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“ Juga maluccinosa Quoy ” One of the species originally included in the subgenus Juga by Adams & Adams (1854), this was listed as an extralimital cerithioidean by Frest & Johannes (2010) with no additional information. This is an incorrect subsequent spelling by Adams & Adams (1854) for Melania moluccensis Quoy & Gaimard, 1834, and is a synonym of Stenomelania erosa (Lesson, 1831) (Glaubrecht & Podlacha 2010)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 81, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Glaubrecht M. & Podlacha K. 2010. Freshwater gastropods from early voyages into the Indo-West Pacific: the ' melaniids' (Cerithioidea, Thiaridae) from the French ' La Coquille' circumnavigation, 1822 - 1825. Zoosystematics and Evolution 86: 185 - 211. https: // doi. org / 10.1002 / zoos. 201000002"]}

Anculotus nuttalii Reeve, 1861 Described from “ Oregon ”, Hershler & Frest (1996) considered this to be an incorrect subsequent spelling of nuttaliana Lea, 1838. Reeve attributed the name Anculotus nuttalii to “Lea MS”, and did not refer to Paludina nuttaliana I. Lea, 1838. The name is thus neither an incorrect subsequent spelling nor an emendation, but a newly established nominal species founded on its own type material. It was included by Graf (2001), who did not examine the type material. We follow Hershler & Frest (1996) in placing it in the synonymy of Fluminicola nuttalianus (Lithoglyphidae Tryon, 1866). Anculotus Conrad, 1834 is an incorrect subsequent spelling of Anculosa. This species was omitted by Frest & Johannes (2010)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 81, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Hershler R. & Frest T. J. 1996. A review of the North American freshwater snail genus Fluminicola (Hydrobiidae). Smithsonian Contributions to Zoology 583: 1 - 41. https: // doi. org / 10.5479 / si. 00810282.583","Lea I. 1838. Description of new freshwater and land shells. Transactions of the American Philosophical Society 6: 1 - 154. [Observations on the genus Unio, etc., 2: 1 - 152.] https: // doi. org / 10.2307 / 1005319","Graf D. L. 2001. The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of Recent North America, north of Mexico. Walkerana 12: 1 - 124.","Tryon G. W. 1866. Monograph of the family Strepomatidae. American Journal of Conchology 2: 14 - 52.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61."]}

Genus Juga H. Adams & A. Adams, 1854 Vibex (Juga) H. & A. Adams, 1854. Type species: Melania silicula A. Gould, 1847, by subsequent designation (Baker 1963: 35). Juga (Calibasis) D.W. Taylor, 1966. Type species: Melania (? Goniobasis) acutifilosa Stearns, 1890, by original designation. Syn. nov. Juga (Oreobasis) D.W. Taylor, 1966. Type species: Melania newberryi I. Lea, 1860, by original designation. Juga (Idabasis) D.W. Taylor, 1966 †. Type species: Juga chrysopylica D.W. Taylor, 1966 †, by original designation. Syn. nov. Diagnosis Shell dextral, smooth to strongly plicate, lirate or cancellate, medium to large in size, ~ 1.5–3.5 cm in length. Operculum corneous, brown, paucispiral with eccentric nucleus. Gonochoristic, oviparous, with broad, shallow, triangular ovipositor pore; convoluted gonoductal groove, proximal albumen gland u-shaped; seminal receptacle present; tightly interlocking folds of lateral and medial laminae of prostate gland; gelatinous, finger-like egg masses with large numbers of eggs. Radula taenioglossate, rachidian basal cusps weakly developed or lacking. Midgut with comparatively narrow glandular pad and wide, shallow, crescentic groove; digestive gland duct vestibule receiving several ducts of digestive gland. Remarks Until recently, extant species of Juga were distributed among three subgenera, based on the sculpture of the early whorls (Taylor 1966): Juga s. str. with strong and sometimes shouldered plications or ribs, Calibasis with spiral or lirate sculpture sometimes giving a frilled appearance, and Oreobasis with essentially smooth early whorls. Taylor (1966) also recognized a fourth subgenus with cancellate early sculpture, Idabasis, based on the Blancan fossil, Juga chrysopylica Taylor, 1966. According to Frest & Johannes (2010: 7), Taylor further subdivided Oreobasis into two sections in an unpublished checklist dated from 1977 which we have not seen. Strong & Frest (2007) synonymized Oreobasis with Juga s. str. given their morphological similarity in reproductive and alimentary anatomy. Campbell et al. (2016) concluded that none of the three extant subgenera were monophyletic but that clades of equivalent rank merited recognition. The molecular analysis of Strong & Whelan (2019) confirmed that the strength and persistence of spiral and axial sculpture are variable within and between species, sometimes among conspecific individuals at a single site, and that subgeneric designations based primarily on shell sculpture are not meaningful. Moreover, the molecular phylogeny did not reveal clades that would merit recognition at the rank of subgenus. Thus, we here further synonymize Calibasis and Idabasis with Juga., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 10, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Gould A. A. 1847. Descriptions of the following species of Melania, from the collection of the Exploring Expedition. Proceedings of the Boston Society of Natural History 2: 222 - 225. https: // doi. org / 10.5962 / bhl. part. 3322","Baker H. B. 1963. Paludomidae (Pleuroceridae). The Nautilus 77: 34 - 35.","Taylor D. W. 1966. Summary of North American Blancan nonmarine mollusks. Malacologia 4: 1 - 172.","Stearns R. E. C. 1890. Scientific results of explorations by the U. S. Fish Commission steamer Albatross. XVII. Descriptions of new West American land, freshwater, and marine shells, with notes and comments. Proceedings of the United States National Museum 13: 205 - 225. https: // doi. org / 10.5479 / si. 00963801.13 - 813.205","Lea I. 1860. Descriptions of four new species of Melanidae of the United States. Proceedings of the Academy of Natural Sciences of Philadelphia 12: 93. Available from https: // www. biodiversitylibrary. org / page / 1801109 [accessed 13 Mar. 2022].","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Strong E. E. & Frest T. 2007. On the anatomy and systematics of Juga from western North America (Gastropoda: Cerithioidea: Pleuroceridae). The Nautilus 121: 43 - 65.","Campbell D. C., Clark S. A., Johannes E. J., Lydeard C. & Frest T. J. 2016. Molecular phylogenetics of the freshwater gastropod genus Juga (Cerithioidea: Semisulcospiridae). Biochemical Systematics and Ecology 65: 158 - 170. https: // doi. org / 10.1016 / j. bse. 2016.01.004","Strong E. E. & Whelan N. V. 2019. Assessing the diversity of Western North American Juga (Semisulcospiridae, Gastropoda). Molecular Phylogenetics and Evolution 136: 87 - 103. https: // doi. org / 10.1016 / j. ympev. 2019.04.009"]}

“ Juga schiedeana Phillipi ” (sic, Philippi) One of the species originally included in the subgenus Juga by Adams & Adams (1854), this was listed by Frest & Johannes (2010) as an extralimital cerithioidean with no additional information. It was established as Melania schiedeana Philippi, 1843 from Mexico and is currently placed in the Pachychilidae as Pachychilus schiedeanus (Thompson 2011)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 83, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Thompson F. G. 2011. An annotated checklist and bibliography of the land and freshwater snails of Mexico and Central America. Florida Museum of Natural History Bulletin 50: 1 - 299."]}

Juga occata (Hinds, 1844) Figs 25–26 Melania occata Hinds, 1844: 9. Goniobasis acutifilosa var. siskiyouensis Pilsbry, 1899: 65–66. Goniobasis acutifilosa pittensis Henderson, 1935a: 134, pl. 4 fig. 11. Melania occata – Hinds 1845: 56–57, pl. 15 fig. 5. Melania shastaensis – Reeve 1860: unpaginated, species 318, pl. 44 fig. 318. Goniobasis acutifilosa pittensis – Henderson 1936b, pl. 2 fig. 11. Juga (Calibasis) occata – Burch & Tottenham 1980: 152, fig. 451. — Burch 1982: 41, fig. 451; 1989: 152, fig. 451. — Frest & Johannes 1993: 61; 1995b: 38; 2010: 10, 44, fig. 2e. — Furnish 2007: 16, fig. 6, above right. — Campbell et al. 2016: 160. Juga (Calibasis) acutifilosa siskiyouensis – Burch & Tottenham 1980: 152. — Burch 1982: 41; 1989: 152. Juga (Calibasis) acutifilosa pittensis – Burch & Tottenham 1980: 152. — Burch 1982: 41; 1989: 152. Juga occata – Turgeon et al. 1988: 65; 1998: 67. — Johannes & Clark 2016: 24. — Johnson et al. 2013: 282. — Strong & Whelan 2019: 89, fig. 4m –q. Juga (Calibasis) acutifilosa – Frest & Johannes 1993: 61 (in part); 1995 b: 37 (in part); 2000a: 283 (in part); 2005: 160 (in part); 2010: 9, 22 (in part). — Furnish 2007: 14, fig. 6, above left. Juga (Calibasis) n. sp. 1 – Frest & Johannes 1995b: 38 (in part). Juga (Calibasis) OTU 3 – Campbell et al. 2016: 160. Juga (Calibasis) OTU 4 – Campbell et al. 2016: 160. Juga (Calibasis) OTU 5 – Campbell et al. 2016: 160. Material examined Melania occata Hinds, 1844 USA • 1 spm (possible syntype, Fig. 25A); “ River Sacramento, California ”; R.B. Hinds leg.; NHMUK 4.9.23.34. Lectotype of Goniobasis acutifilosa var. siskiyouensis Pilsbry, 1899 (designated by Baker 1964: 187) USA • “ Headwaters of Fall River, S.-E. [southeast] Siskiyou Co. ”; R.C. McGregor leg.; ANSP 73508. Paralectotypes of Goniobasis acutifilosa var. siskiyouensis Pilsbry, 1899 (Fig. 25B) USA • 90 spms; same collection data as for lectotype; ANSP 466291 (ex ANSP 73508). Holotype of Goniobasis acutifilosa pittensis Henderson, 1935 (Fig. 25C) USA • “ Fall River Mills, on Pitt [sic, Pit] River, California ”; Ellsworth Bethel leg.; UCM 12080 a. Paratypes of Goniobasis acutifilosa pittensis Henderson, 1935 USA • 2 spms; same collection data as for holotype; UCM 12080b. Other material examined 48 lots, 898 specimens, of which 21 were sequenced. USA – California • 5 spms; Lassen Co., Davis Creek, Lassen NF, above FS22; 40.8325, -121.1183; 1 Sep. 2007; T. Grace leg.; UF 520312 • 54 spms; Lassen Co., Lassen Natl Forest, Davis Creek, on FS Rd 22; [40.8375, -121.1183]; 5 Jun. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; CASIZ 166015 • 39 spms; Lassen Co., Lassen Natl Forest, Davis Creek, on FS Rd 22 [= FS Rd 35N05]; [40.8375, -121.1183]; 5 Jun. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; CASIZ 165958 • 199 spms; Lassen Co., Lassen Natl Forest, Russell Dairy Spring, on 35N85 [= FS Rd 35N05]; [40.8277, -121.0281]; 2001; USFS Lassen Natl Forest Mollusk Survey 2001; CASIZ 166008 • 35 spms; Lassen Co., Russell Dairy Spring Run, Lassen NF, FS22 and 34N2, S side of road; 40.8253, -121.0268; 1 Sep. 2007; T. Grace leg.; UF 520309 • 12 spms; Modoc Co., Goose Lake; [41.9067, -120.3652]; 26 Sep. 1979; CASIZ 85267 • 2 spms; Modoc Co., Modoc Natl Forest, W end of Upper Rush Creek Campground, 3.2 km N of Lower Rush Creek Campground; [41.3015, -120.8461]; 8 Oct. 2007; T. Grace leg.; small spring run, among water cress and on rocks and twigs; UF 548146 • 1 spm; Sacramento Co., Sacramento; [38.5833, -121.5064]; Wetherby leg.; USNM 118914 • 2 spms; Shasta Co., Battle Creek, S boundary of county; [40.3921, -122.1787]; 1898; R.C. McGregor leg.; ANSP 73521 • 49 spms; Shasta Co., Baum Lake; 40.9345, -121.5485; 10 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413151 • 1 spm; same collection data as for preceding; GenBank: MK472482, MK464849, MK480916; USNM 1413146 • 1 spm; same collection data as for preceding; GenBank: MK472483, MK464850, MK480917; USNM 1413147 • 1 spm; same collection data as for preceding; GenBank: MK472484, MK464851, MK480918; USNM 1413148 • 1 spm; same collection data as for preceding; GenBank: MK472485, MK464852, MK480919; USNM 1413149 • 1 spm; same collection data as for preceding; GenBank: MK472486, MK464853, MK480920; USNM 1413150 • 34 spms; Shasta Co., Fall River at Spinner Fall Lodge; 41.0972, -121.5491; 11 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413084 • 1 spm; same collection data as for preceding; GenBank: MK472471, MK464838, MK480906; USNM 1413079 • 1 spm; same collection data as for preceding; GenBank: MK472472, MK464839, MK480907; USNM 1413080 • 1 spm; same collection data as for preceding; GenBank: MK472473, MK464840, MK480908; USNM 1413081 • 1 spm; same collection data as for preceding; GenBank: MK472474, MK464841, MK480909; USNM 1413082 • 1 spm; same collection data as for preceding; GenBank: MK472475, MK464842, MK480910; USNM 1413083 • 29 spms; Shasta Co., Hat Creek Park, Hat Creek, E of Burney at highway bridge; [40.9767, -121.5565]; 16 May 1978; J. Landye leg.; sand and emergent vegetation; USNM 892368 • 50 spms; Shasta Co., Hat Creek Park, Hat Creek, E of Burney at highway crossing; [40.9767, -121.5565]; 16 May 1978; J. Landye leg.; on sand and vegetation; USNM 892375 • 9 spms; Shasta Co., near border of Modoc County, N of McArthur, Big Lake, Crystal Springs; [41.1118, -121.4494]; 16 Jul. 1980; CASIZ 85274 • 55 spms; Shasta Co., Pit River at U.S. Hwy 299; 40.9807, -121.5473; 11 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413091 • 1 spm; same collection data as for preceding; GenBank: MK472476, MK464843, MK480911; USNM 1413085 • 1 spm; same collection data as for preceding; GenBank: MK472477, MK464844, MK480912; USNM 1413086 • 1 spm; same collection data as for preceding; GenBank: MK472478, MK464845; USNM 1413087 • 1 spm; same collection data as for preceding; GenBank: MK472479, MK464846, MK480913; USNM 1413088 • 1 spm; same collection data as for preceding; GenBank: MK472480, MK464847, MK480914; USNM 1413089 • 1 spm; same collection data as for preceding; GenBank: MK472481, MK464848, MK480915; USNM 1413090 • 57 spms; Shasta Co., Pit River, above Lake Britton and upstream from Hat Creek; [40.9805, -121.5476]; 16 May 1978; J. Landye leg.; in pockets along rocky shore in eddies; USNM 892363 • 16 spms; Shasta Co., Rainbow Springs, at headwaters of Fall Mill Creek on Lazy S Ranch adjacent to Thousand Springs Ranch; [41.1111, -121.5511]; 16 May 1978; J. Landye leg.; USNM 892373 • 75 spms; Shasta Co., Rainbow Springs, in headspring on Lazy S Ranch in Thousand Springs Area; [41.1111, -121.5511]; 16 May 1978; J. Landye leg.; on rocks; USNM 892380 • 15 spms; Shasta Co., S side of Hwy 299, Pit River, Lion’s Club picnic area, W of Pit River camping grounds; 40.9866, -121.513; 1 Sep. 2007; T. Grace leg.; USNM 1111864 • 2 spms; Shasta Co., S side of Pit River, water source spring at Pacific Gas and Electric plant; [40.9896, -121.5021]; 16 May 1978; J. Landye leg.; USNM 892372 • 58 spms; Shasta Co., S side of Pit River, water source spring at Pacific Gas and Electric Plant #1; [40.9896, -121.5021]; 16 May 1978; J. Landye leg.; USNM 892365 • 46 spms; Shasta Co., Spring Creek at Spring Creek Rd; 41.1018, -121.5189; 11 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413164 • 1 spm; same collection data as for preceding; GenBank: MK472487, MK464854, MK480921; USNM 1413159 • 1 spm; same collection data as for preceding; GenBank: MK472488, MK464855; USNM 1413160 • 1 spm; same collection data as for preceding; GenBank: MK472489, MK464856, MK480922; USNM 1413161 • 1 spm; same collection data as for preceding; GenBank: MK472490, MK464857, MK480923; USNM 1413162 • 1 spm; same collection data as for preceding; GenBank: MK472491, MK464858; USNM 1413163 • 18 spms; Sacramento River; Trask leg.; USNM 119157 • 2 spms; same collection data as for preceding; U.S. Exploring Expedition; USNM 16251 • 7 spms; Upper Sacramento; USNM 58982 • 3 spms; Sacramento River, headwaters [sic, likely mislocalized]; USNM 56463. – Oregon • 3 spms; Clatsop Co., Astoria [sic, mislocalized]; White leg.; USNM 126074. Description SHELL. Thin to moderately thick, large, reaching ~ 2.75 cm in length; turriform, narrowly or broadly conical to cylindrical in shape, spire height moderate to tall (Fig. 25). Whorls somewhat flattened to convex or angulate with a subsutural ramp, suture weakly to deeply impressed. Aperture oval in shape, lip smooth to crenate, simple to slightly sinuous. Spiral sculpture absent or present, of thickened, scalloped lirae, barely elevated to prominent, highly variable in number and sinuosity, dividing whorl into intervening flattened areas or grooves. Axial sculpture of fine, weakly opisthocyrt to sinuous growth lines; plications lacking. Shell dark to reddish brown, to yellowish green in color, occasionally with a lighter subsutural band; bands otherwise lacking. Interior of aperture light brown to white in color. RADULA. Rachidian squarish, slightly wider than tall, with weakly v-shaped lower margin and projecting outer corners; basal denticles lacking or vestigial (Fig. 26). 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 large, 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 quarter of inner marginal teeth and distal two-thirds of outer marginal teeth; broad outer flanges extending almost to tooth bases. Inner marginal teeth with three to four and outer marginal teeth with four to five flattened denticles. Distribution and ecology In spring-fed creeks to large rivers and lakes, currently restricted to the Pit River system in Lassen, Shasta, and Modoc Counties of northern California (Fig. 21C). Remarks Taylor (1981) emended the type locality of Melania occata to “Sacramento River, California (between American River and mouth)” given that the types were collected during the voyage of the HMS Sulphur which traveled no further upstream than the mouth of the American River. Museum records (e.g., UMMZ 40420, UMMZ 134037) indicate that the species once inhabited the Sacramento River and the lower part of the San Joaquin River watershed to Antioch, California (Henderson 1935a: 96), but was extirpated in the 19 th century from the lower Sacramento River system owing to the effects of mining, pollution, and impoundment (Taylor 1981; Frest & Johannes 1993, 2010; Furnish 2007). Only three river miles of the Upper Sacramento River were unaffected by the Cantara chemical spill in 1991, but surveys found it already extirpated there as well; there are no museum records from the Upper Sacramento River since at least 1940 (Frest & Johannes 1993). The type locality of G. a. var. siskiyouensis was indicated to be the headwaters of the Fall River in southeastern Siskiyou County. However, the Fall River is considered to originate at Thousand Springs in northeastern Shasta County. Bear Creek is a major tributary that joins the river near its origin and flows from southeastern Siskiyou County and may have been the source of the type material. Attempts to collect from Bear Creek by the authors were unsuccessful. The type locality of G. a. pittensis at the confluence of the Fall and Pit Rivers is now impounded, heavily polluted, and eutrophic from agricultural runoff and Juga no longer occurs there. Juga occata has been considered a robust, heavily sculptured, large river form (Fig. 25D–F) (Burch & Tottenham 1980; Taylor 1981; Burch 1982, 1989; Frest & Johannes 1993, 2010), but molecular analyses indicate that this form is conspecific with less heavily sculptured populations that occur in lakes and spring-fed creeks (Strong & Whelan 2019). This includes near topotypic specimens (Fig. 25H–J) of Goniobasis acutifilosa var. siskiyouensis and G. a. pittensis. The classifications of Burch & Tottenham (1980) and Burch (1982, 1989) recognized these two subspecies as valid, but recent classifications (Turgeon et al. 1988, 1998; Johnson et al. 2013) did not consider the rank of subspecies. Frest & Johannes (2010) followed Taylor (1981) in synonymizing siskiyouensis with J. acutifilosa, and pittensis with J. occata, while Campbell et al. (2016: supp. file 1) synonymized both with occata. Distribution The main part of the distribution of Juga occata lies in southern Pit River drainages in Shasta and Lassen Counties where it overlaps with J. douglasi sp. nov., although individuals of the two species are rarely found in syntopy; only three lots with mixtures of the two were found in the USNM collections, and one species is always numerically dominant [USNM 1111864, USNM 1665623 (ex USNM 1111864); USNM 892372, USNM 1665624 (ex USNM 892372); USNM 892373, USNM 1665622 (ex USNM 892373)]. In contrast to J. douglasi, juveniles of occata are lirate, although not always prominently so, and erosion frequently obscures details of the spire; the spire angle in occata is also generally broader and the suture less impressed (Fig. 25X –D’). Sequenced specimens from two sites sampled by Campbell et al. (2016) and lirate museum specimens collected nearby (UF 520309, UF 520312, CASIZ 165958, CASIZ 166008, CASIZ 166015) confirm that the current range of occata extends as far south as central western Lassen County. If reliable, one historical lot (ANSP 73521) collected in the late 19 th century from Battle Creek along the southern border of Shasta County indicates that the well-sculptured form also extended into smaller tributaries of the Sacramento River. Most historical lots from the lower Sacramento and San Joaquin Rivers, including the types, are poorly localized specimens of the typical, sculptured, large river form. We could find no evidence of early teleoconchs with the characteristic lirae south of the Pit watershed and, thus, records in the Sierra Nevada are tentatively hypothesized to represent Juga douglasi. However, given the range of morphological variation seen in species of Juga, it is possible that populations from the Sierra Nevada represent mixtures of J. occata and J. douglasi, or an as yet undescribed form. Common name The common name for this species is the Topaz Juga (Frest & Johannes 1993), erroneously referred to as the Scalloped Juga by Turgeon et al. (1988, 1998), which was followed by Frest & Johannes (2010) and Johnson et al. (2013) (see J. acutifilosa, above). A composite species comprised of J. acutifilosa and J. occata as circumscribed herein has been referred to as the Willow Creek Juga (= Juga (Calibasis) n. sp. 1; Frest & Johannes 1995b)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on pages 74-80, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Hinds R. B. 1844. Descriptions of new species of Melania collected during the voyage of H. M. S. Sulphur. The Annals and Magazine of Natural History 14: 8 - 11. https: // doi. org / 10.1080 / 037454809495124","Pilsbry H. A. 1899. Mollusks collected by R. C. McGregor in northern California. The Nautilus 13: 64 - 67.","Henderson J. 1935 a. West American species of Goniobasis, with a description of new forms. The Nautilus 48: 94 - 99, 130 - 134.","Henderson J. 1936 b. The non-marine Mollusca of Oregon and Washington. Supplement. University of Colorado Studies 23: 251 - 280.","Burch J. B. & Tottenham J. 1980. North American freshwater snails, species list, ranges, and illustrations. Walkerana 3: 1 - 215.","Burch J. B. 1982. Freshwater snails (Mollusca: Gastropoda) of North America. Environmental Monitoring and Support Laboratory, Office of Research and Development, United States Environmental Protection Agency, Cincinnati, Ohio, EPA- 600 / 3 - 82 - 026.","Burch J. B. 1989. North American Freshwater Snails. Malacological Publications, Hamburg, Michigan.","Frest T. J. & Johannes E. J. 1993. Mollusc species of Special Concern within the range of the Northern Spotted Owl. Final report to Forest Ecosystem Management Working Group, USDA Forest Service. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 1995 b. Freshwater mollusks of the Upper Sacramento System, California, with particular reference to the Cantara Spill. 1994 yearly report to California Department of Fish and Game. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Furnish J. L. 2007. Guide to sensitive aquatic mollusks of the U. S. Forest Service Pacific Southwest Region. USDA Forest Service, Pacific Southwest Region, Vallejo, CA.","Campbell D. C., Clark S. A., Johannes E. J., Lydeard C. & Frest T. J. 2016. Molecular phylogenetics of the freshwater gastropod genus Juga (Cerithioidea: Semisulcospiridae). Biochemical Systematics and Ecology 65: 158 - 170. https: // doi. org / 10.1016 / j. bse. 2016.01.004","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1988. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 26.","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1998. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. Second Edition. American Fisheries Society Special Publication 26.","Johannes E. J. & Clark S. A. 2016. Freshwater mollusc declines, local extinctions and introductions in five northern California streams. Tentacle 24: 22 - 25.","Johnson P. D., Bogan A. E., Brown K. M., Burkhead N. M., Cordeiro J. R., Garner J. T., Hartfield P. D., Lepitzki D. A. W., Mackie G. L., Pip E., Tarpley T. A., Tiemann J. S., Whelan N. V. & Strong E. E. 2013. Conservation status of freshwater gastropods of Canada and the United States. Fisheries 38: 247 - 282. https: // doi. org / 10.1080 / 03632415.2013.785396","Strong E. E. & Whelan N. V. 2019. Assessing the diversity of Western North American Juga (Semisulcospiridae, Gastropoda). Molecular Phylogenetics and Evolution 136: 87 - 103. https: // doi. org / 10.1016 / j. ympev. 2019.04.009","Frest T. J. & Johannes E. J. 1995 a. Interior Columbia Basin mollusk species of special concern. Final report: Interior Columbia Basin Ecosystem Management Project, Walla Walla, WA. Contract 43 - 0 E 00 - 4 - 9112.","Frest T. J. & Johannes E. J. 2000 a. A baseline mollusk survey of southwestern Oregon, with emphasis on the Rogue and Umpqua River drainages. Final report prepared for Oregon Natural Heritage Program. Portland, Oregon. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 2005. Springsnails of the Cascade-Siskiyou National Monument and vicinity, Oregon. 2004 report. Final report prepared for World Wildlife Fund, Ashland, Oregon. Deixis Consultants, Seattle, Washington.","Baker H. B. 1964. Type land snails in the Academy of Natural Sciences of Philadelphia part III. Limnophile and thalassophile Pulmonata Part IV. Land and fresh-water Prosobranchia. Proceedings of the Academy of Natural Sciences of Philadelphia 116: 149 - 193.","Taylor D. W. 1981. Freshwater mollusks of California: a distributional checklist. California Fish and Game 67: 140 - 163."]}

