Juga plicifera (I. Lea, 1838)

Figs 4–6

Melania plicifera I. Lea, 1838: 93, pl. 23 fig. 90.

Melania silicula A. Gould, 1847: 224–225 .

Melania rudens Reeve, 1860: unpaginated, species 224, pl. 33 fig. 224.

Goniobasis plicifera var. bulimoides Tryon, 1865: 238, pl. 24 figs 5–6.

Goniobasis plicifera var. oregonensis Tryon, 1865: 238, pl. 24 fig. 4.

Goniobasis hemphilli Henderson, 1935a: 96–97, pl. 4 fig. 1. Syn. nov.

Goniobasis hemphilli dallesensis Henderson, 1935a: 97, pl. 4 fig. 2. Syn. nov.

Melania silicula – Gould 1852: 141–142; “ 1856 ” [1860]: pl. 10 fig. 164–164a; 1862: 46.

Goniobasis hemphilli – Henderson 1936b, pl. 2 fig. 1.

Goniobasis hemphilli dallesensis – Henderson 1936b, pl. 2 fig. 2.

Juga (Juga) plicifera – Burch & Tottenham 1980: 152, fig. 448. — Burch 1982: 41, fig. 448; 1989: 152, fig. 448. — Holznagel & Lydeard 2000: 237.

Juga (Juga) silicula – Burch & Tottenham 1980: 152, fig. 449 (in part). — Burch 1982: 41, fig. 449 (in part); 1989: 152, fig. 449 (in part). — Neitzel & Frest 1992: B.25. — Holznagel & Lydeard 2000: 237. — Strong & Frest 2007: 45. — Johannes 2010b: 5.

Juga (Juga) hemphilli dallesensis – Burch & Tottenham 1980: 152, fig. 455. — Burch 1982: 41; 1989: 152, fig. 455. — Frest & Johannes 1993: 62; 1995a: 172; 2001: 59; 2010: 9, 26, fig. 2b. — Campbell et al. 2016: 160.

Juga silicula – Hawkins & Furnish 1987: 209. — Furnish 1990: 1; 2007: 14, fig. 6, lower left. — Turgeon et al. 1988: 65; 1998: 67. — Lee et al. 2006: 316. — Ó Foighil et al. 2009: 305. — Strong & Köhler 2009: 486. — Strong et al. 2011: 53. — Johnson et al. 2013: 282. — Köhler 2016: 270.

Juga plicifera – Turgeon et al. 1988: 65; 1998: 67. — Lee et al. 2006: 316. — Ó Foighil et al. 2009: 305. — Strong & Köhler 2009: 486. — Johnson et al. 2013: 263, 282, pl. 1.

Juga hemphilli – Turgeon et al. 1988: 65; 1998: 67. — Ó Foighil et al. 2009: 305. — Johnson et al. 2013: 282.

Juga (Juga) plicifera plicifera — Neitzel & Frest 1992: B.25 (in part). — Frest & Johannes 2001: 60; 2010: 11, 46 [as probably valid]. — Johannes 2015: 23. — Campbell et al. 2016: 160.

Juga (Juga) hemphilli hemphilli – Frest & Johannes 1993: 62; 1995a: 173; 2001: 59; 2010: 10, 35. — Campbell et al. 2016: 160.

Juga (Juga) hemphilli n. subsp. – Frest & Johannes 1993: 63.

Juga (Juga) n. sp. 1 – Frest & Johannes 1993: 63 (in part); 1995a: 175 (in part); 2001: 60 (in part).

Juga (Juga) n. sp. 3 – Frest & Johannes 1993: 63.

Juga (Juga) hemphilli n. subsp. 1 – Frest & Johannes 1995a: 174; 2001: 60.

Juga (Juga) silicula silicula – Frest & Johannes 2001: 60; 2010: 11, 50, fig. 2f. — Campbell et al. 2016: fig. 2c.

Juga nigrina – Lydeard et al. 2002: 401. — Strong & Köhler 2009: 486. — Strong et al. 2011: 53.

Juga sp. 1 – Prozorova & Rasshepkina 2004: 90, figs 1, 2.

Juga sp. 2 – Prozorova & Rasshepkina 2004: 91, figs 1, 3.

Juga sp. 3 – Prozorova & Rasshepkina 2004: 91, figs 1, 4.

Juga sp. 4 – Prozorova & Rasshepkina 2004: 91, figs 1, 5.

Juga sp. 5 – Prozorova & Rasshepkina 2004: 91, figs 1, 6.

Juga / Elimia hemphilli – Lee et al. 2006: 314–316, fig. 1a–c.

Juga / Elimia hemphilli dallesensis – Lee et al. 2006: 314–316, fig. 1a–c.

Juga hemphilli dallesensis – Ó Foighil et al. 2009: 305.

Juga (Juga) draytonii – Frest & Johannes 2010: 10, 34 (in part). — Campbell et al. 2016: 160.

Juga (Juga) plicifera bulimoides – Frest & Johannes 2010: 28 [as possibly valid].

Juga (Juga) plicifera oregonensis – Frest & Johannes 2010: 44 [as likely valid]. — Campbell et al. 2016: 160.

Juga (Juga) n. sp. – Johannes 2010a: 7.

Juga (Juga) cf. plicifera – Johannes 2013b: 16.

Juga (Juga) cf. silicula – Johannes 2013b: 19.

Juga (Juga) hemphilli n. subsp. 2 – Johannes 2015: 23.

Juga plicata – Köhler 2016: 270 (error for J. plicifera).

Juga (Juga) OTU 1 – Campbell et al. 2016: 160.

Juga (Juga) OTU 2 – Campbell et al. 2016: 160.

Juga plicifera group – Strong & Whelan 2019: 89.

non Juga (Juga) hemphilli hemphilli – Burch & Tottenham 1980: 152, fig. 454. — Burch 1982: 41, fig. 454; 1989: 152, fig. 454. — Frest & Johannes 2010: fig. 2c. [= Juga bulbosa]

non Juga / Elimia hemphilli – Lee et al. 2006: 316, fig. 1d. [= Elimia doolyensis; see Ó Foighil et al. 2009]

non Juga / Elimia hemphilli dallesensis – Lee et al. 2006: 316, fig. 1d. [= Elimia doolyensis; see Ó Foighil et al. 2009]

Material examined

Lectotype of Melania plicifera I. Lea, 1838 (here designated) (Fig. 4A) USA • “ Wahlamat [sic, Willamette River], near its junction with the Columbia River ”; T. Nuttall leg.; USNM 119244.

Paralectotypes of Melania plicifera I. Lea, 1838 USA • 23 spms; same collection data as for lectotype; USNM 1665628 (formerly USNM 119244X) • 1 spm; same collection data as for lectoype; MCZ 161911.

Lectotype of Melania silicula A. Gould, 1847 (here designated) (Fig. 4B) USA • “ Nisqually, Oregon [Territory]” [possibly Sequalitchew Creek; Strong & Frest 2007; Frest & Johannes 2010]; U.S. Exploring Expedition leg.; USNM 12137.

Paralectotypes of Melania silicula A. Gould, 1847 USA • 2 spms; same collection data as for lectotype; USNM 1665633 (ex USNM 12137).

Lectotype of Melania rudens Reeve, 1860 (here designated) (Fig. 4C) USA • locality unknown; H. Cuming collection; NHMUK 20110170/1.

Paralectotypes of Melania rudens Reeve, 1860 USA • 2 spms; same collection data as for lectotype; NHMUK 20110170/2, NHMUK 20110170/3.

Lectotype of Goniobasis plicifera var. bulimoides Tryon, 1865 (designated by Baker 1964: 180) (Fig. 4D) USA • “ Willamette River, at Eugene City”; W.M. Gabb leg.; ANSP 27542.

