Annotated Checklist of Freshwater 

Truncatelloidean Gastropods of the

Western United States, with an

Illustrated Key to the Genera

U.S. Department of the Interior • Bureau of Land Management

Technical Note 449 June 2017
Cover photo: Live specimen of Pyrgulopsis robusta (Snake River, Idaho). Photo by 

Robert Hershler.

Production services provided by the Bureau of Land Management National Operations Center’s 

Information and Publishing Services Section in Denver, Colorado.

Suggested Citation:

Hershler, R. and H.-P. Liu. 2017. Annotated checklist of freshwater truncatelloidean gastropods 

of the western United States, with an illustrated key to the genera. Technical Note 449. 

U.S. Department of the Interior, Bureau of Land Management, National Operations 

Center, Denver, CO.

BLM/OC/ST-17/003+6524
Annotated Checklist of Freshwater 

Truncatelloidean Gastropods of the 

Western United States, with an 

Illustrated Key to the Genera

Technical Note 449

Authors:

Robert Hershler

Department of Invertebrate Zoology

Smithsonian Institution 

Washington, DC 20013-7012

Hsiu-Ping Liu

Department of Biology

Metropolitan State University of Denver

Denver, CO 80217

U.S. Department of the Interior

Bureau of Land Management

June 2017
Amnicola limosa. Photo by Karolyn Darrow.
Technical Note 449 iii

Acknowledgments

Freya Goetz photographed preserved snails and prepared the figures. Dominique 

Ware helped compile the COI sequence data. Jeanette Howard provided useful 

comments on an early draft of this paper. This project was supported (in part) by 

a contract (# 06132013-1651) from The Nature Conservancy. Our manuscript was 

significantly improved by the input from Barry Roth and two anonymous reviewers. 

We also thank Melissa Dickard for her assistance in facilitating those reviews and 

Linda Hill for carefully editing the manuscript.

Acknowledgments
iv Technical Note 449Pyrgulopsis stearnsiana. Photo by Karolyn Darrow.
Technical Note 449 v

Contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Key to the Freshwater Truncatelloidean Genera of the Western United States . . . . . 9

Species Checklist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85

Contents
Potamopyrgus antipodarum. Photo by Karolyn Darrow.
Technical Note 449 vii

Abstract

This technical note provides an illustrated key to the 13 genera and a checklist for

the 178 currently recognized species of freshwater, inland saline, and estuarine 

truncatelloidean gastropods (primarily belonging to the families Amnicolidae, 

Cochliopidae, Hydrobiidae, Lithoglyphidae, and Tateidae) in the western United 

States. It also outlines methods of specimen preparation and dissection to facilitate use 

of the generic key. The checklist includes bibliographic information for each species 

(and their synonyms), type localities, type material, and geographic distributions.

The appendix includes a comprehensive synopsis of the available DNA barcoding 

sequences (the COI gene, cytochrome c oxidase subunit I) for members of this fauna

in the anticipation that these data will be increasingly used for both taxonomic and 

conservation genetic applications. This technical note serves as an entry portal for 

resource managers, taxonomists, and other individuals interested in this species-rich 

and highly imperiled fauna.

Abstract
Pyrgulopisis californiensis. Photo by Karolyn Darrow.
Technical Note 449 1

Introduction

The western United States contains a large, highly endemic fauna of gastropods 

belonging to the superfamily Truncatelloidea (totaling 178 species, Table 1), which has 

recently become a major focus of conservation actions because of the various threats 

to its typically groundwater-dependent habitats (e.g., Pyrgulopsis, Hershler et al. 

2014a). These snails are a challenging group to study because of their diminutive size 

(the shells of most species are <10 mm long) and rather featureless and monotonous 

shells; identification of these animals to useful taxonomic levels usually requires the 

examination of soft parts, particularly the genitalia. In spite of the increasing interest 

from the conservation community, there are no recently published overviews of this 

fauna, which has mushroomed in size (from 33 to 178 species) and otherwise been 

extensively revised taxonomically (e.g., Hershler 1994; Hershler and Frest 1996; 

Hershler 2001) since the last such compilations were published over 30 years ago 

(Taylor 1975; Burch and Tottenham 1980; Burch 1982). The purpose of this technical 

note is to provide a synopsis of the freshwater truncatelloidean snails in the western 

United States that can serve as an entry point for members of the conservation 

community and other biologists interested in studying these long neglected and still 

poorly known animals. The generic key and accompanying illustrations may also assist 

land managers and conservation biologists in their efforts to track the spread of the 

New Zealand mudsnail (Potamopyrgus antipodarum), a highly invasive pest that is 

easily confused with other truncatelloidean snails (ANS Task Force 2005). This 

technical note includes a simple, illustrated key to differentiate the freshwater 

truncatelloidean genera of the western United States, images of representative shells of 

each genus, and an annotated checklist (and bibliography) for all of the currently 

recognized regional species (175 native, 3 introduced). We also provide a detailed 

compilation of published mitochondrial DNA sequences of the “barcoding” gene 

(COI, cytochrome c oxidase subunit I) for the freshwater truncatelloideans in the 

western United States; these data have proved to be an invaluable tool for taxonomic 

studies of these animals and are also beginning to be used to identify conservation 

management units (e.g., Hurt 2004).

Introduction
Photos by Karolyn Darrow.

Amnicola micrococcus

Colligyrus greggi

Eremopyrgus eganensis

Ipnobius robustus

Littoridinops monroensis

Tryonia angulata

Pyrgulopsis owyheensis

Fluminicola minutissimus

Potamopyrgus antipodarum

Pristinicola hemphilliTaylorconcha serpenticola
Technical Note 449 3

Methods

The geographic scope of 

this technical note is the 

conterminous United States 

west of the Continental 

Divide, which includes all of 

Arizona, California, Idaho, 

Nevada, Oregon, Utah, 

Washington, and portions 

of Colorado, Montana, 

New Mexico, and Wyoming 

(Figure 1). We treat all of 

the currently recognized 

freshwater, inland saline, 

and estuarine taxa of the 

caenogastropod superfamily 

Truncatelloidea that have 

been recorded from this 

region, totaling 178 species

(175 native) in 12 genera 

(Table 1). We also include 

one (freshwater) snail of 

uncertain species status 

belonging to a 13th western 

genus. Three species are 

distributed in estuarine 

waters, while 173 species 

live in inland aquatic 

habitats; two species 

(Potamopyrgus antipodarum, 

Tryonia porrecta) are 

distributed in both of these 

habitats. The technical 

note does not include 

the few semiterrestrial 

truncatelloideans (belonging 

to the families Assimineidae 

and Pomatiopsidae) that are 

distributed in the western

United States.