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; Gen, Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on pages 11-26, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Lea I. 1838. Description of new freshwater and land shells. Transactions of the American Philosophical Society 6: 1 - 154. [Observations on the genus Unio, etc., 2: 1 - 152.] https: // doi. org / 10.2307 / 1005319","Gould A. A. 1847. Descriptions of the following species of Melania, from the collection of the Exploring Expedition. Proceedings of the Boston Society of Natural History 2: 222 - 225. https: // doi. org / 10.5962 / bhl. part. 3322","Tryon G. W. 1865. Review of the goniobases of Oregon and California. American Journal of Conchology 1: 236 - 246.","Henderson J. 1935 a. West American species of Goniobasis, with a description of new forms. The Nautilus 48: 94 - 99, 130 - 134.","Gould A. A. 1852. Mollusca and shells. Volume 12. In: United States Exploring Expedition during the Years 1838, 1839, 1840, 1841, 1842. Under the Command of Charles Wilkes, U. S. N. Gould and Lincoln, Boston. https: // doi. org / 10.5962 / bhl. title. 69333","Gould A. A. 1856 [15 December 1860]. Atlas. Mollusca and shells. In: United States Exploring Expedition during the Years 1838, 1839, 1840, 1841, 1842. Under the Command of Charles Wilkes, U. S. N. Gould and Lincoln, Boston. https: // doi. org / 10.5962 / bhl. title. 69333","Henderson J. 1936 b. The non-marine Mollusca of Oregon and Washington. Supplement. University of Colorado Studies 23: 251 - 280.","Burch J. B. & Tottenham J. 1980. North American freshwater snails, species list, ranges, and illustrations. Walkerana 3: 1 - 215.","Burch J. B. 1982. Freshwater snails (Mollusca: Gastropoda) of North America. Environmental Monitoring and Support Laboratory, Office of Research and Development, United States Environmental Protection Agency, Cincinnati, Ohio, EPA- 600 / 3 - 82 - 026.","Holznagel W. E. & Lydeard C. 2000. A molecular phylogeny of North American Pleuroceridae (Gastropoda: Cerithioidea) based on mitochondrial 16 S rDNA sequences. Journal of Molluscan Studies 66: 233 - 257. https: // doi. org / 10.1093 / mollus / 66.2.233","Burch J. B. 1989. North American Freshwater Snails. Malacological Publications, Hamburg, Michigan.","Neitzel D. A. & Frest T. J. 1992. Survey of Columbia River Basin streams for Columbia Pebblesnail Fluminicola columbiana and Shortface Lanx Fisherola nuttalli. Prepared for the U. S. Department of Energy under contract DE-AC 06 - 76 RLO 1830. Battelle Memorial Institute, Pacific Northwest Laboratories, Richland, Washington, PNL- 8229. https: // doi. org / 10.2172 / 10173267","Strong E. E. & Frest T. 2007. On the anatomy and systematics of Juga from western North America (Gastropoda: Cerithioidea: Pleuroceridae). The Nautilus 121: 43 - 65.","Johannes E. J. 2010 b. Freshwater mollusks found during a survey for Potamopyrgus antipodarum (New Zealand mudsnail) within a five-mile radius of Capitol Lake, Thurston County, Washington. The Dredgings 50: 3 - 5.","Frest T. J. & Johannes E. J. 1993. Mollusc species of Special Concern within the range of the Northern Spotted Owl. Final report to Forest Ecosystem Management Working Group, USDA Forest Service. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 1995 a. Interior Columbia Basin mollusk species of special concern. Final report: Interior Columbia Basin Ecosystem Management Project, Walla Walla, WA. Contract 43 - 0 E 00 - 4 - 9112.","Frest T. J. & Johannes E. J. 2001. Baseline freshwater mollusk survey of Mount Rainier National Park. 2000 yearly report. Final report prepared for U. S. Department of the Interior, National Park Service, Mount Rainier National Park, Ashland, Washington. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Campbell D. C., Clark S. A., Johannes E. J., Lydeard C. & Frest T. J. 2016. Molecular phylogenetics of the freshwater gastropod genus Juga (Cerithioidea: Semisulcospiridae). Biochemical Systematics and Ecology 65: 158 - 170. https: // doi. org / 10.1016 / j. bse. 2016.01.004","Hawkins C. P. & Furnish J. L. 1987. Are snails important competitors in stream ecosystems? Oikos 49: 209 - 220. https: // doi. org / 10.2307 / 3566028","Furnish J. L. 1990. Factors affecting the growth, production and distribution of the stream snail Juga silicula (Gould). Unpublished Ph. D. thesis, Oregon State University.","Furnish J. L. 2007. Guide to sensitive aquatic mollusks of the U. S. Forest Service Pacific Southwest Region. USDA Forest Service, Pacific Southwest Region, Vallejo, CA.","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1988. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 26.","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1998. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. Second Edition. American Fisheries Society Special Publication 26.","Lee T., Kim J. J., Hong H. C., Burch J. B. & O Foighil D. 2006. Crossing the continental divide: the Columbia drainage species Juga hemphilli (Henderson, 1935) is a cryptic member of the eastern North American genus Elimia (Cerithioidea: Pleuroceridae). Journal of Molluscan Studies 72: 314 - 317. https: // doi. org / 10.1093 / mollus / eyl 005","O Foighil D., Lee T., Campbell D. C. & Clark S. A. 2009. All voucher specimens are not created equal: A cautionary tale involving North American pleurocerid gastropods. Journal of Molluscan Studies 75: 305 - 306. https: // doi. org / 10.1093 / mollus / eyp 033","Strong E. E. & Kohler F. 2009. Morphological and molecular analysis of ' Melania' jacqueti Dautzenberg and Fischer, 1906: from anonymous orphan to critical basal offshoot of the Semisulcospiridae (Gastropoda: Cerithioidea). Zoologica Scripta 38: 483 - 502. https: // doi. org / 10.1111 / j. 1463 - 6409.2008.00385. x","Strong E. E., Colgan D. J., Healy J. M., Lydeard C., Ponder W. F. & Glaubrecht M. 2011. Phylogeny of the gastropod superfamily Cerithioidea using morphology and molecules. Zoological Journal of the Linnean Society 162: 43 - 89. https: // doi. org / 10.1111 / j. 1096 - 3642.2010.00670. x","Johnson P. D., Bogan A. E., Brown K. M., Burkhead N. M., Cordeiro J. R., Garner J. T., Hartfield P. D., Lepitzki D. A. W., Mackie G. L., Pip E., Tarpley T. A., Tiemann J. S., Whelan N. V. & Strong E. E. 2013. Conservation status of freshwater gastropods of Canada and the United States. Fisheries 38: 247 - 282. https: // doi. org / 10.1080 / 03632415.2013.785396","Kohler F. 2016. Rampant taxonomic incongruence in a mitochondrial phylogeny of Semisulcospira freshwater snails from Japan (Cerithioidea: Semisulcospiridae). The Journal of Molluscan Studies 82: 268 - 281. https: // doi. org / 10.1093 / mollus / eyv 057","Johannes E. J. 2015. Potential impacts from climate change on freshwater mollusks in the Deschutes, Klamath, and John Day basins. In: Wisseman R. (ed.) Potential Impacts from Climate Change on Freshwater Invertebrates, including Mollusks, in the Deschutes, Klamath, and John Day Basins of Eastern Oregon: 19 - 37.","Lydeard C., Holznagel W. E., Glaubrecht M. & Ponder W. F. 2002. Molecular phylogeny of a circumglobal, diverse gastropod superfamily (Cerithioidea: Mollusca: Caenogastropoda): pushing the deepest phylogenetic limits of mitochondrial LSU rDNA sequences. Molecular Phylogenetics and Evolution 22: 399 - 406. https: // doi. org / 10.1006 / mpev. 2001.1072","Johannes E. J. 2010 a. Impact on Potamopyrgus antipodarum (New Zealand Mudsnail) from the 2009 drawdown of Capitol Lake, Thurston County, Washington. Final report to Washington Department of Fish and Wildlife, Olympia, Washington. Deixis Consultants, SeaTac, Washington.","Johannes E. J. 2013 b. Survey for Potamopyrgus antipodarum (New Zealand Mudsnail) in the Chehalis River Surge Plain Natural Area Preserve, Grays Harbor County, Washington. Final report for Washington State Department of Natural Resources, Pacific Cascade Region, Tumwater Work Center, WA. Deixis Consultants, SeaTac, Washington.","Strong E. E. & Whelan N. V. 2019. Assessing the diversity of Western North American Juga (Semisulcospiridae, Gastropoda). Molecular Phylogenetics and Evolution 136: 87 - 103. https: // doi. org / 10.1016 / j. ympev. 2019.04.009","Baker H. B. 1964. Type land snails in the Academy of Natural Sciences of Philadelphia part III. Limnophile and thalassophile Pulmonata Part IV. Land and fresh-water Prosobranchia. Proceedings of the Academy of Natural Sciences of Philadelphia 116: 149 - 193.","McCoy C. J. Jr. 1964. Type specimens of Recent Gastropoda in the University of Colorado Museum. University of Colorado Studies, Series in Biology 12: 1 - 7.","Branson B. A. 1977. Freshwater and terrestrial Mollusca of the Olympic Peninsula, Washington. The Veliger 19: 310 - 330.","Pilsbry H. A. 1899. Mollusks collected by R. C. McGregor in northern California. The Nautilus 13: 64 - 67.","Goodrich C. 1942. The Pleuroceridae of the Pacific coastal drainage, including the Western Interior Basin. Occasional Papers of the Museum of Zoology, University of Michigan 469: 1 - 4.","Tryon G. W. 1864. Synonymy of the species of Strepomatidae, a family of fluviatile Mollusca inhabiting North America. Part 2. Proceedings of the Academy of Natural Sciences of Philadelphia 16: 35 - 59.","Tryon G. W. 1873. Land and Fresh-water Shells of North America Part IV. Strepomatidae (American Melanians). Smithsonian Institution, Washington, D. C.","Henderson J. 1929. The non-marine Mollusca of Oregon and Washington. University of Colorado Studies 17: 47 - 190.","Tryon G. W. 1866. Monograph of the family Strepomatidae. American Journal of Conchology 2: 14 - 52.","Stearns R. E. C. 1890. Scientific results of explorations by the U. S. Fish Commission steamer Albatross. XVII. Descriptions of new West American land, freshwater, and marine shells, with notes and comments. Proceedings of the United States National Museum 13: 205 - 225. https: // doi. org / 10.5479 / si. 00963801.13 - 813.205","Carpenter P. P. 1857. Report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Report of the British Association for the Advancement of Science 1856: 159 - 368. https: // doi. org / 10.5962 / bhl. title. 60613","Duncan N. 2008. Survey Protocol for Aquatic Mollusk Species: Preliminary Inventory and Presence / Absence Sampling, Version 3.1. Portland, OR. Interagency Special Status / Sensitive Species Program. U. S. Department of Interior, Bureau of Land Management, Oregon / Washington and U. S. Department of Agriculture, Forest Service, Region 6."]}