Paralectotypes of Goniobasis plicifera var. bulimoides Tryon, 1865 USA • 4 spms; same collection data as for lectotype; W.M. Gabb leg.; ANSP 453835 (ex ANSP 27542).

Lectotype of Goniobasis plicifera var. oregonensis Tryon, 1865 (designated by Baker 1964: 186) (Fig. 4E) USA • “ Willamette River, near its junction with the Columbia ”; R.E.C. Stearns leg.; ANSP 27550.

Paralectotypes of Goniobasis plicifera var. oregonensis Tryon, 1865 (Fig, 4E) USA • 6 spms; same collection data as for lectotype; ANSP 467223 (ex ANSP 27550).

Holotype of Goniobasis hemphilli Henderson, 1935 (Fig. 4F) USA • “ Near Portland, Oregon ” [possibly Johnson Creek; Frest & Johannes 1993, 2010]; Hemphill collection; UCM 21167a.

Paratype of Goniobasis hemphilli Henderson, 1935 USA • 1 spm; same collection data as for holotype; UCM 21167b (Henderson 1935a: pl. 4 fig. 1, at right).

Holotype of Goniobasis hemphilli dallesensis Henderson, 1935 (Fig. 4G) USA • “ Mill Creek, The Dalles, Oregon ”; 19 Aug. 1928; J. Henderson and Nelson leg.; UCM 16016a.

Paratypes of Goniobasis hemphilli dallesensis Henderson, 1935 USA • 3 spms; same collection data as for holotype; UCM 16016c (Henderson 1935a: pl. 4 fig. 2).

Remarks Another paratype (UCM 16016b) was not examined (McCoy 1964).

Other material examined 352 lots, 3435 specimens, of which 200 were sequenced.