Table 1. Classification of the freshwater truncatelloidean 

gastropods of the western United States.

Family Genus No. of species

AMNICOLIDAE Amnicola 1

Colligyrus 3

Lyogyrus 1*

COCHLIOPIDAE Eremopyrgus 1

Ipnobius 1

Littoridinops 1**

Spurwinkia 1**

Tryonia 14

HYDROBIIDAE Pyrgulopsis 126

LITHOGLYPHIDAE Fluminicola 25

TATEIDAE Potamopyrgus 1**

UNCERTAIN Pristinicola 1

Taylorconcha 2

* Uncertain species status.

** Introduced to the western United States.

Figure 1. The western United States study area, 

consisting of those states or portions of states that are 

west of the Continental Divide (indicated by the red line).

0 180 360 720km

MONTANA

WYOMING

COLORADO

NEW MEXICO

IDAHO

UTAH

ARIZONA

NEVADA

CALIFORNIA

OREGON

WASHINGTON

Kern R. Owens R.

Gila R.

Verde R.

Colorado R.

White R.

Virgin R.

Green R.

Bear R.

Owyhee R.

Snake R.

Salmon R.

Snake R.

Pit R.

Sacramento R.

Humboldt R.

Sevier R.

Columbia R.

Klamath R.

Black R.

Methods
4 Technical Note 449

Generic Key and Checklist

The key to the genera provided herein uses superficial character states that can be 

readily observed using a dissecting microscope (at 50x) and is not structured 

phylogenetically. We designed this key specifically for the distinctive fauna of the 

western United States, and it has limited utility for other geographic areas. We also 

detail methods of specimen preparation and dissection to facilitate the use of this key. 

Morphological terminology largely follows that of Hershler and Ponder (1998).

The checklist follows the recent changes in the higher taxonomy of the Hydrobiidae 

sensu lato (Kabat and Hershler 1993), which has been split into multiple families based 

on both morphologic evidence and a rapidly growing body of molecular evidence 

(Wilke et al. 2001; Wilke et al. 2013). The families are arranged alphabetically in the 

checklist and the genera and species within each family are also listed in this manner. 

For each genus, images of shells of representative species are provided; types are 

illustrated when feasible. For each species (and junior synonyms thereof), the 

reference to the original description consists of the author(s), date, page(s) and figure 

number(s), type locality, and type material. Extralimital (outside the western United 

States) synonyms are not listed for several species. Type localities are listed as in the 

original publication, with comments or corrections given in brackets. The list of 

secondary types may not be complete for some species.

The institutions and museums where the cited types (and other referenced material) 

are housed are identified by the following acronyms: ALBRCIDA, Orma J. Smith 

Museum of Natural History, The College of Idaho, Caldwell, Idaho (formerly 

Albertson College of Idaho); ANSP, Academy of Natural Sciences of Drexel University, 

Philadelphia, Pennsylvania; CAS, California Academy of Sciences, San Francisco, 

California; LACM, Los Angeles County Museum, Los Angeles, California (now the 

Natural History Museum of Los Angeles County); MCZ, Museum of Comparative 

Zoology, Harvard University, Cambridge, Massachusetts; NHMUK, Natural History 

Museum, London, England; NMW, Naturhistorisches Museum Wien, Vienna, Austria; 

SBMNH, Santa Barbara Museum of Natural History, Santa Barbara, California; UF, 

Florida Museum of Natural History, University of Florida, Gainesville, Florida; 

UMMZ, University of Michigan, Museum of Zoology, Ann Arbor, Michigan; USNM, 

National Museum of Natural History, Smithsonian Institution, Washington, DC; 

UTEP, University of Texas at El Paso, El Paso, Texas (now the Centennial Museum and 

Chihuahuan Desert Gardens).

The common name for each species is listed following Turgeon et al. (1998) and 

Johnson et al. (2013); new (common) names are introduced for several recently 

described species. The geographic distribution of each species is given based on the 

most current published information, with references provided when this source 

Methods

postdates the original descriptions. The taxonomy section lists papers that provide 
Technical Note 449 5

illustrations that are useful for identification purposes (excluding original 

descriptions). The remarks section is primarily used to highlight species requiring 

taxonomic revision.

The COI sequences are available (in GenBank) for 160 of the 178 species of freshwater 

truncatelloideans in the checklist, including some taxonomically undescribed 

populations; these sequences are compiled in the appendix. These summaries are 

arranged by genus and species alphabetically within each family and include (for each 

sequence) the species name, published sample codes, accession numbers, number of 

specimens (N), sequence variation (% pairwise distance), most distant conspecific 

sequences (%), closest interspecific sequences (%), and collection locality (including 

county and state). The sequence divergence comparisons are restricted to populations 

and species in the western United States.

Specimen Preparation and Study

Freshwater truncatelloidean snails usually are locally abundant, enabling ready 

collection of sizeable samples (i.e., >100 specimens). A portion of each sample should 

be directly preserved in concentrated (90-100%) nondenatured ethanol; half of these 

specimens can be subsequently (air-) dried and designated as shell vouchers while the 

rest can be retained (in ethanol) for possible DNA analysis. The remaining portion of 

the live sample should be anesthetized (relaxed) with menthol crystals (prior to 

fixation and preservation) to facilitate examination of soft parts required for 

identification. (Menthol is an organic compound obtained from mint plants that is 

readily available in crystalline form from chemical supply houses.) Relaxed material is 

particularly useful for study of the penis, while pertinent details of the female genitalia 

usually can be obtained from contracted specimens that were directly preserved in 

ethanol. Snails should be relaxed in a large container (such as a 1-pint [473-ml] Mason 

jar) that is nearly filled with habitat water and kept cool and out of the sun. A small 

quantity (about half a teaspoon) of powdered menthol crystals (they can be crushed in 

one’s hand) should be sprinkled over the water surface, after which the container 

should be capped and left undisturbed. The snails usually require about 13 hours for 

proper relaxation, although some species (e.g., Pyrgulopsis robusta) may require 

considerably more time. Once the specimens are anesthetized, at which time the 

head-foot is well extended and insensitive to touch, most of the water should be 

decanted and dilute formalin (10% of stock solution) should be slowly added. After 

4-6 hours of fixation, the material should be rinsed and preserved in 70% ethanol.