Goniobasis rubiginosa I. Lea, 1862 This name was established by Lea (1862: 270; 1863a: 333–334, pl. 38 fig. 193; 1863b: 155–156, pl. 38 fig. 193) based on two specimens sent to him by Wesley Newcomb, M.D., from “ Oregon ”. Tryon (1864, 1865, 1866, 1873) considered the species to be valid, despite having seen specimens only from Lea’s collection, and was confident they would be plicate if the initial whorls were not eroded (Tryon 1865: 244). Pilsbry (1899) also considered the species to be valid. Henderson (1929) identified two specimens as this species in a lot of eight from Ahtanum Creek, Union Gap, in Yakima County, Washington; the other specimens in the lot were identified as Goniobasis draytonii. Henderson (1936b: 274) later admitted that one of the specimens was quite young, and in the other the carina was evidently the “result of disease or injury”. As mentioned above, the lot in question (UCM 15814) appears to be J. bulbosa as circumscribed herein and it does not resemble Lea’s (1863a, 1863b: pl. 38 fig. 193) original illustration of rubiginosa (Fig. 27B). Frest & Johannes (1995a: 246) provided the common name “Rusty Juga ”, and listed J. (J.) rubiginosa among species of uncertain status but considered it a potentially valid species and local endemic of conservation concern owing to agricultural impacts in the Yakima Valley. Later, Frest& Johannes (2010: 6) indicated it could be J. plicifera, but again expressed doubts as to its identity as a Juga. Other authors have considered the species to be conspecific with J. silicula (Goodrich, 1942; Burch & Tottenham 1980; Burch 1982, 1989). J.P.E. Morrison annotated the label of the lectotype, USNM 119297 (Graf 2001; Fig. 27B), that the locality was incorrect, and was “probably North or South Carolina ”. Given that no Juga species are known to possess a single, prominent spiral keel, we agree with the interpretation of Morrison that this species is a mislocalized eastern North American pleurocerid. The paralectotype is filed under Elimia symmetrica (Haldeman, 1841) in the USNM collections. Three paralectotypes in the Newcomb collections at the Paleontological Research Institute (PRI 21730) appear conspecific with the specimens in the USNM., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 82, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Lea I. 1862. Description of a new genus (Goniobasis) of the family Melanidae and eighty-two new species. Proceedings of the Academy of Natural Sciences of Philadelphia 14: 262 - 272. Available from https: // www. biodiversitylibrary. org / part / 84787 [accessed 13 Mar. 2022].","Lea I. 1863 a. New Melanidae of the United States. Journal of the Academy of Natural Sciences of Philadelphia 5: 217 - 356. Available from https: // www. biodiversitylibrary. org / page / 35217921 [accessed 13 Mar. 2022].","Tryon G. W. 1864. Synonymy of the species of Strepomatidae, a family of fluviatile Mollusca inhabiting North America. Part 2. Proceedings of the Academy of Natural Sciences of Philadelphia 16: 35 - 59.","Tryon G. W. 1865. Review of the goniobases of Oregon and California. American Journal of Conchology 1: 236 - 246.","Tryon G. W. 1866. Monograph of the family Strepomatidae. American Journal of Conchology 2: 14 - 52.","Tryon G. W. 1873. Land and Fresh-water Shells of North America Part IV. Strepomatidae (American Melanians). Smithsonian Institution, Washington, D. C.","Pilsbry H. A. 1899. Mollusks collected by R. C. McGregor in northern California. The Nautilus 13: 64 - 67.","Henderson J. 1929. The non-marine Mollusca of Oregon and Washington. University of Colorado Studies 17: 47 - 190.","Henderson J. 1936 b. The non-marine Mollusca of Oregon and Washington. Supplement. University of Colorado Studies 23: 251 - 280.","Lea I. 1863 b. New Melanidae of the United States. In: Observations on the Genus Unio, together with Descriptions of New Species, their Soft Parts, and Embryonic Forms, in the Family Unionidae, and Descriptions of New Genera and Species of the Melanidae 9: 39 - 178.","Frest T. J. & Johannes E. J. 1995 a. Interior Columbia Basin mollusk species of special concern. Final report: Interior Columbia Basin Ecosystem Management Project, Walla Walla, WA. Contract 43 - 0 E 00 - 4 - 9112.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Goodrich C. 1942. The Pleuroceridae of the Pacific coastal drainage, including the Western Interior Basin. Occasional Papers of the Museum of Zoology, University of Michigan 469: 1 - 4.","Burch J. B. & Tottenham J. 1980. North American freshwater snails, species list, ranges, and illustrations. Walkerana 3: 1 - 215.","Burch J. B. 1982. Freshwater snails (Mollusca: Gastropoda) of North America. Environmental Monitoring and Support Laboratory, Office of Research and Development, United States Environmental Protection Agency, Cincinnati, Ohio, EPA- 600 / 3 - 82 - 026.","Burch J. B. 1989. North American Freshwater Snails. Malacological Publications, Hamburg, Michigan.","Graf D. L. 2001. The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of Recent North America, north of Mexico. Walkerana 12: 1 - 124."]}

Juga bulbosa (A. Gould, 1847) Figs 10–11 Melania bulbosa A. Gould, 1847: 225. Goniobasis bairdiana I. Lea, 1862: 267. Syn. nov. Goniobasis draytonii I. Lea, 1862: 264. Syn. nov. Goniobasis hemphilli maupinensis Henderson, 1935a: 97. Syn. nov. Melania bulbosa – Gould 1852: 142; “ 1856 ” [1860]: pl. 10 fig. 163–163a; 1862: 46–47. Goniobasis bairdiana – Lea 1863a: 317–318, pl. 37 fig. 164; 1863b: 139–140, pl. 37 fig. 164. Goniobasis draytonii – Lea 1863a: 300, pl. 37 fig. 134; 1863b: 122, pl. 37 fig. 134. Juga (Oreobasis) bulbosa – Burch & Tottenham 1980: 152 (in part). — Burch 1982: 41 (in part); 1989: 152 (in part). — Frest & Johannes 2010: 9, 26. — Johannes 2015: 24. Juga (Juga) silicula – Burch & Tottenham 1980: 152 (in part). — Burch 1982: 41 (in part); 1989: 152 (in part). 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 (Juga) hemphilli maupinensis – Neitzel & Frest 1992: B.25. — Frest & Johannes 1995a: 173; 2010: 10, 37. — Johannes 2015: 23. — Campbell et al. 2016: 160. Juga (Juga) plicifera plicifera – Neitzel & Frest 1992: B.25 (in part). Juga (Oreobasis) n. sp. 1 – Frest & Johannes 1993: 64; 1995a: 178; 2001: 60. Juga (Oreobasis) n. sp. 2 – Frest & Johannes 1993: 65; 1995a: 179; 1996: 43, 131; 1999: 85, figs 33–34. — Johannes 2015: 23. Juga (Juga) n. sp. 1 – Frest & Johannes 1993: 63 (in part); 1995a: 175 (in part); 2001: 60 (in part). Juga (Juga ?) bairdiana – Frest & Johannes 1995a: 246. Juga (Juga) rubiginosa – Frest & Johannes 1995a: 246. Juga (Oreobasis ?) draytonii – Frest & Johannes 1995a: 247. Juga (Juga) n. sp. 2 – Frest & Johannes 1995a: 176; 2001: 60. — Johannes 2015: 23. Juga bulbosa – Strong & Frest 2007: 56. — Johnson et al. 2013: 282. Juga (Juga) draytonii – Frest & Johannes 2010: 10, 34 (in part). Juga (Juga) bairdiana – Campbell et al. 2016: 160. Juga species 2 – Strong & Whelan 2019: 89, fig. 4a-e. non Oxytrema bulbosa – Morrison 1954: 361, 384, pl. 11 fig. 5. [= Juga caerulea sp. nov.] non Juga bulbosa – Turgeon et al. 1988: 65; 1998: 67. [= Juga newberryi] non Juga (Oreobasis) bulbosa – Neitzel & Frest 1992: B.25. — Frest & Johannes 1995a: 177. [= Juga newberryi] non Juga (Oreobasis) n. sp. 1 – Frest & Johannes 1995b: 39; 2005: 179. — Johannes 2013a: 24. [= Juga canella sp. nov.] non Juga (Oreobasis) n. sp. 2 – Frest & Johannes 2005: 179. [= Juga canella sp. nov.] non Juga (Juga) draytonii – Campbell et al. 2016: 160. [= Juga plicifera] Material examined Lectotype of Melania bulbosa A. Gould, 1847 (designated by Graf 2001: 18) (Fig. 10A) USA • “ Columbia River ” [label: “ Oregon ”]; U.S. Exploring Expedition; J. Drayton leg.; MCZ 169067. Paralectotypes of Melania bulbosa A. Gould, 1847 USA • 9 spms; same collection data as for lectotype; MCZ 169068 • 1 spm; [labels: “ Oregon ”]; MCZ 216750 • 2 spms; “ Oregon ” [Territory]; U.S. Exploring Expedition; USNM 5563. Lectotype of Goniobasis bairdiana I. Lea, 1862 (designated by Graf 2001: 14) (Fig. 10B) USA • “ Columbia River, at Fort George, Oregon [Territory]”; J. Drayton leg.; USNM 119069. Paralectotype of Goniobasis bairdiana I. Lea, 1862 USA • 1 spm; same collection data as for lectotype; USNM 1665627 (formerly USNM 119069 X). Lectotype of Goniobasis draytonii I. Lea, 1862 (designated by Graf 2001: 38) (Fig. 10C) USA • “ Fort George, Oregon ”; J. Drayton leg.; USNM 118993. Paralectotypes of Goniobasis draytonii I. Lea, 1862 USA • 6 spms; same collection data as for lectotype; USNM 1665629 (formerly USNM 118993 X) • 12 spms; “Walla, Oregon ” [sic, Walla Walla, Oregon Territory]; received from Lady K. Douglas; USNM 119124. Holotype of Goniobasis hemphilli maupinensis Henderson, 1935 (Fig. 10D) USA • “ Deschutes River, Maupin, Oregon ”; 11 Aug. 1931; J. and B.R. Henderson leg.; UCM 17754 a. Remarks 1 paratype (UCM 17754 b, same collection data as for holotype) not examined (McCoy 1964). Other material 114 lots, 678 specimens, of which 75 were sequenced. USA – Washington • 32 spms; Klickitat Co., Major Creek; 45.7154, -121.3509; 6 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413188 • 1 spm; same collection data as for preceding; GenBank: MK472168, MK464540; USNM 1413185 • 1 spm; same collection data as for preceding; GenBank: MK472169, MK464541; USNM 1413186 • 1 spm; same collection data as for preceding; GenBank: MK472170, MK464542; USNM 1413187 • 40 spms; Skamania Co., Dog Creek; 45.7106, -121.6715; 5 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413232 • 1 spm; same collection data as for preceding; GenBank: MK472171, MK464543; USNM 1413227 • 1 spm; same collection data as for preceding; GenBank: MK472172, MK464544; USNM 1413228 • 1 spm; same collection data as for preceding; GenBank: MK472173, MK464545; USNM 1413229 • 1 spm; same collection data as for preceding; GenBank: MK472174, MK464546; USNM 1413230 • 1 spm; same collection data as for preceding; GenBank: MK472175, MK464547; USNM 1413231 • 8 spms; Yakima Co., Ahtanum Creek, south of Union Gap; [46.5369, -120.4751]; UCM 15814. – “ Oregon ” [Territory] • 1 spm; USNM 59058 • 3 spms; same collection data as for preceding; USNM 56427 • 1 spm; same collection data as for preceding; USNM 36948 • 2 spms; same collection data as for preceding; USNM 12141 • 6 spms; same collection data as for preceding; USNM 12209. – Oregon • 10 spms; Hood River Co., Post Canyon Dr; 45.697, -121.5749; 8 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295063 • 1 spm; same collection data as for preceding; GenBank: MK472148, MK464520; USNM 1295056 • 1 spm; same collection data as for preceding; GenBank: MK472149, MK464521; USNM 1295057 • 1 spm; same collection data as for preceding; GenBank: MK472150, MK464522; USNM 1295058 • 1 spm; same collection data as for preceding; GenBank: MK472151, MK464523, MK480749; USNM 1295059 • 1 spm; same collection data as for preceding; GenBank: MK472158, MK464530; USNM 1295060 • 1 spm; same collection data as for preceding; GenBank: MK472159, MK464531; USNM 1295061 • 1 spm; same collection data as for preceding; GenBank: MK472160, MK464532; USNM 1295062 • 1 spm; same collection data as for preceding; GenBank: MK472147, MK464519; USNM 1295055 • 20 spms; Hood River Co., SH 281, Hood River; 45.6971, -121.5231; 8 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295095 • 1 spm; same collection data as for preceding; GenBank: MK472152, MK464524; USNM 1295087 • 1 spm; same collection data as for preceding; GenBank: MK472153, MK464525; USNM 1295088 • 1 spm; same collection data as for preceding; GenBank: MK472154, MK464526; USNM 1295089 • 1 spm; same collection data as for preceding; GenBank: MK472155, MK464527, MK480750; USNM 1295090 • 1 spm; same collection data as for preceding; GenBank: MK472156, MK464528; USNM 1295091 • 1 spm; same collection data as for preceding; GenBank: MK472161, MK464533; USNM 1295092 • 1 spm; same collection data as for preceding; GenBank: MK472162, MK464534; USNM 1295093 • 1 spm; same collection data as for preceding; GenBank: MK472163, MK464535; USNM 1295094 • 28 spms; Hood River Co., West Fork Neal Creek at 1700 Rd; 45.5421, -121.5222; 5 Aug. 2015; Jennings leg.; USNM 1413445 • 1 spm; same collection data as for preceding; GenBank: MK472202; USNM 1413438 • 1 spm; same collection data as for preceding; GenBank: MK472203, MK464574; USNM 1413439 • 1 spm; same collection data as for preceding; GenBank: MK472204, MK464575; USNM 1413440 • 1 spm; same collection data as for preceding; GenBank: MK472205; USNM 1413441 • 1 spm; same collection data as for preceding; GenBank: MK472206, MK464576; USNM 1413442 • 1 spm; same collection data as for preceding; USNM 1413443 • 1 spm; same collection data as for preceding; GenBank: MK472207, MK464577; USNM 1413444 • 1 spm; Jefferson Co., Deschutes River at junction with Dry Creek; 44.7844, -121.1959; 8 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413422 • 25 spms; Jefferson Co., Deschutes River, 100 yds. S of Hwy 26 bridge at Warm Springs; [44.7595, -121.2277]; 2 Aug. 1971; J. Landye leg.; USNM 892362 • 1 spm; Jefferson Co., U.S. Hwy 26; 44.7584, -121.2272; 9 Jul. 2014; J.T. Garner and N.V. Whelan leg.; GenBank: MK472157, MK464529; USNM 1295201 • 59 spms; Sherman Co., confluence of Muddy Hollow and China Hollow; 45.6471, -120.8086; 6 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413310 • 1 spm; same collection data as for preceding; GenBank: MK472181, MK464553; USNM 1413305 • 1 spm; same collection data as for preceding; GenBank: MK472182, MK464554, MK480751; USNM 1413306 • 1 spm; same collection data as for preceding; GenBank: MK472183, MK464555; USNM 1413307 • 1 spm; same collection data as for preceding; GenBank: MK472184, MK464556; USNM 1413308 • 1 spm; same collection data as for preceding; GenBank: MK472185; USNM 1413309 • 54 spms; Sherman Co., Muddy Hollow 2; 45.6311, -120.7933; 6 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413316 • 1 spm; same collection data as for preceding; GenBank: MK472186, MK464557, MK480752; USNM 1413311 • 1 spm; same collection data as for preceding; GenBank: MK472187, MK464558, MK480753; USNM 1413312 • 1 spm; same collection data as for preceding; GenBank: MK472188, MK464559, MK480754; USNM 1413313 • 1 spm; same collection data as for preceding; GenBank: MK472189, MK464560; USNM 1413314 • 1 spm; same collection data as for preceding; GenBank: MK472190, MK464561; USNM 1413315 • 1 spm; Wasco Co., Oak Springs; 45.2216, -121.0827; 8 Sep. 2015; E.E. Strong and P. Bouchet leg.; GenBank: MK472164, MK464536; USNM 1413165 • 1 spm; same collection data as for preceding; GenBank: MK472165, MK464537; USNM 1413166 • 1 spm; same collection data as for preceding; GenBank: MK472166, MK464538; USNM 1413167 • 1 spm; same collection data as for preceding; GenBank: MK472167, MK464539; USNM 1413168 • 4 spm; Skamania Co., above Bonneville Dam, Sullivan’s Id., Columbia River; [45.6916, -121.882]; H.W. Krieger leg.; USNM 468428 • 23 spms; Wasco Co., Chenoweth Creek on Chenoweth Rd; [45.6289, -121.2283]; 1 Aug. 1983; J. Landye leg.; USNM 1152452 • 45 spms; Wasco Co., Fifteenmile Creek; 45.4309, -121.2249; 6 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413359 • 1 spm; same collection data as for preceding; GenBank: MK472191, MK464562; USNM 1413354 • 1 spm; same collection data as for preceding; GenBank: MK472192, MK464563; USNM 1413355 • 1 spm; same collection data as for preceding; GenBank: MK472193, MK464564, MK480755; USNM 1413356 • 1 spm; same collection data as for preceding; GenBank: MK472194, MK464565; USNM 1413357 • 1 spm; same collection data as for preceding; GenBank: MK472195, MK464566; USNM 1413358 • 1 spm; Wasco Co., Gate Creek at the 4820 Rd; 45.2193, -121.4305; 9 Jul. 2015; Jennings leg.; GenBank: MK472215; USNM 1413455 • 1 spm; Wasco Co., Gate Creek at the 4820 Rd; 45.2193, -121.4305; 9 Jul. 2015; Jennings leg.; USNM 1413462 • 19 spms; Wasco Co., Gate Creek at the 4820 Rd; 45.2192, -121.4305; 5 Aug. 2015; Jennings leg.; USNM 1413454 • 1 spm; same collection data as for preceding; GenBank: MK472211, MK464581, MK480757; USNM 1413450 • 1 spm; same collection data as for preceding; GenBank: MK472212, MK464582; USNM 1413451 • 1 spm; same collection data as for preceding; GenBank: MK472213, MK464583; USNM 1413452 • 1 spm; same collection data as for preceding; GenBank: MK472214, MK464584; USNM 1413453 • 40 spms; Wasco Co., Harpham Flat; 45.1375, -121.122; 8 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413304 • 1 spm; same collection data as for preceding; GenBank: MK472176, MK464548; USNM 1413299 • 1 spm; same collection data as for preceding; GenBank: MK472177, MK464549; USNM 1413300 • 1 spm; same collection data as for preceding; GenBank: MK472178, MK464550; USNM 1413301 • 1 spm; same collection data as for preceding; GenBank: MK472179, MK464551; USNM 1413302 • 1 spm; same collection data as for preceding; GenBank: MK472180, MK464552; USNM 1413303 • 26 spms; Wasco Co., I-84 eastbound at The Dalles Dam just past exit 88 (Frest spring #3 site); 45.6256, -121.1059; 18 May 2017; K. Van Norman leg.; USNM 1592625 • 1 spm; same collection data as for preceding; USNM 1592629 • 1 spm; same collection data as for preceding; USNM 1592626 • 1 spm; same collection data as for preceding; USNM 1592627 • 1 spm; same collection data as for preceding; USNM 1592628 • 1 spm; Wasco Co., South Fork Gate Creek at the 4830 Rd; 45.196, -121.4132; 15 Jul. 2015; Jennings leg.; GenBank: MK472216, MK464585, MK480758; USNM 1413456 • 29 spms; Wasco Co., South Fork Gate Creek at the 4830 Rd; 45.196, -121.4132; 5 Aug. 2015; Jennings leg.; USNM 1413461 • 1 spm; same collection data as for preceding; GenBank: MK472217, MK464586; USNM 1413457 • 1 spm; same collection data as for preceding; GenBank: MK472218, MK464587; USNM 1413458 • 1 spm; same collection data as for preceding; GenBank: MK464588; USNM 1413459 • 1 spm; same collection data as for preceding; GenBank: MK472219, MK464589; USNM 1413460 • 1 spm; Wasco Co., South Junction; 44.8602, -121.0604; 8 Sep. 2015; E.E. Strong and P. Bouchet leg.; GenBank: MK472196, MK464567; USNM 1413423 • 1 spm; same collection data as for preceding; GenBank: MK472197, MK464568; USNM 1413424 • 1 spm; same collection data as for preceding; GenBank: MK472198, MK464569; USNM 1413425 • 1 spm; same collection data as for preceding; GenBank: MK472199, MK464570; USNM 1413426 • 1 spm; same collection data as for preceding; GenBank: MK472200, MK464571; USNM 1413427 • 1 spm; Wasco Co., Souva Creek off the 4830120 Rd; 45.2081, -121.448; 8 Jul. 2015; Jennings leg.; GenBank: MK472201, MK464573; USNM 1413437 • 1 spm; Wasco Co., Souva Creek off the 4830120 Rd; 45.2085, -121.4483; 8 Jul. 2015; Jennings leg.; USNM 1413448 • 1 spm; same collection data as for preceding; GenBank: MK472210, MK464580; USNM 1413449 • 1 spm; Wasco Co., Souva Creek off the 4830120 Rd; 45.209, -121.4493; 8 Jul. 2015; Jennings leg.; GenBank: MK472208, MK464578; USNM 1413446 • 1 spm; Wasco Co., Souva Creek off the 4830120 Rd; 45.2094, -121.45; 8 Jul. 2015; Jennings leg.; GenBank: MK472209, MK464579; USNM 1413447 • 25 spms; Wasco Co., Souva Creek off the 4830120 Rd; 45.2081, -121.4481; 5 Aug. 2015; Jennings leg.; USNM 1413436 • 1 spm; same collection data as for preceding; GenBank: MK464572, MK480756; USNM 1413435 • 48 spms; Wasco Co., Warm Springs River; 44.9672, -121.4687; 6 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413407 • 1 spm; same collection data as for preceding; USNM 1413402 • 1 spm; same collection data as for preceding; USNM 1413403 • 1 spm; same collection data as for preceding; USNM 1413404 • 1 spm; same collection data as for preceding; USNM 1413405 • 1 spm; same collection data as for preceding; USNM 1413406 • 33 spms; Wasco Co., Warm Springs River, at Hwy 26 Crossing NW of Warm Springs River; [44.9669, -121.4678]; 2 Aug. 1971; J. Landye leg.; on rocks; USNM 892374 • 7 spms; Wasco Co., Warm Springs River, near Kah-Nee-Ta Warm Spring Area by highway bridge; [44.8664, -121.2164]; 23 Apr. 1975; J. Landye leg.; on sand; USNM 892376 • 2 spms; Willamette River [sic, mislocalized]; S. Smith leg.; USNM 321825. Description SHELL. Thick, large, reaching ~ 2.75 cm in length; turriform, narrowly conical in shape, spire height moderate to tall (Fig. 10). Whorls somewhat flattened to moderately convex, suture weakly to moderately impressed. Aperture oval in shape, lip smooth and simple. Spiral sculpture of numerous fine striae, usually indistinct but occasionally weakly to moderately developed, variable in strength and number; sporadic indistinct elevated lirae. Axial sculpture of fine, sinuous growth lines; plications present or absent, confined to upper part of spire; plications regular and even, weakly to moderately developed, occasionally shouldered, orthocline to weakly opisthocyrt, variable in strength and number. Shell black, dark or reddish brown, to tan in color, occasionally with a lighter subsutural band; reddish brown bands present or absent, one to three in number and variable in width. Interior of aperture white, yellowishbrown to dark purple in color; bands when present visible inside aperture. RADULA. Rachidian narrowly rectangular, wider than tall, basal margin flattened between bluntly rounded median and lateral projections; basal denticles lacking or vestigial (Fig. 11). Upper margin slightly concave with cutting edge bearing central elongate, conical cusp, and two stout, conical denticles on each side. Lateral teeth with prominent triangular cusp flanked by two inner and two to three outer, triangular denticles, and occasionally 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 of inner marginal teeth and extending almost to tooth bases of outer marginal teeth; broad outer flanges extending almost to tooth bases. Inner marginal teeth with four and outer marginal teeth with four to five flattened denticles. Distribution and ecology In springs, spring runs, and spring-fed creeks to large rivers in the central and eastern Columbia River Gorge drainages in north-central Oregon and south-central Washington, including the Mt. Hood National Forest, Columbia Gorge National Scenic Area, and Lower Deschutes River below Pelton Dam (Fig. 7C). Remarks The type locality of Goniobasis bairdiana and G. draytonii, both ostensibly from the Columbia River at Fort George [= Astoria] collected by J. Drayton, is likely erroneous (Frest & Johannes 2010), as the species does not extend to the western Columbia Gorge and there is no evidence it ever has. All sequenced specimens and historical lots in museum collections from the terminal part of, Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on pages 33-41, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Gould A. A. 1847. Descriptions of the following species of Melania, from the collection of the Exploring Expedition. Proceedings of the Boston Society of Natural History 2: 222 - 225. https: // doi. org / 10.5962 / bhl. part. 3322","Lea I. 1862. Description of a new genus (Goniobasis) of the family Melanidae and eighty-two new species. Proceedings of the Academy of Natural Sciences of Philadelphia 14: 262 - 272. Available from https: // www. biodiversitylibrary. org / part / 84787 [accessed 13 Mar. 2022].","Henderson J. 1935 a. West American species of Goniobasis, with a description of new forms. The Nautilus 48: 94 - 99, 130 - 134.","Gould A. A. 1852. Mollusca and shells. Volume 12. In: United States Exploring Expedition during the Years 1838, 1839, 1840, 1841, 1842. Under the Command of Charles Wilkes, U. S. N. Gould and Lincoln, Boston. https: // doi. org / 10.5962 / bhl. title. 69333","Gould A. A. 1856 [15 December 1860]. Atlas. Mollusca and shells. In: United States Exploring Expedition during the Years 1838, 1839, 1840, 1841, 1842. Under the Command of Charles Wilkes, U. S. N. Gould and Lincoln, Boston. https: // doi. org / 10.5962 / bhl. title. 69333","Gould A. A. 1862. Otia Conchologica: Descriptions of Shells and Mollusks, from 1839 to 1862. Gould and Lincoln, Boston. https: // doi. org / 10.5962 / bhl. title. 12860","Lea I. 1863 a. New Melanidae of the United States. Journal of the Academy of Natural Sciences of Philadelphia 5: 217 - 356. Available from https: // www. biodiversitylibrary. org / page / 35217921 [accessed 13 Mar. 2022].","Burch J. B. & Tottenham J. 1980. North American freshwater snails, species list, ranges, and illustrations. Walkerana 3: 1 - 215.","Burch J. B. 1982. Freshwater snails (Mollusca: Gastropoda) of North America. Environmental Monitoring and Support Laboratory, Office of Research and Development, United States Environmental Protection Agency, Cincinnati, Ohio, EPA- 600 / 3 - 82 - 026.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Johannes E. J. 2015. Potential impacts from climate change on freshwater mollusks in the Deschutes, Klamath, and John Day basins. In: Wisseman R. (ed.) Potential Impacts from Climate Change on Freshwater Invertebrates, including Mollusks, in the Deschutes, Klamath, and John Day Basins of Eastern Oregon: 19 - 37.","Neitzel D. A. & Frest T. J. 1992. Survey of Columbia River Basin streams for Columbia Pebblesnail Fluminicola columbiana and Shortface Lanx Fisherola nuttalli. Prepared for the U. S. Department of Energy under contract DE-AC 06 - 76 RLO 1830. Battelle Memorial Institute, Pacific Northwest Laboratories, Richland, Washington, PNL- 8229. https: // doi. org / 10.2172 / 10173267","Frest T. J. & Johannes E. J. 1995 a. Interior Columbia Basin mollusk species of special concern. Final report: Interior Columbia Basin Ecosystem Management Project, Walla Walla, WA. Contract 43 - 0 E 00 - 4 - 9112.","Campbell D. C., Clark S. A., Johannes E. J., Lydeard C. & Frest T. J. 2016. Molecular phylogenetics of the freshwater gastropod genus Juga (Cerithioidea: Semisulcospiridae). Biochemical Systematics and Ecology 65: 158 - 170. https: // doi. org / 10.1016 / j. bse. 2016.01.004","Frest T. J. & Johannes E. J. 1993. Mollusc species of Special Concern within the range of the Northern Spotted Owl. Final report to Forest Ecosystem Management Working Group, USDA Forest Service. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 2001. Baseline freshwater mollusk survey of Mount Rainier National Park. 2000 yearly report. Final report prepared for U. S. Department of the Interior, National Park Service, Mount Rainier National Park, Ashland, Washington. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 1996. Additional information on certain mollusk species of special concern occurring within the range of the Northern Spotted Owl. Report to Oregon State Office, BLM. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 1999. Field guide to survey and manage freshwater mollusk species. United States Department of the Interior; Bureau of Land Management, Oregon State office; United States Fish and Wildlife Service, Northwest Regional Ecosystems Office; and United States Department of Agriculture, Forest Service, Region 6, Portland, Oregon. BLM / OR / WA / PL- 99 / 045 + 1792.","Frest T. J. & Johannes E. J. 1995 b. Freshwater mollusks of the Upper Sacramento System, California, with particular reference to the Cantara Spill. 1994 yearly report to California Department of Fish and Game. Deixis Consultants, Seattle, Washington.","Strong E. E. & Frest T. 2007. On the anatomy and systematics of Juga from western North America (Gastropoda: Cerithioidea: Pleuroceridae). The Nautilus 121: 43 - 65.","Johnson P. D., Bogan A. E., Brown K. M., Burkhead N. M., Cordeiro J. R., Garner J. T., Hartfield P. D., Lepitzki D. A. W., Mackie G. L., Pip E., Tarpley T. A., Tiemann J. S., Whelan N. V. & Strong E. E. 2013. Conservation status of freshwater gastropods of Canada and the United States. Fisheries 38: 247 - 282. https: // doi. org / 10.1080 / 03632415.2013.785396","Strong E. E. & Whelan N. V. 2019. Assessing the diversity of Western North American Juga (Semisulcospiridae, Gastropoda). Molecular Phylogenetics and Evolution 136: 87 - 103. https: // doi. org / 10.1016 / j. ympev. 2019.04.009","Morrison J. P. E. 1954. The relationships of Old and New World melanians. Proceedings of the United States National Museum 103: 357 - 394. https: // doi. org / 10.5479 / si. 00963801.103 - 3325.357","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1988. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 26.","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1998. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. Second Edition. American Fisheries Society Special Publication 26.","Frest T. J. & Johannes E. J. 2005. Springsnails of the Cascade-Siskiyou National Monument and vicinity, Oregon. 2004 report. Final report prepared for World Wildlife Fund, Ashland, Oregon. Deixis Consultants, Seattle, Washington.","Johannes E. J. 2013 a. Extraordinary endemic springsnail radiation: justification for expanding the Cascade-Siskiyou National Monument, Southwest Oregon. Tentacle 21: 23 - 26.","Graf D. L. 2001. The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of Recent North America, north of Mexico. Walkerana 12: 1 - 124.","McCoy C. J. Jr. 1964. Type specimens of Recent Gastropoda in the University of Colorado Museum. University of Colorado Studies, Series in Biology 12: 1 - 7.","Smith A. G. 1937. The type locality of Oreohelix strigosa (Gould). The Nautilus 50: 73 - 77","Carpenter P. P. 1857. Report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Report of the British Association for the Advancement of Science 1856: 159 - 368. https: // doi. org / 10.5962 / bhl. title. 60613","Tryon G. W. 1865. Review of the goniobases of Oregon and California. American Journal of Conchology 1: 236 - 246.","Roscoe E. J. 1963. Some goniobases in western United States. The Nautilus 77: 43 - 47.","Burch J. B. 1989. North American Freshwater Snails. Malacological Publications, Hamburg, Michigan.","Campbell D. C. 2019. Semisulcospiridae Morrison, 1952. In: Lydeard C. & Cummings K. S. (eds) Freshwater Mollusks of the World: A Distribution Atlas: 81 - 85. Johns Hopkins University Press. Baltimore, MD.","Pilsbry H. A. 1899. Mollusks collected by R. C. McGregor in northern California. The Nautilus 13: 64 - 67.","Walker B. 1918. A synopsis of the classification of the fresh-water Mollusca of North America, North of Mexico, and a catalogue of the more recently described species, with notes. Miscellaneous Publications of the University of Michigan Museum of Zoology 6: 1 - 213. https: // doi. org / 10.5962 / bhl. title. 46279","Goodrich C. 1942. The Pleuroceridae of the Pacific coastal drainage, including the Western Interior Basin. Occasional Papers of the Museum of Zoology, University of Michigan 469: 1 - 4.","Henderson J. 1936 b. The non-marine Mollusca of Oregon and Washington. Supplement. University of Colorado Studies 23: 251 - 280.","Henderson J. 1929. The non-marine Mollusca of Oregon and Washington. University of Colorado Studies 17: 47 - 190.","Furnish J. L. 2005. Sensitive aquatic mollusks of the U. S. Forest Service Pacific Southwest Region. USDA Forest Service, Pacific Southwest Region, Vallejo, CA.","Duncan N. 2008. Survey Protocol for Aquatic Mollusk Species: Preliminary Inventory and Presence / Absence Sampling, Version 3.1. Portland, OR. Interagency Special Status / Sensitive Species Program. U. S. Department of Interior, Bureau of Land Management, Oregon / Washington and U. S. Department of Agriculture, Forest Service, Region 6.","USDA [United States Department of Agriculture], USDI [United States Department of the Interior]. 1994. Standards and guidelines for management of habitat for late-successional and old-growth forest related species within the range of the Northern Spotted Owl. Attachment A to the record of decision for amendments to Forest Service and Bureau of Land Management planning documents within the range of the Northern Spotted Owl. Available from https: // www. fs. usda. gov / Internet / FSE _ DOCUMENTS / fseprd 479477. pdf [accessed 13 Mar. 2022].","USDA [United States Department of Agriculture], USDI [United States Department of the Interior]. 2000. Final supplemental Environmental Impact Statement for amendment to the survey and manage, protection buffer, and other mitigation measures standards and guidelines Volume II - Appendices. BLM / OR / WA / PT- 00 / 065 + 1792. Available from https: // www. fs. usda. gov / r 6 / reo / survey-and-manage / 2001 / surveymanage- 2001 - fseis-vol-ii-appendices. pdf [accessed 13 Mar. 2022].","USFWS [United States Fish and Wildlife Service]. 2011. Endangered and threatened wildlife and plants; 90 - day finding on a petition to list 29 mollusk species as threatened or endangered with critical habitat. Federal Register 76: 61826 - 61853.","USFWS [United States Fish and Wildlife Service]. 2012. Endangered and threatened wildlife and plants; 12 - month finding on a petition to list 14 aquatic mollusks as endangered or threatened. Federal Register 77: 57922 - 57948."]}