USA – Washington • 5 spms; Clallam Co., Lake Crescent; [48.0581, -123.8132]; 19 Jul. 1935; V. Scheffer leg.; USNM 756742 • 5 spms; Clark Co., Burnt Bridge Creek Bridge at NE 110 th Ave, Vancouver; 45.6581, -122.5601; 23 Jun. 2015; K. Van Norman leg.; USNM 1413042 • 1 spm; same collection data as for preceding; GenBank: MK472579, MK464944; USNM 1413037 • 1 spm; same collection data as for preceding; GenBank: MK472580, MK464945, MK480955; USNM 1413038 • 1 spm; same collection data as for preceding; GenBank: MK472581, MK464946, MK480956; USNM 1413039 • 1 spm; same collection data as for preceding; USNM 1413040 • 1 spm; same collection data as for preceding; GenBank: MK472582, MK464947; USNM 1413041 • 38 spms; Clark Co., NW 179 th St; 45.7503, -122.7146; 6 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295164 • 1 spm; same collection data as for preceding; GenBank: MK472553, MK464920; USNM 1295163 • 1 spm; same collection data as for preceding; GenBank: MK472549, MK464916; USNM 1295159 • 1 spm; same collection data as for preceding; GenBank: MK472550, MK464917; USNM 1295160 • 1 spm; same collection data as for preceding; GenBank: MK472551, MK464918, MK480948; USNM 1295161 • 1 spm; same collection data as for preceding; GenBank: MK472552, MK464919; USNM 1295162 • 12 spms; Clark Co., Washington River, Vancouver; Clench leg.; USNM 1501083 • 1 spm; Cowlitz Co., Abernathy Creek behind Abernathy Fish Technology Center; 46.2267, -123.1478; 1 Aug. 2017; M. Piteo leg.; GenBank: KF680626; USNM 1597945 • 1 spm; same collection data as for preceding; GenBank: KF680627; USNM 1597944 • 45 spms; Grays Harbor Co., Black River; 46.8299, -123.1859; 4 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413220 • 1 spm; same collection data as for preceding; GenBank: MK472616, MK464981; USNM 1413215 • 1 spm; same collection data as for preceding; GenBank: MK472617, MK464982; USNM 1413216 • 1 spm; same collection data as for preceding; GenBank: MK472618, MK464983, MK480970; USNM 1413217 • 1 spm; same collection data as for preceding; GenBank: MK472619, MK464984; USNM 1413218 • 1 spm; same collection data as for preceding; GenBank: MK472620, MK464985; USNM 1413219 • 9 spms; Grays Harbor Co., Lake Quinault; [47.4729, -123.8683]; United States Fish Commission (USFC); Rutter and Brady leg.; USNM 218271 • 1 spm; Grays Harbor Co., Lake Quinault, Falls Creek Campground, boat launch E of private docks along South Shore Rd; 47.4701, -123.8457; 11 Aug. 2015; C. Fallon and E. Blevins leg.; GenBank: MK472682, MK465049, MK481000; USNM 1413471 • 1 spm; same collection data as for preceding; GenBank: MK472683, MK465050, MK481001; USNM 1413472 • 1 spm; same collection data as for preceding; GenBank: MK472684, MK465051; USNM 1413473 • 1 spm; same collection data as for preceding; GenBank: MK472685, MK465052; USNM 1413474 • 1 spm; Grays Harbor Co., Lake Quinault at Gatton Creek Campground, along rocky beach; 47.4738, -123.8392; 11 Aug. 2015; C. Fallon and E. Blevins leg.; GenBank: MK472676, MK465043, MK480995; USNM 1413465 • 1 spm; same collection data as for preceding; GenBank: MK472677, MK465044, MK480996; USNM 1413466 • 1 spm; same collection data as for preceding; GenBank: MK472678, MK465045, MK480997; USNM 1413467 • 1 spm; Jefferson Co., Quinault River, just N of S Shore Rd, between Fletcher and Bunch Canyons; 47.5297, -123.6987; 11 Aug. 2015; C. Fallon and E. Blevins leg.; GenBank: MK472679, MK465046, MK480998; USNM 1413468 • 1 spm; same collection data as for preceding; GenBank: MK472680, MK465047; USNM 1413469 • 1 spm; same collection data as for preceding; GenBank: MK472681, MK465048, MK480999; USNM 1413470 • 4 spms; King Co., L. Washington; [47.6214, -122.2557]; R.Z. Fahs leg.; USNM 169076 • 53 spms; Klickitat Co., Major Creek; 45.7154, -121.3509; 6 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413184 • 1 spm; same collection data as for preceding; GenBank: MK472604, MK464970, MK480965; USNM 1413182 • 1 spm; same collection data as for preceding; GenBank: MK472605, MK480966; USNM 1413183 • 41 spms; Lewis Co .. Ames Creek; 46.4516, -121.9929; 5 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413226 • 1 spm; same collection data as for preceding; GenBank: MK472621, MK464986, MK480971; USNM 1413221 • 1 spm; same collection data as for preceding; GenBank: MK472622, MK464987, MK480972; USNM 1413222 • 1 spm; same collection data as for preceding; GenBank: MK472623, MK464988; USNM 1413223 • 1 spm; same collection data as for preceding; GenBank: MK472624, MK464989; USNM 1413224 • 1 spm; same collection data as for preceding; GenBank: MK472625, MK464990; USNM 1413225 • 50 spms; Lewis Co., Cowlitz River at Spencer Rd Trout Hatchery; 46.4851, -122.7259; 5 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413175 • 1 spm; same collection data as for preceding; GenBank: MK472594, MK464960, MK480961; USNM 1413170 • 1 spm; same collection data as for preceding; GenBank: MK472595, MK464961; USNM 1413171 • 1 spm; same collection data as for preceding; GenBank: MK472596, MK464962; USNM 1413172 • 1 spm; same collection data as for preceding; GenBank: MK472597, MK464963; USNM 1413173 • 1 spm; same collection data as for preceding; GenBank: MK472598, MK464964, MK480962; USNM 1413174 • 46 spms; Lewis Co., Cowlitz River, at I-5 Crossing W of Toledo; [46.4149, -122.8897]; 24 May 1978; J. Landye leg.; on mud; USNM 892367 • 61 spms; Lewis Co., from stream in Lewis and Clark State Park [Lacamas Creek]; [46.5183, -122.8039]; 15 Jun. 1937; C.E. Burt leg.; USNM 473497 • 48 spms; Lewis Co., Little Falls [= Vader]; [46.395, -122.9578]; Arnold leg.; USNM 187284 • 49 spms; Lewis Co., Olequa Creek, tributary to Cowlitz River near Little Falls [= Vader]; [46.395, -122.9578]; R. Arnold leg.; USNM 181105 • 34 spms; Lewis Co., South Fork Newaukum River; 46.5751, -122.8353; 5 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413214 • 1 spm; same collection data as for preceding; GenBank: MK472611, MK464976, MK480968; USNM 1413209 • 1 spm; same collection data as for preceding; GenBank: MK472612, MK464977; USNM 1413210 • 1 spm; same collection data as for preceding; GenBank: MK472613, MK464978; USNM 1413211 • 1 spm; same collection data as for preceding; GenBank: MK472614, MK464979; USNM 1413212 • 1 spm; same collection data as for preceding; GenBank: MK472615, MK464980, MK480969; USNM 1413213 • 11 spms; Mason Co., Spider Lake, off of FR 23; 47.4068, -123.4372; 13 Aug. 2015; C. Fallon Mason and E. Blevins leg.; USNM 1413464 • 1 spm; same collection data as for preceding; GenBank: MK472675, MK465042, MK480994; USNM 1413463 • 5 spms; Pacific Co., Holcomb, Willapa River; [46.5699, -123.6154]; USFC; Rutter leg.; USNM 109895 • 1 spm; Pacific Co., Nasel, Nasel River [sic, Naselle, Naselle River]; [46.367, -123.8103]; USFC; Rutter leg.; USNM 109893 • 12 spms; Pierce Co., Ft Lewis, Sequalitchew Springs (main water source of Ft Lewis); [47.1116, -122.6016]; 19 Oct. 1951; Lt A. Mayfield leg.; USNM 598456 • 1 spm; Pierce Co., Lake George; [46.7878, -121.9036]; U.S. Exploring Expedition; USNM 5565 • 26 spms; Pierce Co., Roy; [47.0055, -122.5431]; 18 Aug. 1897; V. Bailey leg.; USNM 251946 • 6 spms; Pierce Co., S of Tacoma, in a brook; W.J. Eyerdam leg.; USNM 509403 • 6 spms; Pierce Co., Sequalitchew L., near creek; [47.1124, -122.6163]; 25 Apr. 1904; USFC; USNM 218270 • 9 spms; Pierce Co., Steilacoom Lake, near Tacoma; [47.1643, -122.5376]; 20 Oct. 1949; USNM 1294910 • 7 spms; Pierce Co., Steilacoon River [sic, Steilacoom = Chambers Creek]; [47.1948, -122.5528]; Smithsonian leg.; USNM 118931 • 2 spms; Pierce Co., Tacoma; [47.2538, -122.4349]; Hare leg.; USNM 133697 • 21 spms; Pierce Co., Tacoma, Steilacoom Lake; [47.1643, -122.5376]; 20 Oct. 1949; A.F. Bartsch leg.; USNM 601879 • 47 spms; Pierce Co., H.V. Chase leg.; USNM 742590 • 1 spm; Sherman Co., at Biggs Junction, downstream from Wallula Gap, Columbia River; [45.6727, -120.8346]; 10 Feb. 1949; J.J. Davis leg.; USNM 653137 • 72 spms; Skamania Co., Le Bong Creek at Stevenson; 45.6976, -121.9128; 5 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413181 • 1 spm; same collection data as for preceding; GenBank: MK472599, MK464965; USNM 1413176 • 1 spm; same collection data as for preceding; GenBank: MK472600, MK464966; USNM 1413177 • 1 spm; same collection data as for preceding; GenBank: MK472601, MK464967, MK480963; USNM 1413178 • 1 spm; same collection data as for preceding; GenBank: MK472602, MK464968, MK480964; USNM 1413179 • 1 spm; same collection data as for preceding; GenBank: MK472603, MK464969; USNM 1413180 • 10 spms; Skamania Co., Skamania at Skelton Rd; 45.6219, -122.0505; 5 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413208 • 1 spm; same collection data as for preceding; GenBank: MK472606, MK464971; USNM 1413203 • 1 spm; same collection data as for preceding; GenBank: MK472607, MK464972; USNM 1413204 • 1 spm; same collection data as for preceding; GenBank: MK472608, MK464973; USNM 1413205 • 1 spm; same collection data