Juga acutifilosa ( Stearns, 1890 )

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), European Journal of Taxonomy 848, pp. 1-97 : 26-33

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: 38), Taylor (1981) did not mention nor explicitly synonymize the two with J. nigrina in his checklist of freshwater mollusks of California. The range for nigrina , given by Taylor (1981: 149) as “northern California and adjacent parts of Oregon and Nevada ”, might imply inclusion of laurae and interioris , but is ambiguous.

Distribution

The species as redefined herein occurs in the upper Pit River drainage in northern California and Great Basin drainages in northeastern California, southeastern Oregon, and northwestern Nevada. Based on the presence of elevated spiral lirae, J. acutifilosa erroneously has been considered to extend to Siskiyou County, California, and Jackson County in south-central Oregon ( Frest & Johannes 1993, 2000a, 2010; Furnish 2007; Johnson et al. 2013), although Frest & Johannes (2010) acknowledged the species to be composite. As defined by Frest & Johannes (1993, 2000a, 2005, 2010: table 5), J. acutifilosa would contain species referable to J. acutifilosa , J. occata , and J. canella sp. nov. as redefined herein.

USNM 1296706 is a lot of 11 specimens ostensibly collected in Jun. 1948 from Convict Creek, a Great Basin tributary of the Owens River in Mono County, California. This record lies far to the south of the currently accepted distribution of Juga and its provenance is unreliable. A lot from Paiute Canyon in southern Humboldt County, Nevada, collected in 1991 under the auspices of the Great Basin Spring Snail Project, represents a southern extension of the range as reported elsewhere ( Strong & Frest 2007; Frest & Johannes 2010; Campbell et al. 2016). The species is more broadly distributed than previously appreciated and includes populations of smooth, dark morphs attributed to J. interioris and J. laurae from northwestern Nevada.

Common name

The common name for this species is the Scalloped Juga ( Frest & Johannes 1993, 2000a). Turgeon et al. (1988, 1998) incorrectly referred to this species as the Topaz Juga (= J. occata ) which was followed by Frest & Johannes (1995b, 2010) and Johnson et al. (2013). This may have been the result of the erroneous synonymy of Goniobasis acutifilosa pittensis with the frequently reddish-brown J. acutifilosa rather than with J. occata , for which a green- or yellow-brown coloration is more typical ( Frest & Johannes 2010; see J. occata below). Synonyms have been known as the Oasis Juga (for J. laurae ) and Smooth Juga (for J. interioris ) (e.g., Turgeon et al. 1988, 1998; Frest & Johannes 2010; Johnson et al. 2013). As mentioned above, populations from south-central Oregon ( Frest & Johannes 1993, 2000a, 2005) referred to as the Scalloped Juga represent a different species (= Juga canella sp. nov.; see account below). 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). The specimen with a prominent spiral keel figured as J. acutifilosa (USNM 425495) in Johnson et al. (2013), is Elimia arachnoidea (Anthony, 1854) (USNM 28749) from Tennessee.

Figures

Fig. 8. Shell morphology of Juga acutifilosa (Stearns, 1890). A–C. Type material. D–J. Sequenced vouchers. A. Melania acutifilosa Stearns, 1890. Lectotype, USNM 60596. B. Goniobasis laurae Goodrich, 1944. Paratype, MCZ 221159. C. Goniobasis interioris Goodrich, 1944. Paratype, MCZ 221163. D–E. Topotypic (Melania acutifilosa Stearns, 1890). D. USNM 1413153. E. USNM 1413156. F. Topotypic (Goniobasis laurae Goodrich, 1944), USNM 1413201. G. Topotypic (Goniobasis interioris Goodrich, 1944), USNM 1413191. H. USNM 1239161. I. USNM 1295048. J. USNM 1413073. K. USNM 854634. L. USNM 1152451. M. CASIZ 183594. N. CASIZ 183597. O. UF 520310. P. USNM 858178. Q. USNM 874271. R–S. CASIZ 183594. Scale bar = 1 cm.

Fig. 9. Radular morphology of Juga acutifilosa (Stearns, 1890). A. View of the anterior radular ribbon. B. Detail of cutting edge of rachidian and lateral teeth. C. Detail of lateral teeth cusps. D. Detail of rachidian cusps. E. Inner and outer marginal teeth. F. Detail of inner and outer marginal teeth cusps. (D and F modified from Strong & Frest 2007). Scale bars: A = 200 µm; B–C = 100 µm; D = 50 µm; E–F = 20 µm.

Fig. 7. Distribution maps. A. Juga plicifera (I. Lea, 1838). B. Juga acutifilosa (Stearns, 1890). C. Juga bulbosa (A. Gould, 1847). Red stars, type localities; black dots, sequenced specimens; gray dots, unsequenced museum material. Abbreviations: CA = California; NV = Nevada; OR = Oregon; WA = Washington.