Juga newberryi (I. Lea, 1860) Figs 19–20 Melania newberryi I. Lea, 1860: 93. Goniobasis newberryi – Lea 1863a: 300–301, pl. 37 fig. 135; 1863b: 122–123, pl. 37 fig. 135. Juga (Oreobasis) bulbosa – Burch & Tottenham 1980: 152, fig. 452 (in part). — Burch 1982: 41, fig. 452 (in part); 1989: 152, fig. 452 (in part). — Neitzel & Frest 1992: B.25. — Frest & Johannes 1995a: 177. Juga bulbosa – Turgeon et al. 1988: 65; 1998: 67. Juga (Oreobasis) n. sp. 3 – Frest & Johannes 1995a: 179. — Johannes 2015: 23. Juga (Oreobasis) n. sp. 4 – Frest & Johannes 1995a: 180. — Johannes 2015: 23. Juga (Oreobasis) newberryi – Strong & Frest 2007: 54. — Frest & Johannes 2010: 10, 39. — Johannes 2015: 23. — Campbell et al. 2016: 160. Juga newberryi – Johnson et al. 2013: 282. — Strong & Whelan 2019: 89. Juga (Oreobasis) OTU 1 – Campbell et al. 2016: 160. non Juga (Oreobasis) n. sp. 3 – Frest & Johannes 1993: 65; 1996: 44, 132; 1999: 89, figs 35–36. [= Juga canella sp. nov.] non Juga newberryi – Strong & Köhler 2009: 486. [= Juga nigrina] Material examined Lectotype of Melania newberryi I. Lea, 1860 (designated by Graf 2001: 70) (Fig. 19A) USA • “ Upper des Chutes [sic, Deschutes] River, Oregon Territory”; J.S. Newberry, M.D. leg.; USNM 118961. Paralectotypes of Melania newberryi I. Lea, 1860 USA • 12 spms; same collection data as for lectotype; USNM 1665630 (formerly USNM 118961X). Other material examined 42 lots, 380 specimens, of which 27 were sequenced. USA – “ Oregon ” [Territory] • 2 spms; U.S. Exploring Expedition; USNM 12174. – Oregon • 24 spms; Jefferson Co., Deschutes River at junction with Dry Creek; 44.7844, -121.1959; 8 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413421 • 1 spm; same collection data as for preceding; USNM 1413414 • 1 spm; same collection data as for preceding; USNM 1413415 • 1 spm; same collection data as for preceding; USNM 1413416 • 1 spm; same collection data as for preceding; USNM 1413417 • 1 spm; same collection data as for preceding; USNM 1413418 • 1 spm; same collection data as for preceding; USNM 1413419 • 1 spm; same collection data as for preceding; USNM 1413420 • 56 spms; Jefferson Co., Deschutes River at Rainbow Landing; 44.7578, -121.2272; 6 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413434 • 1 spm; same collection data as for preceding; GenBank: MK472315, MK464682, MK480826; USNM 1413428 • 1 spm; same collection data as for preceding; GenBank: MK472316, MK464683, MK480827; USNM 1413429 • 1 spm; same collection data as for preceding; GenBank: MK472317, MK464684, MK480828; USNM 1413430 • 1 spm; same collection data as for preceding; GenBank: MK472318, MK464685, MK480829; USNM 1413431 • 1 spm; same collection data as for preceding; GenBank: MK472319, MK464686, MK480830; USNM 1413432 • 1 spm; same collection data as for preceding; GenBank: MK472320, MK464687, MK480831; USNM 1413433 • 30 spms; Jefferson Co., Deschutes River at RM 97.2 to 97.4 on east side channel, ca 0.3–0.6 km S of U.S. 526 bridge and mouth of Shitike Creek at Rainbow Landing; 44.7575, -121.2267; 13 Aug. 2000; T. Frest leg.; USNM 1100660 • 67 spms; Jefferson Co., Opal Springs; 44.4905, -121.2981; 7 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413335 • 1 spm;same collection data as for preceding; GenBank: MK472312, MK464679, MK480823; USNM 1413332 • 1 spm; same collection data as for preceding; GenBank: MK472313, MK464680, MK480824; USNM 1413333 • 1 spm; same collection data as for preceding; GenBank: MK472314, MK464681, MK480825; USNM 1413334 • 1 spm; same collection data as for preceding; GenBank: MK472311, MK464678; USNM 1413331 • 1 spm; same collection data as for preceding; GenBank: MK472310, MK464677, MK480822; USNM 1413330 • 33 spms; Jefferson Co., U.S. Hwy 26; 44.7584, -121.2272; 9 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295086 • 1 spm; same collection data as for preceding; GenBank: MK472303, MK464670, MK480820; USNM 1295085 • 1 spm; same collection data as for preceding; GenBank: MK472301, MK464668, MK480818; USNM 1295083 • 1 spm; same collection data as for preceding; GenBank: MK472302, MK464669, MK480819; USNM 1295084 • 1 spm; same collection data as for preceding; GenBank: MK472299, MK464666, MK480816; USNM 1295081 • 1 spm; same collection data as for preceding; GenBank: MK472300, MK464667, MK480817; USNM 1295082 • 1 spm; same collection data as for preceding; GenBank: MK472296, MK464663, MK480813; USNM 1295078 • 1 spm; same collection data as for preceding; GenBank: MK472298, MK464665, MK480815; USNM 1295080 • 1 spm; same collection data as for preceding; GenBank: MK472297, MK464664, MK480814; USNM 1295079 • 1 spm; same collection data as for preceding; GenBank: MK472294, MK464661, MK480811; USNM 1295076 • 1 spm; same collection data as for preceding; GenBank: MK472295, MK464662, MK480812; USNM 1295077 • 25 spms; Jefferson Co., Willow Creek, at inflow to Lake Simtustus, Madras West, 2400 ft S, 1800 ft E of NW corner; [44.6718, -121.2278]; 2 Aug. 1971; J. Landye leg.; in substrate; USNM 905120 • 109 spms; Wasco Co., Oak Springs; 45.2216, -121.0827; 8 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413323 • 1 spm; same collection data as for preceding; GenBank: MK472305, MK464672; USNM 1413318 • 1 spm; same collection data as for preceding; GenBank: MK472306, MK464673; USNM 1413319 • 1 spm; same collection data as for preceding; GenBank: MK472307, MK464674; USNM 1413320 • 1 spm; same collection data as for preceding; GenBank: MK472308, MK464675; USNM 1413321 • 1 spm; same collection data as for preceding; GenBank: MK472309, MK464676; USNM 1413322 • 1 spm; same collection data as for preceding; GenBank: MK472304, MK464671, MK480821; USNM 1413317. Description SHELL. Moderately thick, small for the genus, reaching ~ 1.5 cm in length; broadly to elongately conical in shape, spire height low to moderate (Fig. 19). Whorls convex to somewhat flattened, shouldered, suture deeply impressed, body whorl usually inflated. Aperture broadly oval in shape, lip smooth, simple. Spiral sculpture lacking. Axial sculpture of fine, weakly prosocline to opisthocyrt growth lines; plications lacking. Shell dark purple to tan in color, usually with three reddish brown bands, variable in width. Interior of aperture white to dark purple in color; bands when present visible inside aperture. Columella occasionally tinged reddish-purple. RADULA. Rachidian narrowly rectangular, wider than tall, with angular to convex lower margin and projecting outer corners; basal denticles lacking or vestigial (Fig. 20). 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 half to two-thirds of shafts; broad outer flanges extending almost to tooth bases. Inner marginal teeth with four to five and outer marginal teeth with six to seven flattened denticles. See also Strong & Frest (2007). Distribution and ecology In the lower Middle and Lower Deschutes River in central Oregon, and associated springs, spring runs and spring-fed creeks in the Deschutes River drainage (Fig. 21A). Remarks Tryon (1864, 1865, 1866, 1873) viewed Goniobasis newberryi as valid and distinct but remarked that this species and G. bulbosa were “exactly similar in outline” (Tryon 1864: 53; 1865: 246; 1873: 255) and “certainly extremely closely allied” (Tryon 1865: 245), differing only in the presence or absence of bands (Tryon 1866: 34). Pilsbry (1899) subsequently synonymized the two, which was followed by most authors and resulted in confusion about the identity and distribution of Juga newberryi for over 100 years (see Strong & Frest 2007). Most usages of the name bulbosa in the literature in fact refer to J. newberryi (e.g., Tryon 1865: fig. 17; 1866: fig. 217; 1873: fig. 496; Roscoe 1963: pl. 4 figs 5–6; Burch & Tottenham 1980: fig. 452; Burch 1982, 1989: fig. 452; Turgeon et al. 1988, 1998). This confusion was compounded by the fact that the species is inexplicably rare in museum collections (see e.g., Henderson 1936b) despite occurring in abundance at some sites. Apart from the types and material collected for the present research, holdings of newberryi in the USNM collections include only three lots, one from the U.S. Exploring Expedition, one collected in 1971 and one in 2000. What little museum material there is in some cases contains mixtures of the two species (see account for J. bulbosa, above). Although Frest & Johannes (2010: 26) could not determine its identity, the specimen figured by Tryon as Goniobasis bulbosa (1865: fig. 17; 1866: fig. 217; 1873: fig. 496) is the larger of the two specimens in USNM 12174 (Fig. 19N), entered in the handwritten ledger catalogue on May 10, 1876, identified as “ Strepoma bulbosa Gould ” with no other information than being from “ Oregon ” [Territory]. Tryon (1865: 246) claimed the specimen to be, “the largest I have seen, and considerably exceeds the dimensions of Dr. Gould’s type specimen”. There can be no doubt that it represents a specimen of J. newberryi given the low spire and inflated body whorl. Paralectotype lot MCZ 169068 comprises a mixture of bulbosa and newberryi, as does USNM 5563 which contains Gould’s (1860: fig. 163–163a) figured specimen of bulbosa (Fig. 10E–F). The erroneous synonymy of newberryi with bulbosa notwithstanding, this is a distinctive Deschutes River endemic, particularly the low spired, inflated morph. However, banded subadults of J. bulbosa (e.g., Fig. 10S) may be confused for J. newberryi without careful scrutiny when they co-occur (e.g., Fig. 19C–D) but can be distinguished by the degree of banding, spire height, and inflation of the body whorl. The early teleoconchs of bulbosa are frequently eroded but when preserved may be plicate, whereas they are invariably smooth in newberryi. The tall slender morph of newberryi (Fig. 19G–H, L–M) also may be difficult to differentiate from bulbosa as the banding patterns can be quite similar. One lot containing dead-collected specimens from the confluence of Willow Creek and the Deschutes River (USNM 905120; Fig. 19L–M) compares favorably in size and proportion with specimens sequenced from Opal Springs (Fig. 19G–H) and are considered to represent newberryi. Specimens of bulbosa sequenced nearby generally have a less deeply impressed suture and are larger in size at adulthood (e.g., Fig. 10P). There are banded and unbanded morphs of both species, even at a single site, but unbanded morphs of newberryi are rare and are usually quite dark, not the tan to golden base color as seen in most historical lots. It is uncertain whether these golden, unbanded forms are merely faded or represent a local variant that has not been recollected. Distribution The type locality of newberryi is the Upper Deschutes River, which is considered the region upstream of Bend, Oregon, but there is no evidence it ever extended this far upriver. Frest & Johannes (1995a, 2010) and Strong & Frest (2007) considered the types to have originated from near Bend without justification and indicated the species occurs sporadically below Pelton Dam to roughly 6 miles above the mouth where it is replaced by “ J. (J.) plicifera ” (= J. bulbosa as circumscribed herein). However, some spring dwelling forms (e.g., Opal Springs, Oak Springs) previously considered an undescribed species (see below) are conspecific with J. newberryi (Strong & Whelan 2019) and place the species above Pelton Dam. Common name The common name for this species is the Banded Juga (Johnson et al. 2013). It has been referred to as the Newberry Juga (Johannes 2015) and the Bulb Juga (Frest & Johannes 1995a; Turgeon et al. 1988, 1998), the latter reflecting the erroneous synonymy with J. bulbosa. Putatively new species that are conspecific with J. newberryi have been referred to as the Crooked River or Opal Springs Juga [= J. (Oreobasis) n. sp. 4] (Fig. 19G–H) and the Oak Springs or Purple Juga (e.g., Frest & Johannes 1995a; Duncan 2008; Johannes 2015; Fallon et al. 2020). The latter is a dark, unbanded form (Fig. 19E–F)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on pages 55-60, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Lea I. 1860. Descriptions of four new species of Melanidae of the United States. Proceedings of the Academy of Natural Sciences of Philadelphia 12: 93. Available from https: // www. biodiversitylibrary. org / page / 1801109 [accessed 13 Mar. 2022].","Lea I. 1863 a. New Melanidae of the United States. Journal of the Academy of Natural Sciences of Philadelphia 5: 217 - 356. Available from https: // www. biodiversitylibrary. org / page / 35217921 [accessed 13 Mar. 2022].","Burch J. B. & Tottenham J. 1980. North American freshwater snails, species list, ranges, and illustrations. Walkerana 3: 1 - 215.","Burch J. B. 1982. Freshwater snails (Mollusca: Gastropoda) of North America. Environmental Monitoring and Support Laboratory, Office of Research and Development, United States Environmental Protection Agency, Cincinnati, Ohio, EPA- 600 / 3 - 82 - 026.","Burch J. B. 1989. North American Freshwater Snails. Malacological Publications, Hamburg, Michigan.","Neitzel D. A. & Frest T. J. 1992. Survey of Columbia River Basin streams for Columbia Pebblesnail Fluminicola columbiana and Shortface Lanx Fisherola nuttalli. Prepared for the U. S. Department of Energy under contract DE-AC 06 - 76 RLO 1830. Battelle Memorial Institute, Pacific Northwest Laboratories, Richland, Washington, PNL- 8229. https: // doi. org / 10.2172 / 10173267","Frest T. J. & Johannes E. J. 1995 a. Interior Columbia Basin mollusk species of special concern. Final report: Interior Columbia Basin Ecosystem Management Project, Walla Walla, WA. Contract 43 - 0 E 00 - 4 - 9112.","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1988. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 26.","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1998. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. Second Edition. American Fisheries Society Special Publication 26.","Johannes E. J. 2015. Potential impacts from climate change on freshwater mollusks in the Deschutes, Klamath, and John Day basins. In: Wisseman R. (ed.) Potential Impacts from Climate Change on Freshwater Invertebrates, including Mollusks, in the Deschutes, Klamath, and John Day Basins of Eastern Oregon: 19 - 37.","Strong E. E. & Frest T. 2007. On the anatomy and systematics of Juga from western North America (Gastropoda: Cerithioidea: Pleuroceridae). The Nautilus 121: 43 - 65.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Campbell D. C., Clark S. A., Johannes E. J., Lydeard C. & Frest T. J. 2016. Molecular phylogenetics of the freshwater gastropod genus Juga (Cerithioidea: Semisulcospiridae). Biochemical Systematics and Ecology 65: 158 - 170. https: // doi. org / 10.1016 / j. bse. 2016.01.004","Johnson P. D., Bogan A. E., Brown K. M., Burkhead N. M., Cordeiro J. R., Garner J. T., Hartfield P. D., Lepitzki D. A. W., Mackie G. L., Pip E., Tarpley T. A., Tiemann J. S., Whelan N. V. & Strong E. E. 2013. Conservation status of freshwater gastropods of Canada and the United States. Fisheries 38: 247 - 282. https: // doi. org / 10.1080 / 03632415.2013.785396","Strong E. E. & Whelan N. V. 2019. Assessing the diversity of Western North American Juga (Semisulcospiridae, Gastropoda). Molecular Phylogenetics and Evolution 136: 87 - 103. https: // doi. org / 10.1016 / j. ympev. 2019.04.009","Frest T. J. & Johannes E. J. 1993. Mollusc species of Special Concern within the range of the Northern Spotted Owl. Final report to Forest Ecosystem Management Working Group, USDA Forest Service. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 1996. Additional information on certain mollusk species of special concern occurring within the range of the Northern Spotted Owl. Report to Oregon State Office, BLM. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 1999. Field guide to survey and manage freshwater mollusk species. United States Department of the Interior; Bureau of Land Management, Oregon State office; United States Fish and Wildlife Service, Northwest Regional Ecosystems Office; and United States Department of Agriculture, Forest Service, Region 6, Portland, Oregon. BLM / OR / WA / PL- 99 / 045 + 1792.","Strong E. E. & Kohler F. 2009. Morphological and molecular analysis of ' Melania' jacqueti Dautzenberg and Fischer, 1906: from anonymous orphan to critical basal offshoot of the Semisulcospiridae (Gastropoda: Cerithioidea). Zoologica Scripta 38: 483 - 502. https: // doi. org / 10.1111 / j. 1463 - 6409.2008.00385. x","Graf D. L. 2001. The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of Recent North America, north of Mexico. Walkerana 12: 1 - 124.","Tryon G. W. 1864. Synonymy of the species of Strepomatidae, a family of fluviatile Mollusca inhabiting North America. Part 2. Proceedings of the Academy of Natural Sciences of Philadelphia 16: 35 - 59.","Tryon G. W. 1865. Review of the goniobases of Oregon and California. American Journal of Conchology 1: 236 - 246.","Tryon G. W. 1866. Monograph of the family Strepomatidae. American Journal of Conchology 2: 14 - 52.","Tryon G. W. 1873. Land and Fresh-water Shells of North America Part IV. Strepomatidae (American Melanians). Smithsonian Institution, Washington, D. C.","Pilsbry H. A. 1899. Mollusks collected by R. C. McGregor in northern California. The Nautilus 13: 64 - 67.","Roscoe E. J. 1963. Some goniobases in western United States. The Nautilus 77: 43 - 47.","Henderson J. 1936 b. The non-marine Mollusca of Oregon and Washington. Supplement. University of Colorado Studies 23: 251 - 280.","Duncan N. 2008. Survey Protocol for Aquatic Mollusk Species: Preliminary Inventory and Presence / Absence Sampling, Version 3.1. Portland, OR. Interagency Special Status / Sensitive Species Program. U. S. Department of Interior, Bureau of Land Management, Oregon / Washington and U. S. Department of Agriculture, Forest Service, Region 6.","Fallon C., Hietala-Henschell K. & Jordan S. F. 2020. Interagency Special Status / Sensitive Species Program (ISSSSP) Species Fact Sheet: Juga newberryi. USDA Forest Service Region 6 and USDI Bureau of Land Management Oregon State Office. Available from https: // www. fs. usda. gov / r 6 / [accessed 13 Mar. 2022]."]}