as for preceding; GenBank: MK472610, MK464975, MK480967; USNM 1413207 • 1 spm; same collection data as for preceding; GenBank: MK472609, MK464974; USNM 1413206 • 3 spms; Skamania Co., Stevenson; [45.695, -121.8928]; V. Jackson leg.; USNM 169055 • 12 spms; Thurston Co., Black Lake; [46.9881, -122.9793]; USFC; Rutter leg.; USNM 109862 • 14 spms; Thurston Co., Black River, N of bridge along SE 128 Ave, Littlerock; 46.902, -123.0239; 28 Jun. 2015; K. Van Norman leg.; USNM 1413048 • 1 spm; same collection data as for preceding; GenBank: MK464948, MK480957; USNM 1413043 • 1 spm; same collection data as for preceding; GenBank: MK472583, MK464949; USNM 1413044 • 1 spm; same collection data as for preceding; GenBank: MK472584, MK464950; USNM 1413045 • 1 spm; same collection data as for preceding; GenBank: MK472585, MK464951; USNM 1413046 • 1 spm; same collection data as for preceding; GenBank: MK472586, MK464952, MK480958; USNM 1413047 • 1 spm; same collection data as for preceding; GenBank: MK472587, MK464953; USNM 1413049 • 1 spm; same collection data as for preceding; GenBank: MK472588, MK464954; USNM 1413050 • 16 spms; Thurston Co., creek on Evergreen State College, Snyder Cove; [47.0842, -122.9742]; 10 Aug. 1983; J. Landye leg.; USNM 1152454 • 5 spms; Thurston Co., creeks near Olympia; Randolph leg.; USNM 130099 • 24 spms; Thurston Co., McAllister Creek at Steilacoom Rd Bridge, near McAllister Creek Hatchery; 47.0501, -122.7262; 29 Sep. 2005; T. Frest and E. Johannes leg.; USNM 1100657 • 19 spms; Thurston Co., Millersylvania State Park, Deep Lake, in picnic area; [46.9088, -122.9109]; 24 May 1978; J. Landye leg.; USNM 892364 • 12 spms; Thurston Co., Muck Creek just upstream from Nisqually River; [46.9975, -122.6264]; 10 Aug. 1983; J. Landye leg.; USNM 1152371 • 54 spms; Thurston Co., Nisqually River; 46.9334, -122.5608; 4 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413239 • 1 spm; same collection data as for preceding; GenBank: MK472626, MK464991, MK480973; USNM 1413233 • 1 spm; same collection data as for preceding; GenBank: MK472627, MK464992, MK480974; USNM 1413234 • 1 spm; same collection data as for preceding; GenBank: MK472628, MK464993; USNM 1413235 • 1 spm; same collection data as for preceding; GenBank: MK472629, MK464994; USNM 1413236 • 1 spm; same collection data as for preceding; GenBank: MK472630, MK464995, MK480975; USNM 1413237 • 1 spm; same collection data as for preceding; GenBank: MK472631, MK464996; USNM 1413238 • 35 spms; Thurston Co., Skookumchuck River; 46.7954, -122.7609; 4 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413245 • 1 spm; same collection data as for preceding; GenBank: MK472632, MK464997; USNM 1413240 • 1 spm; same collection data as for preceding; GenBank: MK472633, MK464998, MK480976; USNM 1413241 • 1 spm; same collection data as for preceding; GenBank: MK472634, MK464999; USNM 1413242 • 1 spm; same collection data as for preceding; GenBank: MK472635, MK465000; USNM 1413243 • 1 spm; same collection data as for preceding; GenBank: MK472636, MK465001; USNM 1413244 • 15 spms; Thurston Co., Skookumchuck River, Bucoda Volunteer Park, Bucoda; 46.7961, -122.8672; 28 Jun. 2015; K. Van Norman leg.; USNM 1413060 • 1 spm; same collection data as for preceding; GenBank: MK472589, MK464955, MK480959; USNM 1413055 • 1 spm; same collection data as for preceding; GenBank: MK472590, MK464956; USNM 1413056 • 1 spm; same collection data as for preceding; GenBank: MK472591, MK464957; USNM 1413057 • 1 spm; same collection data as for preceding; GenBank: MK472592, MK464958, MK480960; USNM 1413058 • 1 spm; same collection data as for preceding; GenBank: MK472593, MK464959; USNM 1413059 • 13 spms; Walla Walla Co., Walla Walla; USNM 119156 • 9 spms; Walla Walla Co., Walula [sic, Wallula], Walla Walla River; USFC; USNM 109897 • 26 spms; Whitman Co .. Snake Riv. tributary between Wawawai and Bishop; [46.6022, -117.3548]; 1 Apr. 1958; Richards leg.; USNM 795562 • 16 spms; Coloting River [sic, mislocalized], Washington Territory; Suckley leg.; USNM 119152 • 2 spms; Salmon River; USNM 1294901. — “ Oregon ” [Territory] • 11 spms; USNM 58979 • 7 spms; same collection data as for preceding; USNM 59211 • 5 spms; same collection data as for preceding; USNM 58975 • 6 spms; same collection data as for preceding; USNM 12135 • 6 spms; same collection data as for preceding; Newcomb leg.; USNM 119285 • 9 spms; Columbia River; Drayton leg.; USNM 119158. – Oregon • 103 spms; Benton Co., 2 mi from Corvallis, Willamette River; [44.5291, -123.2538]; 5 Jul. 1929; L. Sinitsin leg.; USNM 531451 • 134 spms; Benton Co., Alsea River; 13 Apr. 1958; Walter leg.; USNM 795571 • 5 spms; Benton Co., Corvallis; [44.5655, -123.2555]; 22 Jul. 1949; A.L. Mehring leg.; USNM 715700 • 48 spms; Benton Co., Corvallis, Mr. Little’s Ranch; 31 Jul. 1929; L. Sinitsin leg.; USNM 531205 • 1 spm; Benton Co., Corvallis, Oak Creek; [44.557, -123.282]; 14 Jul. 1929; L. Sinitsin leg.; USNM 531114 • 21 spms; Benton Co., Mary’s River, Mary’s River Park, Philomath; 44.5343, -123.3735; 14 Jul. 2015; K. Van Norman leg.; USNM 1413036 • 1 spm; same collection data as for preceding; GenBank: MK472575, MK464940; USNM 1413031 • 1 spm; same collection data as for preceding; GenBank: MK472576, MK464941, MK480953; USNM 1413032 • 1 spm; same collection data as for preceding; USNM 1413033 • 1 spm; same collection data as for preceding; GenBank: MK472577, MK464942, MK480954; USNM 1413034 • 1 spm; same collection data as for preceding; GenBank: MK472578, MK464943; USNM 1413035 • 1 spm; Clackamas Co., left bank of Clackamas River, 100 ft south (upstream) of mouth of Whale Creek; 45.1109, -122.0725; 19 May 2014; J. Williamson leg.; USNM 1413021 • 1 spm; same collection data as for preceding; GenBank: MK472569; USNM 1413022 • 1 spm; same collection data as for preceding; USNM 1413023 • 1 spm; same collection data as for preceding; USNM 1413024 • 4 spms; Clackamas Co., near Estacada, N fork of Molalla River [sic, possibly Clackamas River]; Jul. 1943; D. Macfarlane leg.; USNM 518284 • 12 spms; Clackamas Co., Oregon City; [45.3586, -122.6092]; L.B. Elliott leg.; USNM 509456 • 62 spms; Clackamas Co., Oswego, from a creek [possibly Tryon Creek]; [45.424, -122.661]; L. Bruner and H.F. Wickham leg.; USNM 504256 • 20 spms; Clackamas Co., small creek near Tawney’s Hotel [Tawney’s Mountain Home], 12 mi from Mount Hood; [45.3165, -121.9475]; Aug. 1916; J.G. Malone leg.; ANSP 115580 • 1 spm; Clackamas Co., Whale Creek, between confluence with Clackamas River and upstream 45 feet; 45.111, -122.0731; 19 May 2014; J. Williamson leg.; USNM 1413019 • 1 spm; same collection data as for preceding; USNM 1413020 • 5 spms; Clatsop Co., Astoria; [46.1882, -123.8275]; ANSP 264447 • 7 spms; same collection data as for preceding; ANSP 27546 • 15 spms; Clatsop Co., Astoria, Columbia River; [46.1882, -123.8275]; W.H. Dall leg.; USNM 119012 • 41 spms; Clatsop Co., at confluence with Bear Creek, Maki Rd; 46.1543, -123.6672; 7 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295182 • 1 spm; same collection data as for preceding; GenBank: MK472559, MK464926; USNM 1295177 • 1 spm; same collection data as for preceding; GenBank: MK472560, MK464927; USNM 1295178 • 1 spm; same collection data as for preceding; GenBank: MK472561, MK464928, MK480949; USNM 1295179 • 1 spm; same collection data as for preceding; GenBank: MK472562, MK464929; USNM 1295180 • 1 spm; same collection data as for preceding; GenBank: MK472563; USNM 1295181 • 2 spms; Clatsop Co., Columbia River, near Astoria; [46.1882, -123.8275]; H. Hemphill leg.; UF 80844 • 4 spms; same collection data as for preceding; E. White leg.; UF 80863 • 41 spms; Clatsop Co., Green Mountain Rd, ca 300 m upstream of confluence of North Fork and South Fork Klaskanine Rivers; 46.0865, -123.741; 7 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295069 • 1 spm; same collection data as for preceding; GenBank: MK472535, MK464901; USNM 1295064 • 1 spm; same collection data as for preceding; GenBank: MK472536, MK464902; USNM 1295065 • 1 spm; same collection data as for preceding; GenBank: MK472537, MK464903, MK480940; USNM 1295066 • 1 spm; same collection data as for preceding; GenBank: MK472538, MK464904, MK480941; USNM 1295067 • 1 spm; same collection data as for preceding; GenBank: MK472539, MK464905, MK480942; USNM 1295068 • 13 spms; Clatsop Co., just downstream of Gnat Creek Fish Hatchery, along U.S. Hwy 30; 46.177, -123.5034; 7 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295006 • 1 spm; same collection data as for preceding; GenBank: MK472519, MK464885, MK480933; USNM 1295001 • 1 spm; same collection data as for preceding; GenBank: MK472520, MK464886, MK480934; USNM 1295002 • 1 spm; same collection data as for preceding; GenBank: MK472521, MK464887, MK480935; USNM 1295003 • 1 spm; same collection data as for preceding; GenBank: MK472522, MK464888, MK480936; USNM 1295004 • 1 spm; same collection data as for preceding; GenBank: MK472523, MK464889; USNM 