Alcohol-preserved snails can be quickly separated from their shells by placing them in 

a small quantity of concentrated hydrochloric acid (which dissolves the calcareous 

shell). The appearance of the distal portion of the oviduct—whether it is glandular 

(Figure 2A) or thin-walled and containing brooded young (Figure 2B)—can be readily 

Methods
6 Technical Note 449

determined without dissection. The bursa copulatrix can be viewed by pinning the 

animal, cutting the mantle along the left side of the head-foot, and pulling this tissue 

over (as shown in Figure 2A) to expose the oviduct and associated structures 

(Figure 2C-D). The penis is attached to the “neck” of the snail behind the snout and 

usually extends beyond the mantle edge (Figure 3A-C); both the upper (dorsal) and 

lower (ventral) surfaces of the penis should be examined for glands, which are 

relatively large and quite obvious; the internal penial glands of amnicolids are clearly 

visible in appropriately prepared specimens (Figure 3D). We recommend that workers 

practice the methods of anesthetizing, preserving, and dissecting snails before 

applying them to essential specimens.

Figure 2. Photographs of female snails. A. Oviparous specimen having a thickened, 

glandular oviduct (Pyrgulopsis). Dashed black lines indicate the incisions made 

prior to pulling the tissue over (in the direction of the curved white arrow) to 

expose the left side of the oviduct and the associated structures (shown in C-D). 

B. Ovoviviparous specimen with thin-walled oviduct containing brooded young 

(Tryonia). C-D. Left side of glandular oviduct and associated structures (Pyrgulopsis). 

Bp, brood pouch; Bu, bursa copulatrix; Cm, columellar muscle; Dn, dissecting needle; 

Glo, glandular oviduct; Rm, rectum; Sr, seminal receptacle.

Methods
Technical Note 449 7

Figure 3. Photographs of male snails. A. Relaxed specimen with extended penis 

(Pyrgulopsis). B-C. Specimens with portion or all of pallial roof removed to show 

attachment of penis to “neck.” (Pyrgulopsis, Taylorconcha, respectively). D. Penis 

with internal “tubular” gland (Amnicola). Gs, glandular strip, Ie, inner edge of penis; 

Oe, outer edge of penis; Pd, penial duct; Pn, penis; Tg, tubular gland; Tgd, tubular 

gland duct.

Methods
Spurwinkia salsa. Photo by Karolyn Darrow.
Technical Note 449 9

Key to the Freshwater 

Truncatelloidean Genera of the 

Western United States

1. Female oviduct glands well developed (oviparous reproductive mode) 

(Figure 2A), shell globose to turriform ________________________2

Female oviduct glands reduced, serving as brood pouch 

(ovoviviparous) (Figure 2B), shell conical to turriform ___________11

2. Penial glands absent (Figure 3B-C)____________________________3

Penial glands present (Figure 3A, D) __________________________7

3. Terminal papilla of (distal end of) penis large (Figure 4A); 

estuarine _______________________________________ Spurwinkia

Terminal papilla of (distal end of) penis very small or absent 

(Figure 4B-G); inland aquatic _______________________________4

Key to the Freshwater Truncatelloidean Genera of the Western United States

Tp

Pd

Pd

A B

C D

E

F G

Pd

Pd Pd

Pd

Pd Figure 4. 

Drawings of penes 

(dorsal aspect). 

A. Spurwinkia salsa. 

B. Fluminicola fuscus. 

C. Fluminicola virens. 

D. Taylorconcha 

serpenticola. 

E. Pyrgulopsis lockensis. 

F. Pyrgulopsis millenaria. 

G. Pristinicola hemphilli. 

Distal edge to left. 

Pd, penial duct; 

Tp, terminal papilla.
10 Technical Note 449

4. Bursa copulatrix absent (Figure 5A) ________________ Taylorconcha

Bursa copulatrix present (Figure 5B-D) ________________________5

Key to the Freshwater Truncatelloidean Genera of the Western United States

Glo Glo

Glo

Glo

Bu

Bu

Bu

Sr

Pw A B

C D

Figure 5.

Drawings of (female) 

glandular oviducts 

and associated 

structures (left side). 

A. Taylorconcha 

serpenticola. 

B. Fluminicola 

ahjumawi. 

C. Pyrgulopsis

militaris. 

D. Pristinicola

hemphilli. 

Anterior to left. 

Bu, bursa copulatrix; 

Glo, glandular oviduct; 

Pw, posterior wall of 

pallial cavity; 

Sr, seminal receptacle.

5. Bursa copulatrix overlapping right side of albumen gland

(Figure 5B); protoconch sculptured with spiral lines 

(Figure 6A) ____________________________________ Fluminicola

Bursa copulatrix overlapping left side of albumen gland 

(Figure 5C-D); protoconch surface wrinkled or pitted

(Figure 6B) ______________________________________________6

Figure 6. 

Scanning electron 

micrographs 

showing sculpture 

on shell apical 

whorls. 

A. Fluminicola 

gustafsoni. 

B. Pyrgulopsis 

castaicensis.
Technical Note 449 11

6. Shell near planispiral to turriform, rarely pupiform;

whorls usually convex; operculum amber-colored

(Figure 7A); penis rectangular to elongate

(<4x longer than wide); penial duct narrow

(Figure 4E-F)____________________________ Pyrgulopsis (in part)

Shell pupiform; whorls near flat; operculum reddish

(Figure 7B); penis vermiform (>6x longer than wide);

penial duct wide (Figure 4G) ______________________ Pristinicola

Key to the Freshwater Truncatelloidean Genera of the Western United States

Figure 7. 