“ Melania wahlamatensis Lea ” This was correctly listed as a nomen nudum (Frest & Johannes 2010), but only on the authority of Graf (2001), who attributed it to Binney (1860) while noting that Hannibal (1912: 178) gave the authority as “‘Lea,’ Carpenter, 1857 (nude name)”. Indeed, the first mention we can find is Carpenter (1857: 211, 325), who listed it from the Sacramento River and credited it to a draft manuscript of Gould on the U.S. Exploring Expedition shells. Given the locality, it would be referable to J. occata, not a “form of plicifera ” as speculated by Frest & Johannes (2010). The name was listed by Taylor (1975a) as Juga wahlamatensis with no indication that it was not an available name., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 83, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Graf D. L. 2001. The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of Recent North America, north of Mexico. Walkerana 12: 1 - 124.","Binney W. G. 1860. No. 6. Fluviatile Gasteropoda. In: Check lists of the shells of North America. Prepared for the Smithsonian Institution by Isaac Lea, P. P. Carpenter, W. M. Stimpson, W. G. Binney, and Temple Prime. Smithsonian Miscellaneous Collections 2: 7 - 13. https: // doi. org / 10.5962 / bhl. title. 18985","Hannibal H. 1912. A synopsis of the recent and Tertiary freshwater Mollusca of the Californian province, based upon an ontogenetic classification. Proceedings of the Malacological Society of London 10 (2): 112 - 166; 10 (3): 167 - 211. Available from https: // www. biodiversitylibrary. org / page / 15237075 [accessed 13 Mar. 2022].","Carpenter P. P. 1857. Report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Report of the British Association for the Advancement of Science 1856: 159 - 368. https: // doi. org / 10.5962 / bhl. title. 60613","Taylor D. W. 1975 a. Index and bibliography of Late Cenozoic freshwater Mollusca of western North America. Claude W. Hibbard Memorial Volume 1. Papers on Paleontology (Museum of Paleontology, University of Michigan) 10: 1 - 384."]}