1295005 • 19 spms; Clatsop Co., old U.S. Hwy 30; 46.1663, -123.673; 7 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1294987 • 1 spm; same collection data as for preceding; GenBank: MK472513; USNM 1294981 • 1 spm; same collection data as for preceding; GenBank: MK472514, MK464880; USNM 1294982 • 1 spm; same collection data as for preceding; GenBank: MK472515, MK464881, MK480930; USNM 1294983 • 1 spm; same collection data as for preceding; GenBank: MK472516, MK464882; USNM 1294984 • 1 spm; same collection data as for preceding; GenBank: MK472517, MK464883, MK480931; USNM 1294985 • 1 spm; same collection data as for preceding; GenBank: MK472518, MK464884, MK480932; USNM 1294986 • 25 spms; Clatsop Co., Tweedle Rd; 45.8963, -123.5544; 7 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1294980 • 1 spm; same collection data as for preceding; GenBank: MK472507, MK464874, MK480928; USNM 1294974 • 1 spm; same collection data as for preceding; GenBank: MK472508, MK464875; USNM 1294975 • 1 spm; same collection data as for preceding; GenBank: MK472509, MK464876; USNM 1294976 • 1 spm; same collection data as for preceding; GenBank: MK472510, MK464877, MK480929; USNM 1294977 • 1 spm; same collection data as for preceding; GenBank: MK472511, MK464878; USNM 1294978 • 1 spm; same collection data as for preceding; GenBank: MK472512, MK464879; USNM 1294979 • 22 spms; Clatsop Co., Youngs River Falls, just upstream of Youngs River Rd; 46.0675, -123.7888; 7 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295023 • 1 spm; same collection data as for preceding; GenBank: MK472524, MK464890; USNM 1295018 • 1 spm; same collection data as for preceding; GenBank: MK472525, MK464891; USNM 1295019 • 1 spm; same collection data as for preceding; GenBank: MK472526, MK464892; USNM 1295020 • 1 spm; same collection data as for preceding; GenBank: MK472527, MK464893; USNM 1295021 • 1 spm; same collection data as for preceding; GenBank: MK472528, MK464894, MK480937; USNM 1295022 • 20 spms; Columbia Co., McBride Creek, Smith Rd; [45.8912, -122.8178]; 29 Jun. 2002; T. Frest and E. Johannes leg.; USNM 1270862 • 29 spms; Columbia Co., Scappoose Creek on road to Pittsburg and Vernonia; [45.7638, -122.88]; J. Landye leg.; USNM 1152453 • 53 spms; Columbia Co., U.S. Hwy 30, Scapoose; 45.7708, -122.8791; 7 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295043 • 1 spm; same collection data as for preceding; GenBank: MK472529, MK464895; USNM 1295037 • 1 spm; same collection data as for preceding; GenBank: MK472530, MK464896, MK480938; USNM 1295038 • 1 spm; same collection data as for preceding; GenBank: MK472531, MK464897; USNM 1295039 • 1 spm; same collection data as for preceding; GenBank: MK472532, MK464898; USNM 1295040 • 1 spm; same collection data as for preceding; GenBank: MK472533, MK464899, MK480939; USNM 1295041 • 1 spm; same collection data as for preceding; GenBank: MK472534, MK464900; USNM 1295042 • 28 spms; Deschutes Co., just downstream of U.S. Hwy 20 (FS campground); 44.3567, -121.6107; 16 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1294973 • 1 spm; same collection data as for preceding; GenBank: MK472504, MK464871; USNM 1294970 • 1 spm; same collection data as for preceding; GenBank: MK472503, MK464870, MK480926; USNM 1294969 • 1 spm; same collection data as for preceding; GenBank: MK472502, MK464869; USNM 1294968 • 1 spm; same collection data as for preceding; GenBank: MK472505, MK464872, MK480927; USNM 1294971 • 1 spm; same collection data as for preceding; GenBank: MK472506, MK464873; USNM 1294972 • 55 spms; Gilliam Co., Columbia River, 5 mi W of Arlington; [45.7109, -120.3043]; 17 Oct. 1960; D. Bauer leg.; UF 545965 • 23 spms; same collection data as for preceding; UF 545985 • 26 spms; Hood River Co., just downstream of the mouth of Hood River; 45.7149, -121.5135; 15 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1294961 • 1 spm; same collection data as for preceding; GenBank: MK472495, MK464862, MK480924; USNM 1294959 • 1 spm; same collection data as for preceding; GenBank: MK472494, MK464861; USNM 1294958 • 1 spm; same collection data as for preceding; GenBank: MK472493, MK464860; USNM 1294957 • 1 spm; same collection data as for preceding; GenBank: MK472492, MK464859; USNM 1294956 • 1 spm; same collection data as for preceding; GenBank: MK472496, MK464863; USNM 1294960 • 23 spms; Lane Co., 29 mi E of Eugene, near Vida, McKenzie River; [44.1425, -122.5633]; 12 Aug. 1929; L. Sinitsin leg.; USNM 531104 • 6 spms; Lane Co., 4 mi N of Sea Lion Cave, large creek [sic, likely Big Creek]; [44.1744, -124.1153]; 27 May 1971; A.W. Vazquez leg.; USNM 758161 • 22 spms; Lane Co., end of Crossroads Lane (boat ramp); 44.1879, -123.1443; 15 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295176 • 1 spm; same collection data as for preceding; GenBank: MK472554, MK464921; USNM 1295171 • 1 spm; same collection data as for preceding; GenBank: MK472555, MK464922; USNM 1295172 • 1 spm; same collection data as for preceding; GenBank: MK472556, MK464923; USNM 1295173 • 1 spm; same collection data as for preceding; GenBank: MK472557, MK464924; USNM 1295174 • 1 spm; same collection data as for preceding; GenBank: MK472558, MK464925; USNM 1295175 • 10 spms; Lane Co., Eugene; [44.0455, -123.1254]; USNM 431330 • 4 spms; Lane Co., Eugene, Mackenzie River; [44.1124, -123.0468]; USFC; Cramer and Otaki leg.; USNM 109896 • 13 spms; Lane Co., Eugene, Willamette River; [44.0638, -123.1007]; D.T. Jones leg.; USNM 510720 • 38 spms; Lane Co., Long Tom River at Poodle Creek Rd; 44.1434, -123.4298; 16 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413341 • 1 spm; same collection data as for preceding; GenBank: MK472637, MK465002, MK480977; USNM 1413336 • 1 spm; same collection data as for preceding; GenBank: MK472638, MK465003; USNM 1413337 • 1 spm; same collection data as for preceding; GenBank: MK472639, MK465004; USNM 1413338 • 1 spm; same collection data as for preceding; GenBank: MK472640, MK465005; USNM 1413339 • 1 spm; same collection data as for preceding; GenBank: MK472641, MK465006, MK480978; USNM 1413340 • 8 spms; Lane Co., N of Marchfield [sic, Marshfield], Ten Mile Creek; [44.2241, -124.1096]; J. Henderson leg.; USNM 425498 • 88 spms; Lane Co., near Mapleton, Sheverton opposite electric station; [44.0213, -123.8555]; Jul. 1929; L. Sinitsin leg.; where river very slow and shallow; USNM 531107 • 29 spms; Lane Co., Riverlet opp. Mapleton; [44.0333, -123.855]; Jul. 1929; L. Sinitsin leg.; USNM 531105 • 46 spms; Lane Co., Siuslaw Hatchery, creek ca 1 mi from office; [44.0939, -123.7739]; Jul. 1929; L. Sinitsin leg.; USNM 531106 • 28 spms; Lane Co., Siuslaw River at Brickerville; 44.0606, -123.8852; 16 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413395 • 1 spm; same collection data as for preceding; GenBank: MK472662, MK465027; USNM 1413390 • 1 spm; same collection data as for preceding; GenBank: MK472663, MK465028, MK480988; USNM 1413391 • 1 spm; same collection data as for preceding; GenBank: MK472664, MK465029; USNM 1413392 • 1 spm; same collection data as for preceding; GenBank: MK472665, MK465030; USNM 1413393 • 1 spm; same collection data as for preceding; GenBank: MK472666, MK465031; USNM 1413394 • 31 spms; Lane Co., Siuslaw River, ca 1 mi from hatchery in the forest; [44.0939, -123.7739]; Jul. 1929; L. Sinitsin leg.; USNM 531074 • 24 spms; Lane Co., Siuslaw River, Siuslaw hatchery, opp. the house; [44.0939, -123.7739]; Jul. 1929; L. Sinitsin leg.; USNM 531108 • 1 spm; Lane Co., Siuslaw Salmon Hatchery; [44.0939, -123.7739]; 22 May 1929; L. Sinitsin leg.; USNM 531098 • 5 spms; Lane Co., Sutton Lake; [44.0586, -124.0899]; 27 May 1971; A.W. Vazquez leg.; USNM 758160 • 66 spms; Lane Co., Swisshome, Siuslaw Riv.; [44.0553, -123.8014]; Jul. 1929; L. Sinitsin leg.; USNM 531273 • 3 spms; Lincoln Co., Devils Lake; [44.9808, -123.9921]; E.C. Huffman leg.; USNM 593063 • 13 spms; Lincoln Co., just below Nashville, Yaquina River; [44.6531, -123.6077]; 8 Sep. 1977; C. Simmons leg.; USNM 771822 • 21 spms; Lincoln Co., near Eddyville, mi 21 on US 20, large creek (Mary’s Riv.?) [= Little Elk Creek]; [44.6287, -123.7674]; 31 May 1971; A.W. Vasquez leg.; USNM 758152 • 52 spms; Linn Co., Albany; [44.6384, -123.1065]; 1900; H.F. Wickham leg.; USNM 509443 • 37 spms; Linn Co., Calapooia River at McKercher Park; 44.3585, -122.8763; 16 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413347 • 1 spm; same collection data as for preceding; GenBank: MK472642, MK465007; USNM 1413342 • 1 spm; same collection data as for preceding; GenBank: MK472643, MK465008; USNM 1413343 • 1 spm; same collection data as for preceding; GenBank: MK472644, MK465009, MK480979; USNM 1413344 • 1 spm; same collection data as for preceding; GenBank: MK472645, MK465010, MK480980; USNM 1413345 • 1 spm; same collection data as for preceding; GenBank: MK472646, MK465011, MK480981; USNM 1413346 • 52 spms; Linn Co., Thomas Creek at Chapin Park in Scio; 44.7044, -122.8472; 16 