Photographs 

of opercula 

(dorsal aspect). 

A. Pyrgulopsis 

orbiculata. 

B. Pristinicola 

hemphilli.

7. Penial surficial glands (glandular strips or pads)

present (Figure 8A-C) _____________________ Pyrgulopsis (in part)

Penial surficial glands absent ________________________________8

Tg

Gs

Gs

Gp

Pl

Pl

Pa

A B

C D

E F

Tg

Figure 8. 

Drawings of penes 

(A-B, D-F, dorsal side; 

C, ventral side). 

A. Pyrgulopsis militaris. 

B. Pyrgulopsis perforata. 

C. Pyrgulopsis fresti. 

D. Littoridinops monroensis. 

E. Amnicola limosa. 

F. Colligyrus convexus. 

Distal edge to left. 

Gp, glandular pad; 

Gs, glandular strip; 

Pa, glandular papilla; 

Pl, penial lobe; 

Tg, tubular gland.
12 Technical Note 449

8. Penis has glandular papillae (Figure 8D) ____________ Littoridinops

Penis has an internal, tubular gland (Figure 8E-F) ________________9

10. Females have two sperm pouches (bursa copulatrix,

seminal receptacle) (Figure 9A-B)_____________________ Colligyrus

Females have one sperm pouch (bursa copulatrix)

(Figure 9C) ______________________________________ Lyogyrus

9. Shell 3.0-4.5 mm tall; penial lobe medially positioned

(Figure 8E); bursa copulatrix large relative to albumen

gland (Figure 9A) _________________________________ Amnicola

Shell 1.3-3.3 mm tall; penial lobe basally positioned (Figure 8F);

bursa copulatrix small relative to albumen gland

(Figure 9B-C) ___________________________________________10

Key to the Freshwater Truncatelloidean Genera of the Western United States

Figure 9. Female glandular oviducts of amnicolid snails. 

A. Amnicola limosa. B. Colligyrus convexus. 

Anterior to left. C. Lyogyrus sp. 

Bu, bursa copulatrix; Sr, seminal receptacle.

A

Sr

Bu

Sr

Bu

Bu

B

C
Technical Note 449 13Key to the Freshwater Truncatelloidean Genera of the Western United States

Figure 10. 

Drawings

of penes

(dorsal aspect). 

Distal edge

to left. 

A. Potamopyrgus 

antipodarum. 

B. Tryonia

clathrata. 

C. Eremopyrgus 

eganensis. 

D. Ipnobius 

robustus.

11. Males absent in most populations; penial lobes absent

(Figure 10A); white smear often present on inner

side of operculum (Figure 11A, C) ________________ Potamopyrgus

Males common (except in one species of Tryonia), penial lobes 

present (Figure 10B-D); white smear absent on inner side of 

operculum (Figure 11B) ___________________________________12

A B

C D

Pa

Pl

Pa

Figure 11.

Photographs of shells

and opercula.

A. Potamopyrgus antipodarum. 

B. Tryonia imitator. 

C. Operculum (inner side), 

Potamopyrgus antipodarum. 

Ws, white smear.
14 Technical Note 449

12. Penis has square-shaped lobe(s)

(Figure 10C) __________________________________ Eremopyrgus

Penis has papillae (Figure 10B, D) ___________________________13

Key to the Freshwater Truncatelloidean Genera of the Western United States

13. Penis has multiple papillae, including at least one positioned

on inner edge (Figure 10B) ____________________________ Tryonia

Penis has single papilla, positioned on outer edge

(Figure 10D) ______________________________________ Ipnobius
Technical Note 449 15

Species Checklist

Superfamily Truncatelloidea Gray, 1840

 Family Amnicolidae Tryon, 1863

Genus Amnicola Gould and Haldeman in Haldeman, 1840

Amnicola Gould and Haldeman in Haldeman, 1840: 3 [type species, Paludina 

porata Say, 1821 (= Paludina limosa Say, 1817); subsequent designation by 

Herrmannsen 1846: 38].

Remarks: Amnicola (containing six species) lives in a variety of lotic and lentic habitats 

and is distributed in a large portion of eastern North America (north of 

Mexico). One congener (A. limosa) is found in the western United States.

Amnicola limosa (Say, 1817)

Paludina limosa Say, 1817: 125-126. Delaware and Schuylkill Rivers. Types have not 

been found.

Paludina porata Say, 1821: 174. Cayuga Lake [New York]. Lectotype, ANSP 283866; 

paralectotypes, ANSP 368398.

Amnicola ferruginea Calkins 1880: 6, unlabeled figure (on p. 6). Calumet River, Ill. 

[Illinois]. Type has not been found.

Figure 12. Amnicola limosa. A. McWenneger Slough, Flathead County, Montana (University of 

Minnesota Bell Museum of Natural History [BellMNH] 21425). B. Utah Lake, Utah County, Utah 

(USNM 47865). C. Curlew Lake, Ferry County, Washington (USNM 1256491). Scale bars, 1.0 mm.

Species Checklist

AMNICOLIDAE
16 Technical Note 449 AMNICOLIDAEAmnicola orbiculata Lea, 1844: 34. Springfield, Ohio, and Schuylkill? [sic] near 

Philadelphia. Holotype, USNM 121342.

Common name: Mud amnicola.

Distribution: Widely ranging in eastern North America (Burch and Tottenham 1980; 

Clarke 1981); also found in several lakes in Montana and Washington

(Liu et al. 2016). The historic population in Utah Lake appears to be extinct 

(Taylor and Bright 1987).

Taxonomy: Taylor and Bright (1987: fig. 8), Liu et al. (2016).

Remarks: The synonymy does not include the many (extralimital) subspecies of 

Amnicola limosa.

Genus Colligyrus Hershler, 1999

Figure 13. Colligyrus. A-B. C. greggi, springs along Cliff Creek, Sublette County, Wyoming 

(USNM 905410, USNM 883531, respectively). C. C. depressus, springs along Cricket Creek, 

Harney County, Oregon (USNM 860756 [paratype]). D. C. convexus, Lava Creek, Shasta County, 

California (USNM 1004536 [paratype]). Scale bars, 1.0 mm.