Juga caerulea sp. nov. urn:lsid:zoobank.org:act: B1DFDF9D-9B8B-471B-822A-80F281835610 Figs 12–13 Oxytrema bulbosa – Morrison 1954: 361, 384, pl. 11 fig. 5. Juga (Oreobasis) n. sp. 5 – Frest & Johannes 1995a: 181. — Johannes 2015: 27. Juga (Oreobasis) OTU 4 – Campbell et al. 2016: 160. Blue Mountains Juga – Strong & Whelan 2019: 89. Etymology ‘ Caeruleus -a -um ’, Latin for ‘blue’, in reference to the Blue Mountains where it is found, and to the bluish to purplish appearance of the aperture interior of fresh collected shells. Type material Holotype USA • 1 spm (L = 15.02 mm); Oregon, Grant County, Phipps Meadow along U.S. Hwy 26; 44.5816, -118.4419; 10 Jul. 2014; J.T. Garner and N.V. Whelan leg.; GenBank: MK472115, MK464487, MK480730; USNM 1294988. Other material examined 13 lots, 66 specimens, of which 11 were sequenced. USA – Oregon • 37 spms; Grant Co., 10 mi E of Austin, Blue Mountains, stream, John Day River drainage; [44.5987, -118.3364]; alt. 4000 ft; 25 May 1939; V.B. Scheffer leg.; USNM 473801 • 18 spms; Grant Co., Phipps Meadow along U.S. Hwy 26; 44.5816, -118.4419; 10 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295000 • 1 spm; same collection data as for preceding; GenBank: MK472116, MK464488; USNM 1294989 • 1 spm; same collection data as for preceding; GenBank: MK472117, MK464489, MK480731; USNM 1294990 • 1 spm; same collection data as for preceding; GenBank: MK472118, MK464490, MK480732; USNM 1294991 • 1 spm; same collection data as for preceding; GenBank: MK472119, MK464491; USNM 1294992 • 1 spm; same collection data as for preceding; GenBank: MK472120, MK464492; USNM 1294993 • 1 spm; same collection data as for preceding; GenBank: MK472121, MK464493; USNM 1294994 • 1 spm; same collection data as for preceding; GenBank: MK472122, MK464494, MK480733; USNM 1294995 • 1 spm; same collection data as for preceding; GenBank: MK472123, MK464495, MK480734; USNM 1294996 • 1 spm; same collection data as for preceding; GenBank: MK472124, MK464496, MK480735; USNM 1294997 • 1 spm; same collection data as for preceding; GenBank: MK472125, MK464497, MK480736; USNM 1294998 • 1 spm; same collection data as for preceding; GenBank: MK472126, MK464498; USNM 1294999. Description SHELL. Thin, moderate in size, reaching ~ 1.5 cm in length; turriform, conical in shape, spire height moderate (Fig. 12). Whorls convex, suture moderately impressed. Aperture oval in shape, lip smooth, simple to slightly sinuous. Spiral sculpture of indistinct, fine striae, variable in strength and number. Axial sculpture of fine, orthocline to weakly opisthocyrt or sinuous growth lines; plications lacking. Shell dark brown to black, occasionally with a lighter subsutural band. Interior of aperture bluish in color, occasionally tinged with purple. RADULA. Rachidian broadly rectangular, wider than tall, with convex lower margin and projecting outer corners; basal denticles lacking or vestigial (Fig. 13). 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 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 half to two-thirds of shafts; broad outer flanges extending almost to tooth bases. Inner marginal teeth with ~four and outer marginal teeth with ~five flattened denticles. Distribution and ecology In springs and spring runs of the John Day River drainage, northeastern Oregon (Fig. 14A). Remarks This species is characterized by its simple, smooth, dark, lightly constructed shell, without distinguishing sculpture; the only known populations are rather uniform in shape and size. Distribution Juga caerulea sp. nov. is a highly disjunct and restricted species, occurring in springs and spring runs from a few sites within a single spring complex in Phipps Meadow, the headwaters of the Middle Fork of the John Day River in the Malheur National Forest. One lot collected in 1939 (USNM 473801) indicates that historically it may have occurred in small streams in the area; Morrison (1954: 361, 384, pl. 11 fig. 5) identified these specimens as Oxytrema bulbosa and provided a sketch of the head-foot of a female to show the egg laying groove and ovipositor. However, recent efforts to locate the species elsewhere in the Blue Mountains have been unsuccessful (Frest & Johannes 1995a). Phipps Meadow, the source of the type material, has been heavily affected by grazing and Frest & Johannes (1995a) considered existing surveys were sufficient to merit a status of Endangered at the Federal or State level. The Blue Mountain Land Trust recently purchased Phipps Meadow with the goal of protecting and restoring the meadow’s riparian habitat (https://bmlt.org/news/phipps-meadow). Common name The common name for this species is the Blue Mountain Juga (Johannes 2015). It has also been referred to as the Blue Mountains Juga (Frest & Johannes 1995a; Duncan 2008; Strong & Whelan 2019)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on pages 42-45, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Morrison J. P. E. 1954. The relationships of Old and New World melanians. Proceedings of the United States National Museum 103: 357 - 394. https: // doi. org / 10.5479 / si. 00963801.103 - 3325.357","Frest T. J. & Johannes E. J. 1995 a. Interior Columbia Basin mollusk species of special concern. Final report: Interior Columbia Basin Ecosystem Management Project, Walla Walla, WA. Contract 43 - 0 E 00 - 4 - 9112.","Johannes E. J. 2015. Potential impacts from climate change on freshwater mollusks in the Deschutes, Klamath, and John Day basins. In: Wisseman R. (ed.) Potential Impacts from Climate Change on Freshwater Invertebrates, including Mollusks, in the Deschutes, Klamath, and John Day Basins of Eastern Oregon: 19 - 37.","Campbell D. C., Clark S. A., Johannes E. J., Lydeard C. & Frest T. J. 2016. Molecular phylogenetics of the freshwater gastropod genus Juga (Cerithioidea: Semisulcospiridae). Biochemical Systematics and Ecology 65: 158 - 170. https: // doi. org / 10.1016 / j. bse. 2016.01.004","Strong E. E. & Whelan N. V. 2019. Assessing the diversity of Western North American Juga (Semisulcospiridae, Gastropoda). Molecular Phylogenetics and Evolution 136: 87 - 103. https: // doi. org / 10.1016 / j. ympev. 2019.04.009","Duncan N. 2008. Survey Protocol for Aquatic Mollusk Species: Preliminary Inventory and Presence / Absence Sampling, Version 3.1. Portland, OR. Interagency Special Status / Sensitive Species Program. U. S. Department of Interior, Bureau of Land Management, Oregon / Washington and U. S. Department of Agriculture, Forest Service, Region 6."]}

Juga acutifilosa (Stearns, 1890) Figs 8–9 Melania (? Goniobasis) acutifilosa Stearns, 1890: 211–212, pl. 15 fig. 9. Goniobasis laurae Goodrich, 1944: 2–3, fig. 1. Syn. nov. Goniobasis interioris Goodrich, 1944: 3–4, fig. 2. Syn. nov. Juga (Calibasis) acutifilosa acutifilosa – Burch & Tottenham 1980: 152, fig. 450. — Burch 1982: 41, fig. 450; 1989: 152, fig. 450. Juga (Oreobasis) laurae – Burch & Tottenham 1980: 154, fig. 467. — Burch 1982: 42, fig. 467; 1989: 154, fig. 467. — Frest & Johannes 2010: 10, 38. — Campbell et al. 2016: 160. Juga (Oreobasis) interioris – Burch & Tottenham 1980: 154, fig. 466. — Burch 1982: 42, fig. 466; 1989: 154, fig. 466. — Frest & Johannes 2010: 10, 38. — Campbell et al. 2016: 160. Juga acutifilosa – Turgeon et al. 1988: 65; 1998: 67. — Johnson et al. 2013: 282. — Johannes & Clark 2016: 23–24. Juga laurae – Turgeon et al. 1988: 65; 1998: 67. — Johnson et al. 2013: 282. Juga interioris – Turgeon et al. 1988: 65; 1998: 67. — Johnson et al. 2013: 282. Juga (Calibasis) acutifilosa – Frest & Johannes 1993: 61 (in part); 1995b: 37 (in part); 2000a: 283 (in part); 2005: 160 (in part); 2010: 9, 22, fig. 2a (in part). — Strong & Frest 2007: 51. — Campbell et al. 2016: 160, fig. 2a. Juga (Calibasis) n. sp. 1 – Frest & Johannes 1995b: 38 (in part). Juga (Calibasis) OTU 1 – Campbell et al. 2016: 160. Juga (Calibasis) OTU 2 – Campbell et al. 2016: 160. Juga acutifilosa group – Strong & Whelan 2019: 89. non Juga interioris – Lee et al. 2006: 316. — Ó Foighil et al. 2009: 305. [= Juga douglasi sp. nov.] non Juga (Calibasis) acutifilosa – Furnish 2007: 14, fig. 6, above left. [= Juga occata] non Juga acutifilosa – Johnson et al. 2013: pl. 4. [= Elimia arachnoidea] Material examined Lectotype of Melania (? Goniobasis) acutifilosa Stearns, 1890 (designated by Strong & Frest 2007: 51) (Fig. 8A) USA • “ Eagle Lake, California ”; Jun. 1877; H.W. Henshaw leg.; USNM 60596. Paralectotypes of Melania (? Goniobasis) acutifilosa Stearns, 1890 USA • 14 spms; same collection data as for lectotype; USNM 1665625 (formerly USNM 60596X) • 25 spms; same collection data as for lectotype; USNM 1665626 (formerly USNM 60596). Paratypes of Goniobasis laurae Goodrich, 1944 (Fig. 8B) USA • 10 spms; “ Spring west of Home Camp, Long Valley, Washoe County, Nevada ”; 1934; C.L. Hubbs leg.; MCZ 221159 • 1 spm; “ In springs of Grasshopper Valley, Lassen County, California ”; 1942; C.L. Hubbs leg.; SBMNH 35483. Remarks Holotype (UMMZ 160002, “Spring west of Home Camp, Long Valley, Washoe County, Nevada ”. 1934. C.L. Hubbs leg.); 100 spms, paratypes (UMMZ 63452, “ Spring west of Home Camp, Long Valley, Washoe County, Nevada ”; 1934; C.L. Hubbs leg.), possibly lost (Taehwan Lee pers. com.); and 75 spms, paratypes (UMMZ 63453, “ In Boulder Springs, Long Valley ”; 1934; C.L. Hubbs leg.) not examined. Paratypes of Goniobasis interioris Goodrich, 1944 USA • 2 spms; “ Badger Creek, Bitner Ranch, Washoe County, Nevada ”; 1942; C.L. Hubbs leg.; MCZ 221163 (Fig. 8C) • 1 spm; same collection data as for preceding; SBMNH 35482. Remarks Holotype (UMMZ 160005, “ Badger Creek, Bitner Ranch, Washoe County, Nevada ”; 1942; C.L. Hubbs leg.) and 1 spm, paratype (UMMZ 160007, “ Outlet of artesian wells 9 miles west of Gerlach, Washoe County, Nevada ”; 1942; C.L. Hubbs leg.) not examined. Other material examined 83 lots, 3896 specimens, of which 43 were sequenced. USA – Oregon • 24 spms; Lake Co., adjacent to Coleman Lake (dry); 42.0698, -119.8405; 11 Jul. 2014; J.T. Garner and N.V. Whelan leg.; USNM 1295054 • 1 spm; same collection data as for preceding; GenBank: MK472270, MK464638, MK480797; USNM 1295053 • 1 spm; same collection data as for preceding; GenBank: MK472261, MK464629; USNM 1295044 • 1 spm; same collection data as for preceding; GenBank: MK472262, MK464630, MK480795; USNM 1295045 • 1 spm; same collection data as for preceding; GenBank: MK472263, MK464631; USNM 1295046 • 1 spm; same collection data as for preceding; GenBank: MK472264, MK464632; USNM 1295047 • 1 spm; same collection data as for preceding; GenBank: MK472265, MK464633; USNM 1295048 • 1 spm; same collection data as for preceding; GenBank: MK472266, MK464634; USNM 1295049 • 1 spm; same collection data as for preceding; GenBank: MK472267, MK464635; USNM 1295050 • 1 spm; same collection data as for preceding; GenBank: MK472268, MK464636; USNM 1295051 • 1 spm; same collection data as for preceding; GenBank: MK472269, MK464637, MK480796; USNM 1295052 • 4 spms; Lake Co., Stone Corral; [42.6206, -119.6913]; C.C. Engberg leg.; USNM 363039. – California • 218 spms; Lassen Co., Ash Creek, 2600′ E, 2500′ S, 33-38N-11E, NK 10-9 A-3; [41.0903, -120.7133]; 4 Jul. 1972; D.W. Taylor leg.; CASIZ 183594 • 151 spms; Lassen Co., Ash Creek, 2600’W, 1500’N, 4-38N-10E, NK 10-9 A-4; [41.1592, -120.8272]; 30 Aug. 1981; D.W. Taylor leg.; CASIZ 183597 • 6 spms; Lassen Co., Ash Creek, east side of FS R39N50, at Dan Ryan Place; [41.1499, -120.8176]; 18 Aug. 2007; T. Grace leg.; USNM 1111962 • 12 spms; Lassen Co., Ash Creek, Modoc Natl Forest, Dan Ryan Place, E of FS39N50; 41.1499, -120.8176; 18 Aug. 2007; T. Grace leg.; UF 520305 • 6 spms; same collection data as for preceding; UF 520310 • 4 spms; Lassen Co., Ash Creek, NW of Ash Creek Campground, just below FS22; 41.1604, -120.8292; 18 Aug. 2007; T. Grace leg.; UF 520306 • 49 spms; Lassen Co., Big Spring; [40.7574, -120.1233]; 3 Nov. 1986; D. Sada leg.; USNM 858178 • 9 spms; Lassen Co., Eagle Lake [sic, likely Willow Creek]; [40.6043, -120.6936]; Horn leg.; USNM 59145 • 192 spms; Lassen Co., in spring in Murrers Lower Meadow, approx. 2 mi E of Eagle Lake; [40.5897, -120.6993]; 28 Aug. 1979; J. Landye and L. Eng leg.; on rocks; USNM 892378 • 14 spms; Lassen Co., Modoc Natl Forest, Route 139 Willow Creek, NE side of road, just N of the end of Lower McBride Springs and ca 0.64 km E of Hayden Hill Rd; 41.0176, -120.8379; 16 Sep. 2007; T. Grace leg.; USNM 1111797 • 21 spms; Lassen Co., Modoc Natl Forest, Willow Creek, off Route 139 and just upstream from Hayden Hill Rd (Lassen CR 534, FS 37N42) junction, along W side; 41.0192, -120.8517; 16 Sep. 2007; T. Grace leg.; USNM 1111791 • 3 spms; Lassen Co., N end of Murrers Lower Meadow, S-most of 3 springs and E of Eagle Lake; 40.5848, -120.6959; 10 Sep. 1993; T. Frest and E. Johannes leg.; USNM 1100659 • 248 spms; Lassen Co., Sellicks Springs; [40.5668, -120.321]; 27 Aug. 1979; J. Landye and L. Eng leg.; on rocks and sand; USNM 892370 • 42 spms; Lassen Co., spring in Murrer’s Lower Meadow; 40.5875, -120.6973; 10 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413158 • 1 spm; same collection data as for preceding; GenBank: MK472276, MK464644; USNM 1413152 • 1 spm; same collection data as for preceding; GenBank: MK472277, MK464645, MK480803; USNM 1413153 • 1 spm; same collection data as for preceding; GenBank: MK472278, MK464646; USNM 1413154 • 1 spm; same collection data as for preceding; GenBank: MK472279, MK464647; USNM 1413155 • 1 spm; same collection data as for preceding; GenBank: MK472280, MK464648; USNM 1413156 • 1 spm; same collection data as for preceding; GenBank: MK472281, MK464649; USNM 1413157 • 103 spms; Lassen Co., Spring near Susanville, drains into Honey Lake [possibly Cady Spring]; [40.4158, -120.7106]; W.H. Dall leg.; 4700 ft; USNM 118564 • 17 spms; Lassen Co., Wendel, Secret Valley; [40.3495, -120.2341]; Henshaw leg.; USNM 198875 • 35 spms; Lassen Co., Willow Creek; 10 Oct. 1959; D.W. Taylor leg.; USNM 1151932 • 107 spms; Lassen Co., Willow Creek at Hayden Hill Rd; 41.0202, -120.8526; 10 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413078 • 1 spm; same collection data as for preceding; GenBank: MK472271, MK464639, MK480798; USNM 1413073 • 1 spm; same collection data as for preceding; GenBank: MK472272, MK464640, MK480799; USNM 1413074 • 1 spm; same collection data as for preceding; GenBank: MK472273, MK464641, MK480800; USNM 1413075 • 1 spm; same collection data as for preceding; GenBank: MK472274, MK464642, MK480801; USNM 1413076 • 1 spm; same collection data as for preceding; GenBank: MK472275, MK464643, MK480802; USNM 1413077 • 40 spms; Lassen Co., Willow Creek off dirt road to east, and 0.4 km S of Murrers Lower Meadow; 40.5722, -120.6963; 10 Sep. 1993; T. Frest and E. Johannes leg.; USNM 1100658 • 239 spms; Lassen Co., Willow Creek, S of Adin on Hwy 139 near Willow Creek campground; [41.0131, -120.826]; 31 Jul. 1983; J. Landye leg.; USNM 1152451 • 6 spms; Lassen Co., Willow Fork; Ex. + Surv. W of 100 Merid.; USNM 198955 a • 7 spms; same collection data as for preceding; USNM 198955 • 22 spms; Modoc Co., Ash Creek, Modoc Natl Forest, S of FS39N50 bridge; 41.1499, -120.8176; 8 Oct. 2007; T. Grace leg.; UF 520304 • 14 spms; Modoc Co., Likely, W side of Modoc CR 63, 2.6 mi W of Hwy 395, Smokey Charley Spring run, 0.24 km E of source; 1360 m a.s.l.; [41.2248, -120.5533]; 18 Aug. 2007; T. Grace leg.; on stones and among weeds; USNM 1111961. _ Nevada • 12 spms; Humboldt Co., Cabin Springs, Area B, W of cabins 150 m; 30 Aug. 1979; J. Landye et al. leg.; on stones; USNM 892377 • 115 spms; Humboldt Co., Little Smoky Creek No. 1; [41.2082, -119.3235]; 16 Jun. 1991; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 874224 • 411 spms; Humboldt Co., Little Smoky Creek No. 3; [41.2082, -119.3235]; 16 Jun. 1991; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 874222 • 20 spms; Humboldt Co., Paiute Canyon, Upper Paiute Creek; [39.7772, -119.5049]; 9 Aug. 1991; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 874271 • 222 spms; Washoe Co., Big Hole Spring, 8 mi NW of Gerlach; [40.7076, -119.4692]; 6 Aug. 1991; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 874273 • 35 spms; Washoe Co., Bitner Ranch; 41.7364, -119.469; 9 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413195 • 1 spm; same collection data as for preceding; GenBank: MK472282, MK464650; USNM 1413189 • 1 spm; same collection data as for preceding; GenBank: MK472283, MK464651, MK480804; USNM 1413190 • 1 spm; same collection data as for preceding; GenBank: MK472284, MK464652, MK480805; USNM 1413191 • 1 spm; same collection data as for preceding; GenBank: MK472285, MK464653; USNM 1413192 • 1 spm; same collection data as for preceding; GenBank: MK472286, MK464654; USNM 1413193 • 1 spm; same collection data as for preceding; GenBank: MK472287, MK464655, MK480806; USNM 1413194 • 405 spms; Washoe Co., Boulder Reservoir Spring; [41.3462, -119.7473]; 2 Aug. 1991; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 874265 • 211 spms; Washoe Co., Deep Hole Spring, 9 mi WNW of Gerlach; [40.7211, -119.4828]; 30 Aug. 1979; J. Landye et al. leg.; in areas of flow on rocks and vegetation; USNM 892369 • 34 spms; Washoe Co., Divine Spring run near Home Camp; 41.354, -119.854; 9 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413202 • 1 spm; same collection data as for preceding; GenBank: MK472288, MK464656; USNM 1413196 • 1 spm; same collection data as for preceding; GenBank: MK472289, MK464657, MK480807; USNM 1413197 • 1 spm; same collection data as for preceding; GenBank: MK472290, MK480808; USNM 1413198 • 1 spm; same collection data as for preceding; GenBank: MK472291, MK464658, MK480809; USNM 1413199 • 1 spm; same collection data as for preceding; GenBank: MK472292, MK464659; USNM 1413200 • 1 spm; same collection data as for preceding; GenBank: MK472293, MK464660, MK480810; USNM 1413201 • 23 spms; Washoe Co., Hays Canyon Range, E side of range, unnamed spring; 3 Aug. 1991; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 874267 • 22 spms; Washoe Co., Long Valley, 1 mi NE of Middle Lake, unnamed spring; [41.7801, -119.6705]; 12 Nov. 1992; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 854634 • 339 spms; Washoe Co., N of Little High Rock Reservoir, unnamed spring; [41.2545, -119.4295]; 17 Jun. 1991; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 874220 • 15 spms; Washoe Co., North Little High Rock Canyon Wilderness, Black Rock Desert; 41.2594, -119.4344; 29 Sep. 2011; D. Sada leg.; USNM 1239164 • 1 spm; same collection data as for preceding; GenBank: MK472257; USNM 1239160 • 1 spm; same collection data as for preceding; GenBank: MK472255, MK464625; USNM 1239158 • 1 spm; same collection data as for preceding; GenBank: MK472259; USNM 1239162 • 1 spm; same collection data as for preceding; GenBank: MK472254, MK464624, MK480791; USNM 1239156 • 1 spm; same collection data as for preceding; GenBank: MK472251, MK464621, MK480788; USNM 1239153 • 1 spm; same collection data as for preceding; GenBank: MK472253, MK464623, MK480790; USNM 1239155 • 1 spm; same collection data as for preceding; GenBank: MK472258, MK464627, MK480793; USNM 1239161 • 1 spm; same collection data as for preceding; GenBank: MK472260, MK464628, MK480794; USNM 1239163 • 1 spm; same collection data as for preceding; GenBank: MK472252, MK464622, MK480789; USNM 1239154 • 1 spm; same collection data as for preceding; GenBank: MK472256, MK464626, MK480792; USNM 1239159 • 396 spms; Washoe Co., Wall Canyon Reservoir, 3 mi above canyon, unnamed spring; [41.2048, -119.7822]; 2 Aug. 1991; Great Basin Spring Snail Project; G. Vinyard leg.; USNM 874269. Description SHELL. Thin to moderately thick, large, reaching ~ 2.85 cm in length; turriform, narrowly conical to cylindrical in shape, spire height moderate to tall (Fig. 8). Whorls somewhat flattened to convex or angulate with a subsutural ramp, suture weakly to deeply impressed. Aperture oval in shape, lip smooth to crenate, simple to slightly sinuous. Spiral sculpture absent or present, of thickened, scalloped lirae, barely elevated to prominent, highly variable in number and sinuosity, dividing whorl into intervening flattened areas or grooves. Axial sculpture of fine, orthocline, or weakly opisthocyrt to sinuous growth lines; plications lacking. Shell black, brown or reddish brown, rarely yellowish green in color, occasionally with a lighter subsutural band; bands otherwise lacking. Interior of aperture light brown to white in color. RADULA. Rachidian narrowly rectangular, wider than tall, basal margin slightly concave between bluntly rounded median and lateral projections; basal denticles lacking or vestigial (Fig. 9). Upper margin slightly concave with cutting edge bearing central, elongate conical cusp, and two stout, conical denticles on each side. Lateral teeth with prominent triangular cusp flanked by two inner and three to four outer, triangular denticles, and occasionally 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; broad outer flanges extending almost to tooth bases. Inner marginal teeth with five and outer marginal teeth with six flattened denticles. See also Strong & Frest (2007). Distribution and ecology In springs, spring runs, and spring-fed creeks from Modoc and Lassen Counties in the upper Pit River drainage in northern California and in Great Basin drainages from northeastern California (Lassen County, Honey Lake basin), southeastern Oregon (Lake County), and northwestern Nevada, primarily in Washoe County, and adjacent parts of western Humboldt County (Fig. 7B). Remarks The type material of Melania acutifilosa was collected during the geographical surveys of the territory of the United States west of the 100 th meridian (Wheeler 1879). The type locality is “Eagle Lake” (Stearns 1890), later corrected by Taylor (1981) to head of Willow Creek without justification. Strong & Frest (2007) followed this, explaining that no populations are currently found in Eagle Lake or in the small tributaries that flow into the lake during winter. Moreover, specimens of Juga have not been recorded from the lake itself during subsequent collecting efforts (e.g., by J. Maillard in 1923; Hanna 1924) nor from drift and dredged material (Frest & Johannes 2010) but have been reported from several unnamed springs and creeks nearby in collections made by R.C. McGregor in 1898 (Pilsbry 1899; in the collections of the ANSP). Frest & Johannes (2010) noted that Taylor (1981) did not distinguish between the “headwaters” or “Murrer’s Meadows” as the source of the types. Given their view that different taxonomical species are found in the headwaters versus further downstream, they concluded the lirate types were most likely from the head of Willow Creek. This is likely correct, as Willow Spring at the head of Willow Creek in Murrer’s Upper Meadow was the site of an altitude measurement taken during the survey (Wheeler 1879: 130) and lies roughly two miles to the east of the shore of Eagle Lake. Frest & Johannes (2010) considered USNM 198955 from “Willow Fork” also collected during the expedition as possible topotypes; the lot was annotated “Eagle Lake vicinity” by J.P.E. Morrison who separated the original lot into lirate and smooth forms (USNM 198955 and USNM 198955a), which are here considered conspecific. This is a highly variable species, ranging from lirate to smooth forms, and like J. canella sp. nov. with which this species has been frequently confused, these forms can co-occur at a single site. Juga laurae and J. interioris have been considered valid species in recent classifications (Burch & Tottenham 1980; Burch 1982, 1989; Turgeon et al. 1988, 1998; Frest & Johannes 2010; Johnson et al. 2013). However, molecular data (Strong & Whelan 2019) show that topotypic specimens (Fig. 8F–G) of these two nominal species are conspecific with disjunct populations from adjacent parts of Oregon and California, including from the type locality of M. acutifilosa (Fig. 8D–E), and their synonymy with J. acutifilosa here is new. Juga laurae and J. interioris were established by Goodrich (1944), who found them so wanting in definite characters of the shell, operculum, and radula that he was at first hesitant to describe them as new (Goodrich 1935), but later upon doing so, felt unable to ascertain their affinities (Goodrich 1944: 1). Despite statements to the contrary (Frest & Johannes 2010: 3, Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on pages 26-33, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Stearns R. E. C. 1890. Scientific results of explorations by the U. S. Fish Commission steamer Albatross. XVII. Descriptions of new West American land, freshwater, and marine shells, with notes and comments. Proceedings of the United States National Museum 13: 205 - 225. https: // doi. org / 10.5479 / si. 00963801.13 - 813.205","Goodrich C. 1944. Pleuroceridae of the Great Basin. Occasional Papers of the Museum of Zoology, University of Michigan 485: 1 - 11.","Burch J. B. & Tottenham J. 1980. North American freshwater snails, species list, ranges, and illustrations. Walkerana 3: 1 - 215.","Burch J. B. 1982. Freshwater snails (Mollusca: Gastropoda) of North America. Environmental Monitoring and Support Laboratory, Office of Research and Development, United States Environmental Protection Agency, Cincinnati, Ohio, EPA- 600 / 3 - 82 - 026.","Burch J. B. 1989. North American Freshwater Snails. Malacological Publications, Hamburg, Michigan.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Campbell D. C., Clark S. A., Johannes E. J., Lydeard C. & Frest T. J. 2016. Molecular phylogenetics of the freshwater gastropod genus Juga (Cerithioidea: Semisulcospiridae). Biochemical Systematics and Ecology 65: 158 - 170. https: // doi. org / 10.1016 / j. bse. 2016.01.004","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1988. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 26.","Turgeon D. D., Quinn J. F., Bogan A. E., Coan E. V., Hochberg F. G. et al. 1998. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks. Second Edition. American Fisheries Society Special Publication 26.","Johnson P. D., Bogan A. E., Brown K. M., Burkhead N. M., Cordeiro J. R., Garner J. T., Hartfield P. D., Lepitzki D. A. W., Mackie G. L., Pip E., Tarpley T. A., Tiemann J. S., Whelan N. V. & Strong E. E. 2013. Conservation status of freshwater gastropods of Canada and the United States. Fisheries 38: 247 - 282. https: // doi. org / 10.1080 / 03632415.2013.785396","Johannes E. J. & Clark S. A. 2016. Freshwater mollusc declines, local extinctions and introductions in five northern California streams. Tentacle 24: 22 - 25.","Frest T. J. & Johannes E. J. 1993. Mollusc species of Special Concern within the range of the Northern Spotted Owl. Final report to Forest Ecosystem Management Working Group, USDA Forest Service. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 1995 b. Freshwater mollusks of the Upper Sacramento System, California, with particular reference to the Cantara Spill. 1994 yearly report to California Department of Fish and Game. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 2000 a. A baseline mollusk survey of southwestern Oregon, with emphasis on the Rogue and Umpqua River drainages. Final report prepared for Oregon Natural Heritage Program. Portland, Oregon. Deixis Consultants, Seattle, Washington.","Frest T. J. & Johannes E. J. 2005. Springsnails of the Cascade-Siskiyou National Monument and vicinity, Oregon. 2004 report. Final report prepared for World Wildlife Fund, Ashland, Oregon. Deixis Consultants, Seattle, Washington.","Strong E. E. & Frest T. 2007. On the anatomy and systematics of Juga from western North America (Gastropoda: Cerithioidea: Pleuroceridae). The Nautilus 121: 43 - 65.","Strong E. E. & Whelan N. V. 2019. Assessing the diversity of Western North American Juga (Semisulcospiridae, Gastropoda). Molecular Phylogenetics and Evolution 136: 87 - 103. https: // doi. org / 10.1016 / j. ympev. 2019.04.009","Lee T., Kim J. J., Hong H. C., Burch J. B. & O Foighil D. 2006. Crossing the continental divide: the Columbia drainage species Juga hemphilli (Henderson, 1935) is a cryptic member of the eastern North American genus Elimia (Cerithioidea: Pleuroceridae). Journal of Molluscan Studies 72: 314 - 317. https: // doi. org / 10.1093 / mollus / eyl 005","O Foighil D., Lee T., Campbell D. C. & Clark S. A. 2009. All voucher specimens are not created equal: A cautionary tale involving North American pleurocerid gastropods. Journal of Molluscan Studies 75: 305 - 306. https: // doi. org / 10.1093 / mollus / eyp 033","Furnish J. L. 2007. Guide to sensitive aquatic mollusks of the U. S. Forest Service Pacific Southwest Region. USDA Forest Service, Pacific Southwest Region, Vallejo, CA.","Wheeler G. M. 1879. Annual report upon the geographical surveys of the territory of the United States west of the 100 th meridian, in the states and territories of California, Colorado, Kansas, Nebraska, Nevada, Oregon, Texas, Arizona, Idaho, Montana, New Mexico, Utah, Washington, and Wyoming: being Appendix OO of the annual report of the Chief of Engineers for 1879. Govt. Print. Off., Washington. https: // doi. org / 10.3133 / 70039940","Taylor D. W. 1981. Freshwater mollusks of California: a distributional checklist. California Fish and Game 67: 140 - 163.","Hanna G. D. 1924. Freshwater mollusks of Eagle Lake, California. Proceedings of the California Academy of Sciences, 4 th Series 13: 131 - 136.","Pilsbry H. A. 1899. Mollusks collected by R. C. McGregor in northern California. The Nautilus 13: 64 - 67.","Goodrich C. 1935. A species of Goniobasis new to the Great Basin. The Nautilus 49: 66."]}