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413413 • 1 spm; same collection data as for preceding; GenBank: MK472670, MK465037; USNM 1413408 • 1 spm; same collection data as for preceding; GenBank: MK472671, MK465038; USNM 1413409 • 1 spm; same collection data as for preceding; GenBank: MK472672, MK465039, MK480993; USNM 1413410 • 1 spm; same collection data as for preceding; GenBank: MK472673, MK465040; USNM 1413411 • 1 spm; same collection data as for preceding; GenBank: MK472674, MK465041; USNM 1413412 • 58 spms; Marion Co., Drift Creek on Cascade Highway NE; 44.9693, -122.8094; 16 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413371 • 1 spm; same collection data as for preceding; GenBank: MK472652, MK465017, MK480984; USNM 1413366 • 1 spm; same collection data as for preceding; GenBank: MK472653, MK465018; USNM 1413367 • 1 spm; same collection data as for preceding; GenBank: MK472654, MK465019; USNM 1413368 • 1 spm; same collection data as for preceding; GenBank: MK472655, MK465020, MK480985; USNM 1413369 • 1 spm; same collection data as for preceding; GenBank: MK472656, MK465021, MK480986; USNM 1413370 • 1 spm; Marion Co., Mehama; [44.7889, -122.618]; C.V. Piper leg.; USNM 225925 • 15 spms; Marion Co., Salem; [44.9446, -123.0461]; M. Witter leg.; USNM 509400 • 8 spms; same collection data as for preceding; A.K. F. leg.; USNM 509454 • 2 spms; Marion Co., Salem; [44.9446, -123.0461]; USNM 534927 • 2 spms; Multnomah Co., Benson State Recreation Area, along Interstate 84, near Multnomah Falls; 45.577, -122.1262; 8 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295075 • 1 spm; same collection data as for preceding; GenBank: MK472540, MK464906, MK480943; USNM 1295070 • 1 spm; same collection data as for preceding; GenBank: MK472541, MK464907, MK480944; USNM 1295071 • 1 spm; same collection data as for preceding; GenBank: MK472542, MK464908; USNM 1295072 • 1 spm; same collection data as for preceding; GenBank: MK472543, MK464909; USNM 1295073 • 1 spm; same collection data as for preceding; GenBank: MK472544, MK464910, MK480945; USNM 1295074 • 30 spms; Multnomah Co., Benson State Recreation Area, along Interstate 84, near Multnomah Falls; 45.5767, -122.13; 8 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295200 • 1 spm; same collection data as for preceding; GenBank: MK472564, MK464930; USNM 1295195 • 1 spm; same collection data as for preceding; GenBank: MK472565, MK464931; USNM 1295196 • 1 spm; same collection data as for preceding; GenBank: MK472566, MK464932; USNM 1295197 • 1 spm; same collection data as for preceding; GenBank: MK472567, MK464933; USNM 1295198 • 1 spm; same collection data as for preceding; GenBank: MK472568, MK464934, MK480950; USNM 1295199 • 18 spms; Multnomah Co., Coopey Creek above falls at Angels Rest trail crossing, Columbia River Gorge Natl Scenic Area; [45.5616, -122.1629]; 22 Jul. 2015; R. Wisseman leg.; USNM 1438822 • 2 spms; Multnomah Co., E of Portland; H. Hemphill leg.; USNM 30585 • 5 spms; Multnomah Co., near Horse Tail Falls; [45.5896, -122.0687]; E.C. Huffman leg.; USNM 593061 • 3 spms; Multnomah Co., Portland; Lewis leg.; USNM 715183 • 22 spms; same collection data as for preceding; L.B. Elliott leg.; USNM 509453 • 16 spms; same collection data as for preceding; H. Hemphill leg.; USNM 59132 • 51 spms; same collection data as for preceding; 27 Jun. 1905; M.W. Lyon Jr. leg.; USNM 187283 • 12 spms; same collection data as for preceding; May 1895; W. Palmer leg.; USNM 184298 • 4 spms; same collection data as for preceding; B. Shimek leg.; USNM 512975 • 2 spms; same collection data as for preceding; H. Hemphill leg.; USNM 30582 • 2 spms; Multnomah Co., Portland, Johnson Creek; [45.4686, -122.5657]; 14 May 1905; J.E. Benedict leg.; USNM 184189 • 16 spms; Multnomah Co., tributary of Young Creek at old highway crossing, Columbia River Gorge Natl Scenic Area; [45.5466, -122.1978]; 22 Jul. 2015; R. Wisseman leg.; USNM 1665621 (formerly USNM 1438822) • 31 spms; Polk Co., Mill Creek at Mill Creek Park; 44.9866, -123.4259; 16 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413389 • 1 spm; same collection data as for preceding; GenBank: MK472657, MK465022, MK480987; USNM 1413384 • 1 spm; same collection data as for preceding; GenBank: MK472658, MK465023; USNM 1413385 • 1 spm; same collection data as for preceding; GenBank: MK472659, MK465024; USNM 1413386 • 1 spm; same collection data as for preceding; GenBank: MK472660, MK465025; USNM 1413387 • 1 spm; same collection data as for preceding; GenBank: MK472661, MK465026; USNM 1413388 • 34 spms; Polk Co., Mill Creek, Buell County Park, near Sheridan; 45.0229, -123.4181; 14 Jul. 2015; K. Van Norman leg.; USNM 1413030 • 1 spm; same collection data as for preceding; GenBank: MK472570, MK464935, MK480951; USNM 1413025 • 1 spm; same collection data as for preceding; GenBank: MK472571, MK464936, MK480952; USNM 1413026 • 1 spm; same collection data as for preceding; GenBank: MK472572, MK464937; USNM 1413027 • 1 spm; same collection data as for preceding; GenBank: MK472573, MK464938; USNM 1413028 • 1 spm; same collection data as for preceding; GenBank: MK472574, MK464939; USNM 1413029 • 48 spms; Polk Co., Willamette River at Salem Wallace Marine Park; 44.946, -123.044; 16 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413353 • 1 spm; same collection data as for preceding; GenBank: MK472647, MK465012, MK480982; USNM 1413348 • 1 spm; same collection data as for preceding; GenBank: MK472648, MK465013, MK480983; USNM 1413349 • 1 spm; same collection data as for preceding; GenBank: MK472649, MK465014; USNM 1413350 • 1 spm; same collection data as for preceding; GenBank: MK472650, MK465015; USNM 1413351 • 1 spm; same collection data as for preceding; GenBank: MK472651, MK465016; USNM 1413352 • 12 spms; Tillamook Co., 8.7 mi N of Beaver, Fawcett Creek, on U.S. Hwy 101; [45.3897, -123.8031]; 28 May 1971; A.W. Vasquez leg.; USNM 758162 • 7 spms; Tillamook Co., Bewley Creek; [45.3864, -123.8549]; E.C. Huffman leg.; USNM 593060 • 6 spms; Tillamook Co., Farmer Creek; [45.2542, -123.8616]; E.C. Huffman leg.; USNM 593067 • 3 spms; Tillamook Co., Hoquarton Slough; [45.4621, -123.8338]; E.C. Huffman leg.; USNM 593065 • 2 spms; Tillamook Co., Lake Lytle; [45.6239, -123.9401]; E.C. Huffman leg.; USNM 593064 • 10 spms; Tillamook Co., Rockaway, Lake Lytle; [45.6239, -123.9401]; 28 May 1971; A.W. Vazquez leg.; USNM 758159 • 2 spms; Tillamook Co., Trask River; E.C. Huffman leg.; USNM 593066 • 5 spms; Tillamook Co., Wilson River; E.C. Huffman leg.; USNM 593062 • 2 spms; Wasco Co., The Dalles; [45.6030, -121.1800]; H. Hemphill leg.; USNM 30548 • 4 spms; same collection data as for preceding; USNM 30564 • 1 spm; same collection data as for preceding; USNM 509426 • 138 spms; same collection data as for preceding; H.F. Wickham leg.; USNM 509449 • 32 spms; Wasco Co., Thompson City Park, just upstream of Interstate 84, The Dalles; 45.6046, -121.1894; 9 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1294967 • 1 spm; same collection data as for preceding; GenBank: MK472497, MK464864; USNM 1294962 • 1 spm; same collection data as for preceding; GenBank: MK472498, MK464865; USNM 1294963 • 1 spm; same collection data as for preceding; GenBank: MK472499, MK464866, MK480925; USNM 1294964 • 1 spm; same collection data as for preceding; GenBank: MK472500, MK464867; USNM 1294965 • 1 spm; same collection data as for preceding; GenBank: MK472501, MK464868; USNM 1294966 • 52 spms; Washington Co., Tualatin Hills Park, Cedar Mills; 45.5214, -122.8377; 7 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295152 • 1 spm; same collection data as for preceding; GenBank: MK472545, MK464911; USNM 1295147 • 1 spm; same collection data as for preceding; GenBank: MK464912; USNM 1295148 • 1 spm; same collection data as for preceding; GenBank: MK472546, MK464913, MK480946; USNM 1295149 • 1 spm; same collection data as for preceding; GenBank: MK472547, MK464914; USNM 1295150 • 1 spm; same collection data as for preceding; GenBank: MK472548, MK464915, MK480947; USNM 1295151 • 2 spms; Yamhill Co., near McMinnville, Muddy Creek; [45.127, -123.3029]; Jun. 1943; A.W. Macy leg.; USNM 518283 • 48 spms; Yamhill Co., South Yamhill River at McMinnville City Park; 45.2071, -123.1805; 16 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413401 • 1 spm; same collection data as for preceding; GenBank: MK472667, MK465032, MK480989; USNM 1413396 • 1 spm; same collection data as for preceding; GenBank: MK465033, MK480990; USNM 1413397 • 1 spm; same collection data as for preceding; GenBank: MK472668, MK465034, MK480991; USNM 1413398 • 1 spm; same collection data as for preceding; GenBank: MK472669, MK465035, MK480992; USNM 1413399 • 1 spm; same collection data as for preceding; GenBank: MK465036; USNM 1413400 • 2 spms; Willamette River; H.E. Dore leg.; USNM 99305 • 2 spms; same collection data as for preceding; S. Smith leg.; USNM 321801 • 2 spms; same collection data as for preceding; USNM 218569 .