Colligyrus Hershler, 1999: 306-308 [type species: Taylorconcha serpenticola Hershler, 

Frest, Johannes, Bowler and Thompson, 1994; original designation.]

Species Checklist
Technical Note 449 17Species Checklist

Remarks: Colligyrus is widely distributed in the northwestern United States and 

AMNICOLIDAE

typically lives in springs and small streams. Molecular evidence suggests that 

there are undescribed congeners in the Klamath River (Hershler et al. 2003a) 

and Puget Sound (Liu et al. 2015a) basins.

Colligyrus convexus Hershler, Frest, Liu, and Johannes, 2003

Colligyrus convexus Hershler, Frest, Liu, and Johannes, 2003a: 278-280, 282, figs. 2-3, 

4a, 5. Lava Creek at and west of Island Road bridge near mouth to Eastman 

Lake, Lava Creek Ranch, north side of The Island, about 8.3 km north of 

Glenburn, Shasta County, California. Holotype, USNM 1004535; paratypes, 

USNM 1004536.

Common name: Canary duskysnail.

Distribution: Fall River, Hat Creek, and lower Burney Creek drainages (Pit River 

basin), California (Hershler et al. 2003a).

Colligyrus depressus Hershler, 1999

Colligyrus depressus Hershler, 1999: 311, 312, 314, 316, figs. 2E-F, 3B, 6. Unnamed 

springs, Cricket Creek, Silvies River drainage, Harney County, Oregon. 

Holotype, USNM 883876; paratypes, USNM 860756.

Common name: Harney Basin duskysnail.

Distribution: Known only from the type locality.

Colligyrus greggi (Pilsbry, 1935)

Hydrobia greggi Pilsbry, 1935: 93-94, fig. 2. Cliff Creek Canyon, a fork of Hoback 

Canyon, about 29 miles [46.7 km] south of Jackson, Wyoming, in the Snake 

River drainage. Lectotype, ANSP 163812; paralectotypes, ANSP 375735.

Common name: Rocky Mountain duskysnail.

Distribution: Upper Snake River basin (Idaho, Wyoming), northeastern Great Basin 

(Idaho, Utah), lower Columbia River basin (Oregon, Washington) (Hershler 

1999; Liu et al. 2015a).

Taxonomy: Hershler (1999), Liu et al. (2015a).
18 Technical Note 449

Genus Lyogyrus Gill, 1863

Lyogyrus Gill, 1863: 34 [type species: Valvata pupoidea Gould, 1839;

original designation].

Remarks: Lyogyrus (containing nine species) is widely distributed in a variety of 

freshwater and brackish water habitats in North America from the Mississippi 

River basin eastward. Additionally, one congener of uncertain species status 

is found in several lakes in the northwestern United States. Lyogyrus was 

frequently treated in the 20th century literature as a subgenus of Amnicola, 

but is currently recognized as a distinct genus based on morphological and 

molecular evidence (Thompson and Hershler 1991; Liu et al. 2015a).

Lyogyrus sp.

Lyogyrus sp.—Liu et al. 2016: 467-469, figs. 6-8.

Common name: Masked duskysnail.

Distribution: Several lakes in northern Washington and Montana (Liu et al. 2015a).

Remarks: The taxonomic status of this snail is uncertain pending further study of the 

poorly known eastern species of Lyogyrus.

Figure 14. Lyogyrus sp. A. Browns Lake, Powell County, Montana (USNM 

1263003). B. Fish Lake, Chelan County, Washington (USNM 1258915). 

C. Upsata Lake, Powell County, Montana (USNM 1262996). Scale bar, 1.0 mm.

AMNICOLIDAE

Species Checklist
Technical Note 449 19

 Family Cochliopidae Tryon, 1866

Genus Eremopyrgus Hershler, 1999

Figure 15. Eremopyrgus eganensis. A-B. Spring northwest of Clark Spring, 

Steptoe Valley, White Pine County, Nevada (USNM 883529, USNM 860759 

[paratype], respectively). Scale bars, 1.0 mm.

Eremopyrgus Hershler, 1999: 326-328 [type species: Eremopyrgus eganensis Hershler, 

1999; original designation].

Remarks: Eremopyrgus is composed of two locally endemic, spring-dwelling species in 

Nevada and northeastern Mexico (Hershler et al. 2002).

Eremopyrgus eganensis Hershler, 1999

Eremopyrgus eganensis Hershler, 1999: 328, figs. 3C, 12, 13A-C. Spring northwest of 

Clark Spring, Steptoe Valley, White Pine County, Nevada. Holotype, 

USNM 874692; paratypes, USNM 860759.

Common name: Steptoe hydrobe.

Distribution: Several springs in Steptoe Valley, Nevada.

Species Checklist

COCHLIOPIDAE
20 Technical Note 449

Genus Ipnobius Hershler, 2001

Ipnobius Hershler, 2001: 19 [type species, Tryonia robusta Hershler, 1989; 

original designation].

Remarks: Ipnobius is a monotypic genus that is distributed in thermal springs in 

Death Valley, California.

Ipnobius robustus Hershler, 1989

Tryonia robusta Hershler, 1989: 208-211, figs. 51d, 55-58. Nevares Springs, Death 

Valley, Inyo County, California. Holotype, USNM 860411; paratypes, USNM 

853557.

Common name: Robust tryonia.

Distribution: Two spring complexes in central Death Valley, California.

Figure 16. Ipnobius robustus. A-B. Nevares Springs, Death 

Valley, Inyo County, California (USNM 883311, USNM 853557 

[paratype], respectively). Scale bars, 1.0 mm.

COCHLIOPIDAE

Species Checklist
Technical Note 449 21

Genus Littoridinops Pilsbry, 1952

Figure 17. Littoridinops monroensis. A. Sulphur Springs, Hillsborough 

County, Florida (USNM 874865). B. Suisun Bay (San Francisco 

Estuary), Contra Costa County, California (USNM 1087202). 

Scale bars, 1.0 mm.

Littoridinops Pilsbry, 1952: 51 [type species: Amnicola tenuipes Couper in Haldeman, 

1844; original designation].

