Juga bulbosa (A. Gould, 1847 )

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 : 33-41

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 17754a .

Remarks 1 paratype ( UCM 17754b, 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 the Columbia River are conspecific with Juga plicifera (e.g., Fig. 5K–M ). Joseph Drayton was the expedition’s artist and travelled up the Columbia Gorge as far as Fort Walla Walla with a Hudson Bay Company boat bringing supplies to the forts up the river ( Smith 1937; Frest & Johannes 2010), and thus the types could have come from anywhere along this route, likely from tributaries of the main stem Columbia in the central to eastern part of the gorge. The paralectotype lot of draytonii from “Walla” [sic, Walla Walla] (USNM 119124) received via Lady K. Douglas lies well to the east of the known range but its provenance is unreliable ( Carpenter 1857: 162).

Intensive sequencing effort along the Columbia Gorge has revealed only two species ( Strong & Whelan 2019), Juga plicifera and a second species that Strong & Whelan (2019) referred to as Juga species 2 and for which we consider Melania bulbosa to be the earliest available name. The type locality of M. bulbosa , vaguely attributed to the “Columbia River”, is not in conflict with this interpretation. Although considered valid in most historical taxonomical treatments (see Frest & Johannes 2010; table 2), the erroneous synonymy of M. newberryi with bulbosa (see J. newberryi , below), resulted in enormous confusion in the application of the name and most usages of 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) (see below). This confusion was caused, at least in part, by the paucity of historical material in museum collections, and what little there is in some cases contains mixtures of the two species. For example, the paralectotype lot MCZ 169068 is a mixture of bulbosa and newberryi , as is USNM 5563 which contains Gould’s (1860: pl. 10 fig. 163–163a) figured specimen of bulbosa ( Fig. 10E–F ). Strong & Frest (2007) removed newberryi from the synonymy of bulbosa , leaving the status of the latter somewhat unresolved as a likely valid species; this was followed by Frest & Johannes (2010) who reported that they had been unable to recollect this form anywhere in the Deschutes River drainage or Columbia Gorge; accordingly, Campbell et al. (2016) reported they could not assess its status. Johnson et al. (2013) considered bulbosa a valid species and it was indicated to be endangered under AFS criteria and critically imperiled under NatureServe’s global conservation status ranks but tallied as extinct ( Johnson et al. 2013: table 1) ( Campbell et al. 2016; Campbell 2019).

After comprehensive review of the type material in comparison to the morphology of sequenced specimens, we consider the smooth, unbanded lectotypes of Melania bulbosa ( Fig. 10A ) and Goniobasis draytonii ( Fig. 10C ) to fall within the range of variation of Juga species 2 sensu Strong & Whelan (2019) in size, proportion, profile, and sculpture. However, populations precisely matching the distinctive morphology of the types of bulbosa have not been recollected thus far ( Strong & Frest 2007; Frest & Johannes 2010) and may no longer be extant. Frest & Johannes (2010: 26) noted the similarity of the types of bulbosa and of draytonii but distinguished the latter on the basis of its more elongate spire and more numerous whorls after decollation. Frest & Johannes (1993, 1995a, 2001) considered that their J. (J.) n. sp. 1 (Brown Juga ) could match Lea’s types of draytonii , but their composite concept of the species was distributed from the central and western Columbia Gorge in both Oregon and Washington, including the Mt. Hood National Forest, and thus would include individuals of both J. bulbosa and J. plicifera as circumscribed here. The specimen sequenced as J. (J.) draytonii by Campbell et al. (2016) placed in the same clade with J. (J.) plicifera plicifera ; however, the morphology of the type material of draytonii is consistent with the interpretation that it is a synonym of J. bulbosa . Even the comparatively smooth forms of J. plicifera in small Columbia River tributaries from Washington (e.g., Fig. 5H–J ) can be readily distinguished from J. bulbosa by their larger size at adulthood, more cylindrical shape, higher spire, and more shouldered whorls. In Columbia River tributaries on the Oregon side, plicifera tends to be quite large and heavily sculptured (e.g., Fig. 5N–O, R–S ) and that much more distinct from bulbosa . When preserved, the plications in bulbosa are less conspicuous and confined to the upper spire.

Juga bulbosa also includes what has been referred to as J. (Juga) bairdiana by Frest & Johannes (2010) and Campbell et al. (2016). With its plicate early whorls, bairdiana has been viewed as a synonym of plicifera ( Pilsbry 1899; Walker 1918) or silicula ( Goodrich 1942) or as a subspecies of silicula ( Henderson 1935a, 1936b), but sequenced specimens morphologically identical to the type of bairdiana (e.g., Fig. 10H ) ( Strong & Whelan 2019) confirm it is conspecific with bulbosa as circumscribed here. Near topotypic specimens ( Fig. 10I–J ) from the Deschutes River of what has been referred to as J. (J.) hemphilli maupinensis ( Burch & Tottenham 1980; Burch 1982, 1989; Frest & Johannes 2010; Campbell et al. 2016) are also conspecific with bulbosa .

