearing clusters of globular bodies around outer margin; diameter (n = 5) ca. 180-210 μm. Lateral surface of egg covered with pits; micropyles located in an irregular ring nearer collar than anterior pole (Fig. 24); orifices smaller in diameter than adjacent pits and surrounded by slightly raised, oval rims. Anterior surface completely covered with pits, surrounded by raised, follicle cell impression walls; walls thickened, height and thickness irregular; inner dimensions of FCIs relatively small and almost closed in some (Figs. 22-32).

Etymology. The species name honors the Native American Tolowa people whose homeland, based in northwestern California and southwestern Oregon, may have encompassed the site in Del Norte County, California where the type specimen was collected.

Diagnosis. This species appears distinct by virtue of its dark habitus, the bilobed lateral aedeagal lobes, the chorionic detail of the anterior pole of the egg and the barcode sequence of the COI gene. Females may be separated by comparison of subgenital plate shape and size of the median notch. No morphological characters are known which permit separation of larvae of the two species of Sierraperla. Because present data suggests the two species are allopatric we assigned identifications of some larval specimens on this basis when no barcode or adult data were available. Sierraperla cora is known from sites in the Sierra Nevada and Warner Mountains of California and Nevada, whereas S. tolowa is known from the Coast and Cascade mountains of southern Oregon and northern California. A possible exception to the allopatric distribution occurs in Siskiyou Co., California, where populations of both species were discovered near Mt. Shasta. Adult specimens from Big Spring, Mt. Shasta City Park were diagnosed based on the dark habitus, bilobed lateral aedeagal lobes and chorionic detail of the anterior pole (Figs. 24-26), and larval specimens from this site and Black Butte Spring, near Weed were diagnosed by comparing COI sequences. Adults and larvae from the Mt. Shasta City Park site are both identified as S. tolowa. Unfortunately, no adult specimens are available from Black Butte Spring but the larval COI sequence from this population is consistent with that of other S. cora populations.

Distribution. Sierraperla is currently known from a region of northern California, southern Oregon and western Nevada. The northernmost known locality is a site in Jackson Co., Oregon at Pearsony Falls on Mill Creek, the southernmost known locality is Bendorf Spring in El Dorado Co., California, the westernmost site is in Elk River Canyon, Curry Co., Oregon, and the easternmost site is in Douglas Co., Nevada (R. Bottorff, personal communication). Sierraperla appears to be rare along the western slope of the Coast Range but has been collected from upper Freshwater Creek in Humboldt Co., California by a colleague (J.J. Lee, personal communication).

DNA barcoding. Of the 48 specimens submitted, CCDB returned 44 high quality sequences (~92%) meeting the standards for barcode compliance of the Consortium for DNA Barcoding (CBOL) for Sierraperla cora (N = 19), S. tolowa (N = 24) and Soliperla campanula (N = 1). Mean genetic distances among individuals of S. cora and S. tolowa were 0.3% and 0.8% respectively (very similar). The mean genetic distance among specimens of these two species was 9% (quite divergent). Therefore, low levels of intraspecific and high levels of interspecific divergence of DNA barcode sequences among individuals of these taxa provide further support for the recognition of S. tolowa as a species distinct from S. cora (Hebert et al. 2003, Sweeney et al. 2011, Zhou et al. 2009). Mean genetic distances among S. cora, S. tolowa and exemplars of the other western peltoperlid genera (Soliperla and Yoraperla) calculated here are presented in Table 1 for reference.