Remarks: Littoridinops (containing four species) is widely distributed along the 

North American Atlantic Coast from New Jersey to Campeche, Mexico; also 

found on several islands in the West Indies (Hershler and Thompson 1992; 

Vázquez et al. 2012). Members of this genus live in estuarine coastal waters and 

occasionally in inland freshwater habitats.

Littoridinops monroensis (Frauenfeld, 1863)

Hydrobia monroensis Frauenfeld, 1863: 1023. Lakeman Rok, Florida [possibly referring 

to Lake Monroe, Volusia County]. Lectotype, NMW; paralectotypes, NMW 

(per Thompson 1968).

Common name: Cockscomb hydrobe.

Distribution: Florida, Bahama Islands, and Cuba (Thompson 1968; Vázquez et al. 

2012). Introduced to San Francisco Bay (Hershler et al. 2007a).

Taxonomy: Hershler et al. (2007a).

Species Checklist

COCHLIOPIDAE
22 Technical Note 449 COCHLIOPIDAE Species Checklist

Genus Spurwinkia Davis, Mazurkiewicz, and Mandracchia, 1982

Spurwinkia Davis, Mazurkiewicz, and Mandracchia, 1982: 162ff [type species: 

Hydrobia salsa Pilsbry, 1905; original designation].

Remarks: Spurwinkia is a monotypic genus that is distributed in estuarine habitats 

along the North American Atlantic Coast.

Spurwinkia salsa (Pilsbry, 1905)

Paludestrina salsa Pilsbry, 1905: 90-91, pl. 3: fig. 10. Cohasset, Massachusetts. 

Lectotype, ANSP 90445; paralectotypes, ANSP 461848.

Common name: Saltmarsh hydrobe.

Distribution: Widely ranging along the western Atlantic coast from New Brunswick to 

Georgia (Hershler and Thompson 1992; McAlpine et al. 2005). Introduced to

San Francisco Bay (Hershler et al. 2015a).

Taxonomy: Hershler et al. (2015a)

Figure 18. Spurwinkia salsa. A. Near Harvard Bridge, Boston, Massachusetts 

(USNM 435954). B. Eden Landing (San Francisco Estuary), Alameda County, 

California (USNM 1120373). Scale bars, 1.0 mm.
Technical Note 449 23Species Checklist COCHLIOPIDAE

Genus Tryonia Stimpson, 1865

Tryonia Stimpson, 1865: 54 [type species: Tryonia clathrata Stimpson, 1865;

original designation].

Calipyrgula Pilsbry, 1934: 15 [type species: Calipyrgula carinifer Pilsbry, 1934;

original designation].

Figure 19. Tryonia. A. T. angulata, Big Spring, Ash Meadows, Nye County, Nevada (USNM 

883304). B. T. clathrata, Cardy Lamb Spring, Moapa Valley, Clark County, Nevada (USNM 

874355). C. T. imitator, Morro Estuary, San Luis Obispo County, California (USNM 892057). 

D. T. ericae, North Scruggs Spring, Ash Meadows, Nye County, Nevada (USNM 859163 

[paratype]). E. T. porrecta, southern Suisun Bay (San Francisco Estuary), Contra Costa County, 

California (USNM 1087204). F. T. monitorae, Hot Springs, Potts Ranch, Monitor Valley, Nye 

County, Nevada (USNM 860760 [paratype]). Scale bars, 1.0 mm.
24 Technical Note 449 COCHLIOPIDAE Species Checklist

Durangonella Morrison, 1945: 18 [type species: Hydrobia seemani Frauenfeld, 1863; 

original designation].

Hyalopyrgus Thompson, 1968: 43-46 [type species: Bythinella aequicostata Pilsbry, 

1890; original designation].

Paupertryonia Taylor, 1987: 38 [type species: Potamopyrgus cheatumi Pilsbry, 1935; 

original designation].

Remarks: Tryonia is a relatively large genus (33 species) that is widely distributed in 

southwestern North America and is also found in the Florida Peninsula. 

Fourteen congeners are distributed in drainages of the western United States. 

Tryonia usually inhabits thermal springs; a few species live in lakes and/or 

coastal waters. Molecular evidence suggests that Tryonia is paraphyletic 

relative to the monotypic northern Mexican genus Minckleyella; these taxa are 

currently recognized as distinct because of the differences in their female 

reproductive anatomy (Hershler et al. 2011).

Tryonia angulata Hershler and Sada, 1987

Tryonia angulata Hershler and Sada, 1987: 810-811, 814-817, figs. 39a, 40-41, 42a,d, 

43. Fairbanks Spring, Ash Meadows, Nye County, Nevada. Holotype, USNM 

859151; paratypes, UF 93960, USNM 859152.

Common name: Sportinggoods tryonia.

Distribution: Three large spring pools in Ash Meadows, Nevada.

Taxonomy: Hershler (2001).

Tryonia clathrata Stimpson, 1865

Tryonia clathrata Stimpson, 1865: 54, pl. 8, fig. 1. Basin of the Colorado Desert 

[in error, probably Moapa Valley, Nevada; Hershler 1999]. Lectotype (and 

paralectotypes), ANSP 27969.

Common name: Grated tryonia.

Distribution: White River drainage, Nevada (Hershler 2001).

Taxonomy: Hershler (2001).

Tryonia elata Hershler and Sada, 1987

Tryonia elata Hershler and Sada, 1987: 831, figs. 39b, 42c, 53e-h, 56. Point of Rocks 

Springs, Ash Meadows, Nye County, Nevada. Holotype, USNM 859159; 

paratypes, UF 93963, USNM 859160.

Common name: Point of Rocks tryonia.

Distribution: Two small springs at Point of Rocks, Ash Meadows, Nevada.

Taxonomy: Hershler (2001).
Technical Note 449 25Species Checklist COCHLIOPIDAE

Tryonia ericae Hershler and Sada, 1987

Tryonia ericae Hershler and Sada, 1987: 826-831, figs. 39c-d, 42f,i, 53a-d, 54. North 

Scruggs Spring, Ash Meadows, Nye County, Nevada. Holotype, USNM 859162; 

paratypes, UF 93962, USNM 859163.