Description

SHELL. Thick, large, reaching ~ 3.5 cm in length; turriform, narrowly conical to cylindrical in shape, spire high, highly variable in sculpture, color, and banding (Fig. 5). Whorls somewhat flattened to moderately convex, occasionally weakly shouldered or with a subsutural ramp, suture weakly to moderately impressed. Aperture oval in shape, lip smooth, slightly sinuous. Spiral sculpture of numerous, fine striae, typically moderately to strongly developed and rarely indistinct, highly variable in strength and number; sporadic slightly elevated lirae. Axial sculpture of fine, sinuous growth lines, and plications on early whorls when preserved; plications regular and even, moderately to strongly developed, occasionally shouldered, orthocline, opisthocline, to strongly opisthocyrt, sometimes extending full length of shell, but frequently obsolete on body whorl; may be thickened or beaded at intersections with spiral striae. Shell black, dark brown, to olive or tan in color. Bands present or absent, one to three in number and variable in width, reddish-purple in color. Interior of aperture white or cream to dark purple in color, occasionally with yellowish-orange rim inside lip; bands when present visible inside aperture. Base of columella occasionally tinged reddish-purple.

RADULA. Rachidian narrowly rectangular, wider than tall, with convex lower margin and a small, pointed, basal denticle at lower, outer corners (Fig. 6). Upper margin slightly concave with cutting edge bearing central, elongate conical cusp and two to three stout, conical denticles on each side. Lateral teeth with prominent triangular cusp flanked by two to three inner and two to three outer, triangular denticles, and frequently a membranous outermost denticle. Marginal teeth with broadly rounded cutting edges and long, slender, flattened shafts with membranous flanges along inner and outer edges. Narrow inner flanges along distal third to half of shafts; outer flanges extending almost to tooth bases, narrow on inner marginal teeth, broadening towards cutting edges on outer marginal teeth. Inner marginal teeth with five to six, and outer marginal teeth with six to seven flattened denticles.

Distribution and ecology

In small creeks to large rivers primarily west of the Cascades from the Olympic Peninsula of Washington in the north (Branson 1977) to the Willamette River drainage in south-central Oregon (Fig. 7A).

