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).