Common name: Minute tryonia.

Distribution: Two closely proximal springs in Ash Meadows, Nevada.

Taxonomy: Hershler (2001).

Tryonia gilae Taylor, 1987

Tryonia gilae Taylor, 1987: 36-37, fig. 17. Unnamed spring on north side of Gila River 

about 2 mi [3.2 km] north of Bylas, Graham County, Arizona. Holotype, 

LACM 2187; paratypes, UTEP 10063.

Tryonia gilae Hershler in Hershler and Landye, 1988: 43, 47-49, 58, figs. 14c,f, 39a-e, 

40, 41a-c, 42, 43d-i, 44-45, 46a, 47b. Small spring near Bylas, Graham County, 

Arizona. Holotype, USNM 859059; paratypes, USNM 859060.

Common name: Gila tryonia.

Distribution: Springs along the north side of the Gila River between Bylas and 

Pima, Arizona.

Taxonomy: Hershler (2001).

Remarks: Tryonia gilae Hershler is both a junior primary homonym and junior 

subjective synonym of Tryonia gilae Taylor.

Tryonia imitator (Pilsbry, 1899)

Paludestrina imitator Pilsbry, 1899: 124. Santa Cruz, California. Lectotype, ANSP 

6270; paralectotypes, ANSP 457182.

Paludestrina curta Arnold, 1903: 305, pl. VIII: fig. 2. San Pedro bluffs [California], 

Upper San Pedro Series [Pleistocene]. Holotype, USNM 162542.

Common name: Mimic tryonia.

Distribution: Widely ranging along the central and southern California coast 

(Taylor 1981).

Taxonomy: Hershler (2001).
26 Technical Note 449 COCHLIOPIDAE Species Checklist

Tryonia infernalis Hershler, Liu, and Simpson, 2015

Tryonia infernalis Hershler, Liu, and Simpson, 2015b: 114-116, figs. 4-5. Blue Point 

Spring, Clark County, Nevada. Holotype, USNM 883884; paratypes, 

USNM 1266143.

Common name: Blue Point tryonia [name newly introduced herein].

Distribution: Found only at the type locality.

Tryonia margae Hershler, 1989

Tryonia margae Hershler, 1989: 202-205, figs. 48-50, 51a. (Upper) warm spring on 

limestone bench, Grapevine Springs, Death Valley, California. Holotype, 

USNM 860408; paratypes, USNM 857952.

Common name: Grapevine Springs elongate tryonia.

Distribution: Found only at the type locality and a second, closely proximal spring.

Taxonomy: Hershler (2001).

Tryonia monitorae Hershler, 1999

Tryonia monitorae Hershler, 1999: 332, 334, figs. 3D, 13D-G, 15. Hot Springs, Potts 

Ranch, Monitor Valley, Nye County, Nevada. Holotype, USNM 892046; 

paratypes, USNM 860760.

Common name: Monitor tryonia.

Distribution: Known only from the type locality.

Taxonomy: Hershler (2001).

Tryonia porrecta (Mighels, 1848)

Paludina porrecta Mighels, 1848: 22. Oahu [Hawaii]. Type has not been found 

[however, NHMUK 1995123 may be a syntype lot; Hershler 2001].

Amnicola protea Gould, 1855: 129-130. Colorado Desert (Gran Jornado). 

Lectotype, USNM 121074; paralectotypes, USNM 27906, USNM 860867.

Melania exigua Conrad, 1855: 269. Colorado Desert, California. Lectotype (and 

paralectotypes), ANSP 27965.

Pyrgulopsis imminens Taylor, 1950: 30-31, figs. 4-6. Shore of Salton Sea by Fish Springs 

(= Desert Shores), Imperial County, California. Holotype, SBMNH 35497; 

paratypes, SBMNH 35498, SBMNH 35499, USNM 613967.

Pyrgulopsis blakeana Taylor, 1950: 31-32, fig. 7. Holotype, SBMNH 35500; 

paratypes, SBMNH 35501, SBMNH 35502, USNM 613966.

Pyrgulopsis cahuillarum Taylor, 1950: 31-32, fig. 7. Holotype, SBMNH 35503.
Technical Note 449 27Species Checklist

Common name: Desert tryonia.

COCHLIOPIDAE

Distribution: Lower Colorado River basin (Arizona, California, Mexico), Great Basin 

(Nevada, Utah), San Francisco Bay (California), Hawaii. The populations 

in San Francisco Bay (Hershler et al. 2007a) and artificial lakes in Phoenix 

(Hershler et al. 2015a) may have been introduced. (The Hawaiian population 

could be native or prehistorically introduced; Cowie 1997.)

Taxonomy: Hershler (2001), Hershler et al. (2007a).

Remarks: Most populations of T. porrecta are parthenogenetic and entirely composed 

of females (Hershler et al. 1999a; Hershler et al. 2005); one of the San Francisco 

Bay populations contains males (Hershler et al. 2007a).

Tryonia quitobaquitae Hershler in Hershler and Landye, 1988

Tryonia quitobaquitae Hershler in Hershler and Landye, 1988: 50, figs. 39f-h, 41df, 43a-c, 46a, 47a. Quitobaquito Springs, Organ Pipe [Cactus] National 

Monument, Pima County, Arizona. Holotype, USNM 859061; paratypes, 

USNM 859062.

Common name: Quitobaquito springsnail.

Distribution: Two closely proximal springs in the Rio Sonoyta basin, southern 

Arizona.

Taxonomy: Hershler (2001).

Tryonia rowlandsi Hershler, 1989

Tryonia rowlandsi Hershler, 1989: 211-215, figs. 51b, 59-60. Grapevine Springs, 

Grapevine Mountains, Death Valley, Inyo County, California. Holotype, 

USNM 860409; paratypes, USNM 857953.

Common name: Grapevine Springs squat tryonia.

Distribution: Found only at the type locality.

Taxonomy: Hershler (2001).

Tryonia salina Hershler, 1989

Tryonia salina Hershler, 1989: 215-216, figs. 51e, 61-62. Spring brook in Cottonball 

Marsh, Death Valley, Inyo County, California. Holotype, USNM 860410; 

paratypes, USNM 853556.