Remarks

Frest & Johannes (2010) refined the type locality of M. plicifera to the Multnomah Channel, which is indicated on some early maps as the second mouth of the ‘Wahlamat’ or Multnomah River. The terminal portion of the Willamette River near its confluence with the Columbia is also the type locality of Goniobasis plicifera var. oregonensis . Near topotypic specimens have been sequenced from Waterhouse Creek, Tualatin Hills, Washington County (Strong & Whelan 2019) (Fig. 5R–S), and were conspecific with specimens from nearby Johnson Creek, Multnomah County (Campbell et al. 2016), which may have produced Henderson’s type specimens of Goniobasis hemphilli (Frest & Johannes 2010) . Sequalitchew Creek is considered a possible source of Gould’s type material of Melania silicula, a Puget Sound tributary which flowed just to the north of the site of Fort Nisqually between 1833 and 1843, the Hudson’s Bay Co. outpost on the shores of Puget Sound. It was the only settlement in the area at the time and was the first port of call for the U.S. Exploring Expedition, which was the source of Gould’s types (Strong & Frest 2007; Frest & Johannes 2010). Juga no longer occurs in Sequalitchew Creek, but near topotypic specimens from McAllister Creek (Campbell et al. 2016) and the Nisqually River (Strong & Whelan 2019) (Fig. 5A) also placed in this clade. Melania silicula was synonymized with plicifera by Pilsbry (1899), a view not followed by later authors. In earlier treatments, silicula was considered widespread in Washington and Oregon (e.g., Goodrich 1942; Burch & Tottenham 1980, Burch 1982, 1989), but more recently was considered a narrow-range endemic of southeastern Puget Sound (Strong & Frest 2007; Frest & Johannes 2010; Johannes 2010b). Juga hemphilli hemphilli and J. h. dallesensis were regarded as species incerta by Goodrich (1942) but recognized as valid by Burch & Tottenham (1980) and Burch (1982, 1989). Campbell et al. (2016) noted the low genetic divergence of dallesensis from the nominotypical form but did not address their taxonomic status apart from concluding the small morphological differences in the latter could represent individual variation. Frest & Johannes (2010) also considered both as valid, but the inclusion of the name J. h. dallesensis in a heading indicated that they had apparently considered it a possible synonym of J. (O.) bulbosa (Frest & Johannes 2010: 26) . Juga hemphilli was recognized without subdivision by Turgeon et al. (1988, 1998) and Johnson et al. (2013). Thus, the synonymy of these two nominal species with plicifera here is new.

Melania rudens Reeve, 1860 was established without a locality. Tryon (1864) synonymized it with Goniobasis silicula but was later (Tryon 1865: 239) convinced of its validity and considered the species to range from Oregon to California in the Columbia and Sacramento Rivers. Goniobasis silicula rudens was recognized as valid by Henderson (1935a, 1936b) as a form from the Olympic Peninsula. However, most authors have synonymized it with silicula (e.g., Tryon 1873; Henderson 1929; Goodrich 1942; Burch & Tottenham 1980; Burch 1982, 1989) except Pilsbry (1899) who synonymized it with plicifera . Frest & Johannes (2010) considered Juga (Juga) rudens a “dubious taxon”, probably a synonym of J. silicula, and based on the original illustration of Reeve (1860: pl. 33 fig. 224) found the morphology closely comparable to that of specimens from the Nisqually River watershed and nearby tributaries of Puget Sound, the source of the types of Melania silicula . We agree and retain it here.

Tryon (1865) established the varieties Goniobasis plicifera var. bulimoides and G. p. var. oregonensis, distinguished by the length of the shell and the density of revolving striae. He abandoned them soon thereafter, omitting them as either varieties or synonyms (Tryon 1866, 1873). Henderson (1935a, 1936b: 273) acknowledged the “interminably intergrading varieties” and their rather arbitrary separation, yet inexplicably found the names useful, “in order to keep our ideas somewhat clear”. Apart from Goodrich (1942) who synonymized the two with G. plicifera, other 20 th century authors have ignored the names. Frest & Johannes (2010) thought Juga (Juga) plicifera bulimoides could be a valid large, slow river form, and J. (J.) p. oregonensis also likely to be valid. Campbell et al. (2016) noted the low genetic divergence of bulimoides and oregonensis from the nominotypical form and supported the synonymy of J. bulimoides with plicifera but did not address the taxonomic status of J. (J.) p. oregonensis, apart from concluding the small morphological differences could represent individual variation. Analysis of near topotypic specimens consistent with the morphology of the types (Strong& Whelan 2019) indicates that both subspecies represent morphological variation within the range of plicifera .

Frest & Johannes’ (1993, 1995a, 2010) concept of Juga (Juga) draytonii [= “ J. (J.) n. sp. 1 (Brown Juga)”], and possibly of “ J. (J.) hemphilli n. subsp. ” and “ J. (J.) n. sp. 3 (Tall Juga)” are composite. All three occur in the central and western Columbia Gorge in both Oregon and Washington, including the Mt. Hood National Forest, and would include individuals of both J. plicifera and J. bulbosa (Gould, 1847) as circumscribed here. The specimen sequenced as J. (J.) draytonii by Campbell et al. (2016) placed in the same clade with J. (J.) plicifera plicifera; however, the morphology of the type material of draytonii is more consistent with the interpretation that it is a synonym of J. bulbosa (see account below).

Distribution

Several historical records suggest that J. plicifera may once have occurred, at least sporadically, in the Columbia River drainage of southeastern Washington. Two of these date from the late 1800’s and early 1900’s [Walla Walla (USNM 119156; Lea Collection, Lady K. Douglas) and Wallula (USNM 109897; U.S. Fish Commission)] and their provenance is unreliable (Carpenter 1857: 162).Another lot collected in 1958 (USNM 795562; identified as J. hemphilli) places the species in a spring alongside the Snake River, a Columbia River tributary, in Whitman County of far eastern Washington, which lies far to the east of the known distribution. Moreover, this lot contains only preserved soft parts lacking shells, so the identification cannot be confirmed. However, an iNaturalist record from 2019 east of Pendleton, Oregon, shows a plicate specimen that appears to be this species (https://www.inaturalist.org/observations/28180434) and demonstrates the species extends farther east than museum records suggest. Given the non-contiguous distribution of Juga in eastern Oregon and western Nevada, the occurrence of Juga at these distant sites should not be immediately discounted (Frest & Johannes 2010: 18), but we do not include them on our distribution map until they can be confirmed in modern surveys.

Common name

The common name for this species is the Pleated Juga (Turgeon et al. 1988, 1998; Johnson et al. 2013; Johannes 2015). It has also been referred to as the Plicate Juga (Frest & Johannes 2001, 2010). Synonyms have been referred to as the Glass or Glassy Juga ( J. silicula), the Barren Juga ( J. hemphilli hemphilli), Dalles Juga ( J. hemphilli dallesensis), and the Washougal Juga ( J. (J.) hemphilli n. subsp. 1) (Turgeon et al. 1988, 1998; Frest & Johannes 1993, 1995a, 2001, 2010; Duncan 2008; Johannes 2010b, 2015; Johnson et al. 2013). Also conspecific with J. plicifera is the Indian Ford Juga [= Juga (Juga) hemphilli n. subsp. 2], known from Indian Ford Creek near Black Butte in the Deschutes National Forest, Deschutes County, Oregon (Frest & Johannes 1995a; Duncan 2008; Johannes 2015). This corresponds to the site “just downstream of U.S. Hwy 20 (FS campground)” in Strong & Whelan (2019) (Fig. 5Z) and represents an unusual incursion of J. plicifera into the Deschutes River system; it is also the farthest occurrence of Juga up the Deschutes basin (Johannes 2015).