Distribution

As mentioned, well localized, historical specimens of this species are uncommon in museum collections. In the USNM, apart from the material collected for this study, it is represented by four lots collected between 1971 and 1983. The remaining 13 lots, including the type material, were collected in the early 1900’s or earlier and are all poorly or mislocalized: six lots are localized simply to “ Oregon ”, and one lot said to be from the Willamette River (USNM 321825) is mislocalized. Another lot from Sullivan’s Island in the Columbia River (USNM 468428) was catalogued in late 1936 with no collecting date and occurs slightly farther west than the species’ known range, but is comprised of bleached and worn, deadcollected specimens that may have been transported. Goniobasis bulbosa was erroneously reported by Pilsbry (1899) to occur in the Owyhee River, a Snake River tributary, based on a mislocalized specimen record in the ANSP (ANSP 27561), an error perpetuated until recently (e.g., Burch & Tottenham 1980; Burch 1982, 1989; see Frest & Johannes 1995a). This lot was interpreted as nigrina sensu lato by Frest & Johannes (2010) but is a mixed lot of bulbosa and newberryi . Surveys in the Owyhee River drainage and from archeological sites in this drainage have not produced any Juga ( Frest & Johannes 2010: 26) . A lot cited by Henderson (1929, 1936b; UCM 15814) from Ahtanum Creek, Union Gap, in Yakima County, Washington, that he attributed to Goniobasis draytonii and G. rubiginosa , appears to be J. bulbosa as circumscribed herein, and which Frest & Johannes (2010: 35) stated to resemble specimens from the Klickitat Plateau, Washington. One lot of 50 specimens in the ANSP database, from Union Gap in the Yakima area, could not be found (P. Callomon, pers. com.). Attempts to recollect Juga in this area were unsuccessful ( Frest & Johannes 1995a: 246). Campbell et al. (2016: fig. 1) show a specimen record from the Imnaha River, a Snake River tributary in the northeastern corner of Oregon, coded with the same pattern as the point from Ahtanum Creek. Frest and Johannes (2010: 41) have claimed there are no Juga present in any Snake River tributaries, and the map published in their 2010 work ( Frest & Johannes 2010: fig. 1) does not include this point. Campbell et al. (2016) do not provide any information about the basis for this record, and we are unaware of any specimens in museum collections from this drainage.

Common name

The common name for this species is the Bulb Juga ( Frest & Johannes 2010; Johnson et al. 2013; Johannes 2015). Synonyms have been referred to as the Three-Band Juga [= Juga (Juga) n. sp. 2], the One-Band Juga, Brown Juga ( J. draytonii / J. (J.) n. sp. 1), and the Deschutes or Purple-Lipped Juga (= J. hemphilli maupinensis ) (e.g., Frest & Johannes 1993, 1995a, 2001, 2010; Furnish 2005; Duncan 2008; Johannes 2015). Juga bulbosa also includes what has been referred to as the Basalt Juga [= Juga (Oreobasis) n. sp. 2] ( Frest & Johannes 1999; Duncan 2008; Johannes 2015), which has been afforded federal protection under the “survey and manage” provisions of the Northwest Forest Plan ( USDA and USDI 1994, 2000) and petitioned for federal listing under the U.S. Endangered Species Act ( USFWS 2011). However, the 12-month finding on the petition concluded that the species could not be considered a listable entity pending the necessary genetic comparisons to establish its taxonomic validity ( USFWS 2012). Given placement of the Basalt Juga as a synonym of this wider-ranging species, a listing decision can now be made but will require consideration of the full range of J. bulbosa .

Figures

Fig. 10. Shell morphology of Juga bulbosa (A. Gould, 1847). A–D. Type material. G–S. Sequenced vouchers. A. Melania bulbosa A. Gould, 1847. Lectotype, MCZ 169067. B. Goniobasis bairdiana I. Lea, 1862. Lectotype, USNM 119069. C. Goniobasis draytonii I. Lea, 1862. Lectotype, USNM 118993. D. Goniobasis hemphilli maupinensis Henderson, 1935. Holotype, UCM 17754a. E. Melania bulbosa. Figured specimen, USNM 5563. F. Melania bulbosa. Original figure (Gould 1860: pl. 10 fig. 163–163a). G. USNM 1413315. H. USNM 1413307. I–J. Near topotypic (Goniobasis hemphilli maupinensis Henderson, 1935). I. USNM 1413300. J. USNM 1413301. K. USNM 1413426. L. USNM 1413186. M. USNM 1413230. N. USNM 1295091. O. USNM 1295055. P. USNM 1413165. Q. USNM 1413458. R. USNM 1413435. S. USNM 1295201. Scale bar = 1 cm.

Fig. 11. Radular morphology of Juga bulbosa (A. Gould, 1847) (USNM 1413186). 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. Scale bars: A = 200 µm; B–C = 100 µm; D–F = 50 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.

Fig. 5 (previous page). Shell morphology of Juga plicifera (I. Lea, 1838). Sequenced vouchers, arranged roughly from North to South. A. Near topotypic (Melania silicula A. Gould, 1847), USNM 1413235. B. USNM 1413244. C. USNM 1413471. D. USNM 1413219. E. USNM 1413170. F. USNM 1413171. G. USNM 1413222. H. USNM 1413183. I. USNM 1413177. J. USNM 1413204. K. USNM 1295003. L. USNM 1295178. M. USNM 1295022. N. USNM 1294958. O. Topotypic (Goniobasis hemphilli dallesensis Henderson, 1935), USNM 1294964. P. USNM 1295070. Q. USNM 1295196. R–S. Near topotypic (Melania plicifera I. Lea, 1838; Goniobasis plicifera var. oregonensis Tryon, 1865) R. USNM 1295147. S. USNM 1295150. T. USNM 1413397. U. USNM 1413028. V. USNM 1413386. W. USNM 1413370. X. USNM 1413409. Y. USNM 1413344. Z. USNM 1294972. A’. Near topotypic (Goniobasis plicifera var. bulimoides Tryon, 1865), USNM 1295172. B’. USNM 1413394. Scale bar = 1 cm.