Juga douglasi sp. nov. urn:lsid:zoobank.org:act: 8EA208D9-9CE9-4521-8381-7411439C8E1F Figs 17–18 Juga (Oreobasis) nigrina – Frest & Johannes 1995b: 38 (in part). Juga (Oreobasis) “ nigrina ” – Frest & Johannes 1995b: 39 (in part). Juga interioris – Lee et al. 2006: 316. — Ó Foighil et al. 2009: 305. Juga n. spp. – Frest & Johannes 2007: table 4. Juga (Oreobasis) OTU 2 – Campbell et al. 2016: 160. Juga (Oreobasis) OTU 3 – Campbell et al. 2016: 160. Juga species 1 – Strong & Whelan 2019: 89, fig. 4g –l. Etymology Named in honor of the father of the first author, Douglas Wayne Strong. A native Californian, Doug shared with his family his deep appreciation for the natural beauty of northern California, and always enjoyed hearing about the snails that also make their home amongst the majestic redwoods. Type material Holotype USA • 1 spm (L = 15.74 mm); California, Shasta County, Goose Valley Spring; 40.9295, -121.726; 11 Sep. 2015; E.E. Strong and P. Bouchet leg.; GenBank: MK472138, MK464510, MK480745; USNM 1413130. Other material examined 64 lots, 818 specimens, of which 19 were sequenced. USA – California • 13 spms; Alameda Co., Strawberry Creek, Berkeley; [37.871, -122.2635]; 18 Aug. 1926; E. Caruthers leg.; ANSP 145847 • 1 spm; Butte Co., Paradise; [39.7522, -121.6143]; J. Rowell leg.; ANSP 105056 • 51 spms; Butte Co., spring on Chico Creek [possibly Richardson Springs] tributary of Sacramento River; [39.8395, -121.7798]; Henshaw leg.; USNM 128206 • 5 spms; Butte Co., Table Mountain; [39.5831, -121.6192]; W.M. Gabb leg.; ANSP 27568 • 6 spms; same collection data as for preceding; USNM 119284 • 9 spms; El Dorado Co., small stream 24.5 mi E of Placerville; [38.7737, -120.3949]; 10 May 1946; M.L. Walton leg.; ANSP 346017 • 9 spms; Lassen Co., Bob Creek, at crossing of Little Valley Rd (CR 404), Beaver Creek Ranch, 5.5 mi NW of Little Valley; 40.9467, -121.2534; 1 Sep. 2007; T. Grace leg.; USNM 1111865 • 20 spms; Lassen Co., Lassen Natl Forest, Davis Spring, upstream ca 1.5 mi from FS 22 crossing, ca 19 air mi WNW of Spalding; [40.8233, -121.1026]; 15 Jun. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; CASIZ 165974 • 27 spms; Nevada Co., 1 mi NW of North San Juan, small stream between South and North Forks of the Yuba River on Hwy 49 [= Clear Creek]; [39.3792, -121.091]; Jun. 1971; D.D. Chivers leg.; CASIZ 30310 • 8 spms; Nevada Co., Tahoe National Forest, East Fork Creek, 1 mi S of Washington [sic, possibly East Fork Washington Creek]; 22 Aug. 1997; J. Vindum and C. Spencer leg.; CASIZ 113099 • 2 spms; Nevada Co., Tahoe Natl Forest, Sfork of Yuba River, E of Washington along Maybert Rd, Golden Quartz picnic area; [39.356, -120.7466]; 28 Aug. 1997; E.J. Kools leg.; on rocks; CASIZ 113100 • 30 spms; Nevada Co., Tahoe Natl Forest, Washington Creek, 1 mi S of Washington; 39.3446, -120.7948; 28 Aug. 1997; E.J. Kools leg.; CASIZ 112152 • 3 spms; Placer / El Dorado Counties, Middle Fork American River; H. Moores leg.; USNM 118854 • 20 spms; Plumas Co., Butterfly Creek, near Keddie; [40.0071, -120.9624]; Chace and H.N. Lowe leg.; ANSP 212242 • 24 spms; Plumas Co., Domingo Springs, off CR Route 311 (approx. 7 air mi NW of Chester); [40.3609, -121.3473]; 19 Jun. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; J. Brim Box leg.; CASIZ 165953 • 10 spms; Plumas Co., East Fork of Feather River [East Branch North Fork]; [40.0193, -121.1048]; 1898; R.C. McGregor leg.; ANSP 73510 • 5 spms; Plumas Co., Howells [possibly Howells Bridge, along East Branch North Fork Feather River, Belden]; [40.0142, -121.2258]; 29 Aug. 1946; H.P. Chandler leg.; CASIZ 170000 • 8 spms; Plumas Co., Indian Valley; [40.1186, -120.8791]; USNM 58981 • 68 spms; Plumas Co., Lassen Natl Forest, Soldier Creek, off FS Rd 28N36, ca 7 air mi SSW of Chester; [40.2123, -121.2717]; 25 Jun. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; J. Brim Box leg.; CASIZ 165371 • 3 spms; Sacramento Co., spring on Placerville Rd, ca 23 mi E of Placerville; [38.774, -120.4271]; 1916; H.N. Lowe leg.; ANSP 114784 • 14 spms; Shasta Co., Battle Creek 10 mi above mouth, tributary to Sacramento River; [40.4147, -122.0988]; 1898; R.C. McGregor leg.; ANSP 73522 • 30 spms; Shasta Co., Blackberry Creek, Shasta NF, N side of FS50, 5.87 km E of Pit 5 Dam; 40.9669, -121.8284; 16 Sep. 2007; T. Grace leg.; UF 520318 • 23 spms; Shasta Co., Goose Valley Spring; 40.9295, -121.726; 11 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413139 • 1 spm; same collection data as for preceding; GenBank: MK472137, MK464509; USNM 1413129 • 1 spm; same collection data as for preceding; GenBank: MK472139, MK464511; USNM 1413131 • 1 spm; same collection data as for preceding; GenBank: MK472140, MK464512, MK480746; USNM 1413132 • 1 spm; same collection data as for preceding; GenBank: MK472141, MK464513, MK480747; USNM 1413133 • 1 spm; same collection data as for preceding; GenBank: MK472142, MK464514, MK480748; USNM 1413134 • 1 spm; same collection data as for preceding; GenBank: MK472143, MK464515; USNM 1413135 • 1 spm; same collection data as for preceding; GenBank: MK472144, MK464516; USNM 1413136 • 1 spm; same collection data as for preceding; GenBank: MK472145, MK464517; USNM 1413137 • 1 spm; same collection data as for preceding; GenBank: MK472146, MK464518; USNM 1413138 • 12 spms; Shasta Co., Goose Valley, unnamed spring run on W side of Goose Valley Rd and Goose Valley, ca 0.8 km N of Goose Creek, approx. 4.5 mi N of Route 299; 40.9329, -121.7264; 16 Sep. 2007; T. Grace leg.; USNM 1111796 • 15 spms; Shasta Co., Grace Lake [near Shingletown]; [40.4833, -121.8636]; 9 Nov. 1963; CASIZ 57480 • 30 spms; Shasta Co., Hat Creek at Bridge Campground; 40.73, -121.4371; 10 Sep. 2015; E.E. Strong and P. Bouchet leg.; USNM 1413102 • 1 spm; same collection data as for preceding; GenBank: MK472127, MK464499; USNM 1413092 • 1 spm; same collection data as for preceding; GenBank: MK472128, MK464500, MK480737; USNM 1413093 • 1 spm; same collection data as for preceding; GenBank: MK472129, MK464501; USNM 1413094 • 1 spm; same collection data as for preceding; GenBank: MK472130, MK464502, MK480738; USNM 1413095 • 1 spm; same collection data as for preceding; GenBank: MK472131, MK464503, MK480739; USNM 1413096 • 1 spm; same collection data as for preceding; GenBank: MK472132, MK464504, MK480740; USNM 1413097 • 1 spm; same collection data as for preceding; GenBank: MK472133, MK464505, MK480741; USNM 1413098 • 1 spm; same collection data as for preceding; GenBank: MK472134, MK464506, MK480742; USNM 1413099 • 1 spm; same collection data as for preceding; GenBank: MK472135, MK464507, MK480743; USNM 1413100 • 1 spm; same collection data as for preceding; GenBank: MK472136, MK464508, MK480744; USNM 1413101 • 6 spms; Shasta Co., Lamoine, side creek going into Sacramento River; [40.9753, -122.4311]; Aug. 1937; T. Burch and J.Q. Burch leg.; UF 192866 • 5 spms; Shasta Co., Lost Creek, in lava field on N of Wilcox Ranch Rd, W of Hat Creek rim, E of Hat Creek and Hwy 44/89, lower fishing access.; 40.7628, -121.4347; 12 Aug. 2007; T. Grace leg.; USNM 1111863 • 23 spms; Shasta Co., Pit River Canyon (small spring), W side of river 4 mi below Fall River Mills; [40.9851, -121.5126]; C.H. Merriam leg.; USNM 361546 • 108 spms; Shasta Co., Potem Creek, in tributary to Pit River after falls by road; [40.8368, -122.0222]; 17 May 1978; J. Landye leg.; USNM 892366 • 3 spms; Shasta Co., Rainbow Springs, at headwaters of Fall Mill Creek on Lazy S Ranch adjacent to Thousand Springs Ranch; [41.1111, -121.5511]; 16 May 1978; J. Landye leg.; USNM 1665622 (ex USNM 892373) • 1 spm; Shasta Co., S side of Hwy 299, Pit River, Lion’s Club picnic area, W of Pit River camping grounds; 40.9866, -121.513; 1 Sep. 2007; T. Grace leg.; USNM 1665623 (ex USNM 1111864) • 14 spms; Shasta Co., S side of Pit River, water source spring at Pacific Gas and Electric plant; [40.9896, -121.5021]; 16 May 1978; J. Landye leg.; USNM 1665624 (ex USNM 892372) • 18 spms; Shasta Co., Salt Creek Rd, 1 km S of Fenders Ferry Rd [Salt Ck]; 40.8646, -122.1395; 360 m a.s.l.; 18 Oct. 2011; J. Slapcinsky leg.; UF 448882 • 35 spms; Shasta Co., tributary to Shasta Reservoir, 4.7 road mi from McCloud Bridge, E of McCloud arm [= Nosoni Creek]; [40.9113, -122.2078]; 1 Jul. 1997; J.L. Furnish leg.; CASIZ 223387 • 1 spm; Sierra Co., Downieville, Fiddle Creek [sic]; 20 Sep. 1946; H.P. Chandler leg.; CASIZ 170002 • 26 spms; Sierra Co., North Fork, Yuba River, Downieville; [39.5583, -120.8275]; ANSP 125085 • 1 spm; Siskiyou Co., headwaters of Fall River; 1896; R.C. McGregor leg.; ANSP 332330 • 17 spms; Tehama Co., Deer Creek, off FS Rd 28N29, ca 10 air mi NNW of Forest Ranch; [40.0702, -121.7048]; 25 Jul. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; J. Brim Box leg.; CASIZ 165961 • 24 spms; Tehama Co., Deer Creek, off Potato Patch Campground, off SR 32, ca 12 air mi SSE of Mineral; [40.1873, -121.5323]; 23 Jul. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; CASIZ 166030 • 7 spms; Tehama Co., Deer Creek, off SR 32, 0.5 mi N of Alder Creek Campground, ca 11 air mi SE of Mineral; [40.2163, -121.4917]; 23 Jul. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; CASIZ 166361 • 21 spms; Tehama Co., Lassen Natl Forest, Deer Creek, at lower falls, off FS Rd 27N08, approx. 13 air mi S of Mineral; [40.168, -121.5814]; 24 Jul. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; J. Brim Box leg.; CASIZ 165372 • 15 spms; Tehama Co., unnamed spring off [Tehama] County Rd 769, ca 0.4 mi E of SR 36/89, ca 13 air mi WNW of Chester; [40.3406, -121.4605]; 21 Jun. 2001; USFS Lassen Natl Forest Mollusk Survey 2001; CASIZ 166013 • 26 spms; Tehama Co., W Branch of Feather River, Morgan Springs; [40.3846, -121.5137]; 1898; R.C. McGregor leg.; ANSP 73512 • 2 spms; Shaw Creek, at Fire Creek Fire Control Station, Shasta Co. [sic, possibly Shaw Creek, Tehama Co.]; [40.2096, -121.9511]; 17 May 1978; J. Landye leg.; USNM 892381. Description SHELL. Thin, moderate in size, reaching ~ 2 cm in length; turriform, conical in shape, spire height moderate (Fig. 17). Whorls convex, suture moderately impressed. Aperture oval in shape, lip smooth, simple to weakly sinuous. Spiral sculpture of numerous, indistinct, fine striae. Axial sculpture of fine, weakly prosocline to opisthocyrt or sinuous growth lines; irregular varices occasionally present confined to upper spire. Shell light brown to dark purplish- or reddish-brown, occasionally with a lighter subsutural band. Interior of aperture light, yellowish brown to dark purple. RADULA. Rachidian narrowly rectangular, wider than tall, basal margin flattened, with slight, bluntly rounded median and lateral projections; basal denticles lacking or vestigial (Fig. 18). 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 ~three inner and three to five 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 half to third of inner marginal teeth and extending almost to tooth bases of outer marginal teeth; broad outer flanges extending almost to tooth bases. Inner marginal teeth with ~four and outer marginal teeth with ~five flattened denticles. Distribution and ecology Primarily in springs, spring runs, and small creeks in the southern Pit River system; possibly extending into larger creeks and rivers in eastern Sacramento River tributaries and the northern Mokelumne River watershed of the Sierra Nevada (Fig. 14C). Remarks Given the uncertainty of the circumscription of this species in the southern part of its range (see below), the preceding description is based on sequenced and topotypic specimens. Although not present in all specimens, the species may be distinguished by the presence of irregular varices on the upper whorls of some specimens (e.g., Fig. 17V, X–Y). These can be seen in the sequenced holotype and topotypic specimens (Fig. 17A, U) and are more irregular and widely spaced than the plications seen in J. nigrina with which it overlaps in distribution in the western part of its range (see J. nigrina below). Distribution The main part of the distribution of J. douglasi sp. nov. lies in the southern Pit River drainages in Shasta and Lassen Counties where it overlaps with J. occata, although individuals of the two species are rarely found in syntopy; only three lots with mixtures of the two were found in the USNM collections [USNM 1111864, USNM 1665623 (ex USNM 1111864); USNM 892372, USNM 1665624 (ex USNM 892372); USNM 892373, USNM 1665622 (ex USNM 892373)], and one species is always numerically dominant. Sequenced specimens reported in Campbell et al. (2016) expand the distribution into the eastern Sacramento River tributaries of southern Lassen County and northern Plumas County. Specimens consistent with the J. douglasi sp. nov. morphotype from McCloud River (Nosoni Creek; CASIZ 223387) and upper Sacramento River drainages (Lamoine; UF 192866) extend the distribution slightly to the west where it overlaps with J. nigrina, but this should be confirmed with sequenced specimens. Populations from Sacramento River drainages of the western foothills of the Sierra Nevada have been attributed primarily to J. nigrina (e.g., Lowe 1916; Taylor 1981; Frest & Johannes 1995b; Brim Box 2002; Brim Box et al. 2005), rarely to J. interioris (e.g., Lee et al. 2006; Ó Foighil et al. 2009), and more recently were noted to comprise at least one new species (Frest & Johannes 2007). These are difficult to identify with certainty in the absence of molecular data but are here cautiously interpreted as representing J. douglasi sp. nov. When preserved, the early teleoconch is typically smooth, narrow, with convex whorls and a deeply impressed suture, and sometimes (CASIZ 165961, CASIZ 166013) shows the varices seen more frequently in the northwest part of the range. The teleoconchs are usually smooth, but occasionally spirally grooved or with indistinct spiral striae and are rarely banded. If these records can be verified as conspecific with J. douglasi, scattered museum records place the species, at least historically, in eastern Sacramento River drainages in Butte, Sierra, Nevada, El Dorado, and Sacramento Counties in the Feather River, Yuba River, and American River Basins. These areas coincide with the Sierra Nevada goldfields and were heavily affected by placer mining activities including hydraulic mining and dredging. However, modern museum records from the mid 1990’s and surveys by Frest & Johannes (2007; as Juga n. spp.) confirm that Juga is surviving in highly fragmented populations in the southern Lassen National Forest, the Plumas National Forest, and the Tahoe National Forest in Nevada County (CASIZ 112152, 113099, 113100). A 2018 iNaturalist record (https://www.inaturalist.org/observations/15172449) from the Cosumnes River, El Dorado County, indicates that Juga populations are persisting in the Mokelumne River watershed, which borders the Sacramento River watershed to the south. One historical lot (ANSP 145847) from Strawberry Creek on the UC Berkeley campus in the East San Francisco Bay Area contains spirally grooved specimens (Fig. 17T) similar to some from Table Mountain in Butte County [USNM 119284 (Fig. 17Q), ANSP 27568] and North San Juan in Nevada County [CASIZ 30310 (Fig. 17R, Z)] and is hypothesized also to be conspecific with J. douglasi. However, given the range of morphological variation seen in species of Juga, it is possible that populations from the Sierra Nevada represent mixtures of J. occata and J. douglasi, or an as yet undescribed form. Common name The common name for this species is the Douglas Juga. The OTU referred to as J. (O.) nigrina (Frest & Johannes 1995b = Black Juga) from the Upper Sacramento System is a mixture of true J. nigrina and J. douglasi sp. nov. as circumscribed here. The OTU referred to as J. (O.) “ nigrina ” (Frest & Johannes 1995b = Smooth River Juga) from the Upper Sacramento System is a mixture of J. canella sp. nov. and J. douglasi as circumscribed here.

Goniobasis columbiensis Whiteaves, 1905 Frest & Johannes (2010) treated this nominal species elsewhere in their text but omitted it from their list of taxa incorrectly assigned to Juga. It was described from “headwaters of the Columbia River, at Upper Columbia Lake, in the East Kootenay District of British Columbia ” (Whiteaves 1905: 61–62, pl. 2 figs 11–12). Attempts to recollect it at the type locality and nearby sites were unsuccessful (Frest & Johannes 2010), and it is known only from the type material (Fig. 27A). It is considered a mislocalized specimen of the eastern North American pleurocerid, Elimia livescens (Menke, 1830) (Walker 1918; Goodrich 1937, 1942; LaRocque 1953; Frest & Johannes 2010)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 81, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Whiteaves J. F. 1905. Description of a new species of Goniobasis, from British Columbia. The Nautilus 19: 61 - 62.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Walker B. 1918. A synopsis of the classification of the fresh-water Mollusca of North America, North of Mexico, and a catalogue of the more recently described species, with notes. Miscellaneous Publications of the University of Michigan Museum of Zoology 6: 1 - 213. https: // doi. org / 10.5962 / bhl. title. 46279","Goodrich C. 1937. Goniobasis columbiensis Whiteaves. The Nautilus 50: 82 - 84. Goodrich C. 1942. The Pleuroceridae of the Pacific coastal drainage, including the Western Interior Basin. Occasional Papers of the Museum of Zoology, University of Michigan 469: 1 - 4.","Goodrich C. 1942. The Pleuroceridae of the Pacific coastal drainage, including the Western Interior Basin. Occasional Papers of the Museum of Zoology, University of Michigan 469: 1 - 4.","LaRocque A. 1953. Catalogue of the Recent Mollusca of Canada. National Museum of Canada, Bulletins 129: 1 - 406."]}

“ Juga striata Lea ” One of the species originally included in the subgenus Juga by Adams& Adams (1854), Frest & Johannes (2010) correctly indicated the status of Melania striata I. Lea, 1841 as a junior homonym [non Melania striata Sowerby, 1814 (nomen dubium); Melania striata Perry, 1811; Melania striata Schumacher, 1817], for which Lea (1842) himself provided the replacement name, Melania striatula I. Lea, 1842. However, the current taxonomic placement is Elimia striatula (Pleuroceridae) (Burch & Tottenham 1980; Burch 1982, 1989; Johnson et al. 2013; Bieler 2021), not E. perstriata perstriata (I. Lea, 1852) as suggested by Frest & Johannes (2010). The latter was the placement of “ Goniobasis striatula Lea ” of Wheeler (1912) (see Goodrich 1930: 17), which Graf (2001) reported as a nomen nudum., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 83, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Lea I. 1842. Description of new fresh water and land shells. Transactions of the American Philosophical Society 8: 163 - 250 [+ 1 p. contents, 1 p. corrigenda]. https: // doi. org / 10.2307 / 1005238","Burch J. B. & Tottenham J. 1980. North American freshwater snails, species list, ranges, and illustrations. Walkerana 3: 1 - 215.","Burch J. B. 1982. Freshwater snails (Mollusca: Gastropoda) of North America. Environmental Monitoring and Support Laboratory, Office of Research and Development, United States Environmental Protection Agency, Cincinnati, Ohio, EPA- 600 / 3 - 82 - 026.","Burch J. B. 1989. North American Freshwater Snails. Malacological Publications, Hamburg, Michigan.","Johnson P. D., Bogan A. E., Brown K. M., Burkhead N. M., Cordeiro J. R., Garner J. T., Hartfield P. D., Lepitzki D. A. W., Mackie G. L., Pip E., Tarpley T. A., Tiemann J. S., Whelan N. V. & Strong E. E. 2013. Conservation status of freshwater gastropods of Canada and the United States. Fisheries 38: 247 - 282. https: // doi. org / 10.1080 / 03632415.2013.785396","Bieler R. 2021. Isaac Lea's (1792 - 1886) substitutions and other modifications of his own names of molluscan species. Malacologia 64: 1 - 56. https: // doi. org / 10.4002 / 040.064.0101","Wheeler H. E. 1912. The Mollusca of Monte Sano, Alabama. The Nautilus 25: 121 - 127. Available from https: // www. biodiversitylibrary. org / page / 1818312 [accessed 19 Oct. 2022].","Goodrich C. 1930. Goniobases of the vicinity of Muscle Shoals. Occasional Papers of the Museum of Zoology, University of Michigan 209: 1 - 25.","Graf D. L. 2001. The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of Recent North America, north of Mexico. Walkerana 12: 1 - 124."]}

“ Goniobasis plicifera var. leai Hemphill, 1890 ” This was correctly listed by Frest & Johannes (2010) as a nomen nudum, but only on the authority of Coan & Roth (1987). This name was listed in Hemphill’s (1890: 18) catalogue of North American shells for sale as originating from “ Oregon ”. Taylor (1975a) included it as Juga leai with no indication that it was not an available name. Wu & Brandauer (1982) listed “cotypes” UCM 17779 and UCM 17780 collected by H. Hemphill from the Willamette River in Oregon, which have no type status (Coan & Roth 1987). Given the locality, the specimens would be referable to J. plicifera as circumscribed here, not J. (J.) plicifera oregonensis as suggested by Frest & Johannes (2010)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 81, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Hemphill H. 1890. Catalogue of North American Shells Collected and for Sale by Henry Hemphill. Published by the author, San Diego, CA. https: // doi. org / 10.5962 / bhl. title. 21154","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Coan E. & Roth B. 1987. The malacological taxa of Henry Hemphill. The Veliger 29: 322 - 339. Available from https: // www. biodiversitylibrary. org / page / 43061570 [accessed 13 Mar. 2022].","Taylor D. W. 1975 a. Index and bibliography of Late Cenozoic freshwater Mollusca of western North America. Claude W. Hibbard Memorial Volume 1. Papers on Paleontology (Museum of Paleontology, University of Michigan) 10: 1 - 384.","Wu S. - K. & Brandauer N. E. 1982. Type specimens of Recent Mollusca in the University of Colorado Museum. Natural History Inventory of Colorado 7: 1 - 47."]}

Melania buschiana Reeve, 1860 This species was established by Reeve (1860) based on material said to be from California. Carpenter (1872) included it on his list of mollusks from western North America and noted that it resembled the young of Melania scipio Gould, 1847. As noted by Graf (2001), Tryon (1873: 340) included it on his list of “Doubtful and Spurious Species” and considered it to be of East Indian provenance. Both species are now in the synonymy of Stenomelania aspirans (Hinds, 1844) (Thiaridae) (Starmühlner 1993). This species was omitted by Frest & Johannes (2010)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 81, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Carpenter P. P. 1872. The mollusks of Western North America. Smithsonian Miscellaneous Collections 10: 1 - 446. https: // doi. org / 10.5962 / bhl. title. 31923","Gould A. A. 1847. Descriptions of the following species of Melania, from the collection of the Exploring Expedition. Proceedings of the Boston Society of Natural History 2: 222 - 225. https: // doi. org / 10.5962 / bhl. part. 3322","Graf D. L. 2001. The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of Recent North America, north of Mexico. Walkerana 12: 1 - 124.","Tryon G. W. 1873. Land and Fresh-water Shells of North America Part IV. Strepomatidae (American Melanians). Smithsonian Institution, Washington, D. C.","Hinds R. B. 1844. Descriptions of new species of Melania collected during the voyage of H. M. S. Sulphur. The Annals and Magazine of Natural History 14: 8 - 11. https: // doi. org / 10.1080 / 037454809495124","Starmuhlner F. 1993. Ergebnisse der osterreichischen Tonga-Samoa Expedition 1985 des Instituts fur Zoologie der Universitat Wien: Beitrage zur Kenntnis der Suss- und Brackwasser-Gastropoden der Tonga- und Samoa-Inseln (SW-Pazifik). Annalen des Naturhistorischen Museums in Wien, Serie B 94 / 95: 217 - 306.","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61."]}

“ Juga obruta Lea ” One of the species originally included in the subgenus Juga by Adams & Adams (1854), this was incorrectly listed as a possible nomen nudum by Frest & Johannes (2010). This species was established without a locality as Melania obruta I. Lea & H.C. Lea, 1851 and was placed in the Hemisinidae P. Fischer & Crosse, 1891 as Aylacostoma obrutum by Berning (2020)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 82, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61.","Berning M. I. 2020. Cerithioidea. In: Damborenea C., Rogers F. C. & Thorp J. J. (eds) Thorp and Covich's Freshwater Invertebrates, Ed. 4. Vol. 5: Keys to Neotropical and Antarctic Fauna: 354 - 377. Academic Press. https: // doi. org / 10.1016 / C 2015 - 0 - 01546 - 5"]}

Anculosa fusca Haldeman, 1841 This species was described from “ Oregon ” in the genus Anculosa Say, 1821, a junior synonym of Leptoxis Rafinesque, 1819 (Pleuroceridae), where it was retained by Brot (1862, 1868). It was included by Graf (2001) but from early on (e.g., Cooper 1867) was recognized to belong to the genus Fluminicola, where it remains today (Hershler & Frest 1996). It was omitted by Frest & Johannes (2010)., Published as part of Strong, Ellen E., Garner, Jeffrey T., Johnson, Paul D. & Whelan, Nathan V., 2022, A systematic revision of the genus Juga from fresh waters of the Pacific Northwest, USA (Cerithioidea, Semisulcospiridae), pp. 1-97 in European Journal of Taxonomy 848 on page 81, DOI: 10.5852/ejt.2022.848.1993, http://zenodo.org/record/7427764, {"references":["Brot A. 1862. Materiaux pour servir a l'Etude de la Famille des Melaniens: Catalogue systematique des Especes qui composent la Famille des Melaniens. J-G. Fick, Geneva. https: // doi. org / 10.5962 / bhl. title. 11242","Brot A. 1868. Materiaux pour servir a l'Etude de la Famille des Melaniens. Additions et Corrections au Catalogue systematique des Especes qui composent la Famille des Melaniens. J-G. Fick, Geneva. https: // doi. org / 10.5962 / bhl. title. 11249","Graf D. L. 2001. The cleansing of the Augean Stables, or a lexicon of the nominal species of the Pleuroceridae (Gastropoda: Prosobranchia) of Recent North America, north of Mexico. Walkerana 12: 1 - 124.","Cooper J. G. 1867. Geographical Catalogue of the Mollusca found West of the Rocky Mountains, between Latitudes 33 ° and 49 ° North. Geological Survey of California, San Francisco. https: // doi. org / 10.5962 / bhl. title. 13050","Hershler R. & Frest T. J. 1996. A review of the North American freshwater snail genus Fluminicola (Hydrobiidae). Smithsonian Contributions to Zoology 583: 1 - 41. https: // doi. org / 10.5479 / si. 00810282.583","Frest T. J. & Johannes E. J. 2010 - 2011. Review of the species of Juga (western U. S. Cerithioidea, Pleuroceridae, Semisulcospirinae). Malacological Review 43 / 44: 1 - 61."]}