Common name: Cottonball Marsh tryonia.

Distribution: Several springs in Cottonball Marsh, Death Valley (California).

Taxonomy: Hershler (2001).
28 Technical Note 449 COCHLIOPIDAE Species Checklist

Tryonia variegata Hershler and Sada, 1987

Tryonia variegata Hershler and Sada, 1987: 817, 819, 822, 824-826, figs. 39e-g, 

42b,e,g-h, 45-52. Five Springs, Ash Meadows, Nye County, Nevada. Holotype, 

USNM 859166; paratypes, UF 93961, USNM 859167.

Common name: Amargosa tryonia.

Distribution: Amargosa River drainage, from Ash Meadows south to Saratoga Spring 

(Hershler 1989).

Taxonomy: Hershler (2001).

Remarks: Genetic evidence suggests that T. variegata is composed of two 

evolutionarily distinct lineages and requires taxonomic revision 

(Hershler et al. 1999a).
Technical Note 449 29

 Family Hydrobiidae Stimpson, 1865

 Subfamily Nymphophilinae Taylor, 1966

Genus Pyrgulopsis Call and Pilsbry, 1886

Figure 20. Pyrgulopsis. A. P. californiensis, Grapevine Spring, Anza-Borrego Desert State Park, 

San Diego County, SBMNH 4349. B. P. nevadensis, Pyramid Lake, Washoe County, Nevada 

(USNM 590364). C. P. dixensis, spring in Dixie Valley, Pershing County, Nevada 

(USNM 860688 [paratype]). D. P. robusta, Snake River, north of Jackass Butte, Elmore County, 

Idaho (ALBRCIDA 7568). E. P. aardahli, spring at Bramlette Ranch, Benton Valley, Mono County, 

California (USNM 857951 [paratype]). F. P. merriami, Ash Spring, Pahranagat Valley, Lincoln 

County, Nevada (USNM 873395). Scale bars, 1.0 mm.

Species Checklist

HYDROBIIDAE
30 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis Call and Pilsbry, 1886: 9-10 [type species: Pyrgula nevadensis Stearns, 

1883; original designation].

Fontelicella Gregg and Taylor, 1965: 103-104 [type species: Fontelicella californiensis

Gregg and Taylor, 1965; original designation].

Natricola Gregg and Taylor, 1965: 108-109 [type species: Pomatiopsis robusta Walker, 

1908; original designation].

Microamnicola Gregg and Taylor, 1965: 109 [type species: Amnicola micrococcus

Pilsbry in Stearns, 1893; original designation].

Savaginius Taylor, 1966b: 130 [type species: Paludestrina nanna Chamberlain and 

Berry, 1933; original designation].

Nymphophilus Taylor, 1966a: 199-203 [type species: Nymphophilus minckleyi Taylor, 

1966a; original designation].

Mexistiobia Hershler, 1985: 46-47 [type species: Mexistiobia manantiali Hershler, 1985; 

original designation].

Apachecoccus Taylor, 1987: 32 [type species: Apachecoccus arizonae Taylor, 1987; 

original designation].

Yaquicoccus Taylor, 1987: 34 [type species: Yaquicoccus bernadinus Taylor, 1987; 

original designation].

Remarks: Pyrgulopsis contains 143 currently recognized species; 126 are distributed 

in the western United States while the others are found in the Missouri River 

headwaters and in northern Mexico. Pyrgulopsis typically lives in springs and 

spring-fed streams, although a few species live in lakes and/or rivers. The 

systematics of Pyrgulopsis have been unstable in recent decades (see Hershler 

and Thompson 1987; Hershler 1994; Thompson and Hershler 2002; Liu and 

Hershler 2005), and genetic evidence suggests that the genus, as currently 

constituted, may be paraphyletic relative to Floridobia, which is distributed 

in Atlantic Coastal drainages (Hershler et al. 2003b; Liu and Hershler 2005). 

These taxa continue to be recognized as distinct (pending further phylogenetic 

study) because of their broad geographic separation and differences in their 

female reproductive anatomy (Liu and Hershler 2005).

Pyrgulopsis aardahli Hershler, 1989

Pyrgulopsis aardahli Hershler, 1989: 179-181, figs. 10-13. Spring at Bramlette Ranch, 

Benton Valley, Mono County, California. Holotype, USNM 860406; paratypes, 

USNM 857951.

Common name: Benton Valley springsnail.

Distribution: Known only from the type locality.

Taxonomy: Hershler (1994).
Technical Note 449 31Species Checklist HYDROBIIDAE

Pyrgulopsis aloba Hershler, 1998

Pyrgulopsis aloba Hershler, 1998: 60-62, figs. 7F, 19D-E, 33A-B. Spring, northwest of 

Duckwater, Duckwater Valley, Nye County, Nevada. Holotype, USNM 883847; 

paratypes, USNM 860681.

Common name: Duckwater pyrg.

Distribution: Three springs in Duckwater Valley, Nevada.

Pyrgulopsis amargosae Hershler, 1989

Pyrgulopsis amargosae Hershler, 1989: 181-182, figs. 15-16, 17a, 18. Saratoga Spring, 

southern Death Valley, San Bernardino County, California. Holotype, 

USNM 860401; paratypes, USNM 853515.

Common name: Amargosa springsnail.

Distribution: Lower portion of Amargosa River drainage (Tecopa area, southern 

Death Valley), California.

Taxonomy: Hershler (1994).

Remarks: Genetic evidence suggests that P. amargosae is composed of at least two 

evolutionarily distinct lineages and requires taxonomic revision (Hershler and 

Liu 2008; Hershler et al. 2013).

Pyrgulopsis anatina Hershler, 1998

Pyrgulopsis anatina Hershler, 1998: 63-64, figs. 7H, 19H-I, 33F-H. Spring southeast of 

Old Collins Spring, Duckwater Valley, Nye County, Nevada. Holotype, 

USNM 883848; paratypes, USNM 860710.

Common name: Southern Duckwater pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis anguina Hershler, 1998

Pyrgulopsis anguina Hershler, 1998: 110-111, figs. 9K, 23H-J, 44A-E. Big Springs, 

Snake Valley, White Pine County, Nevada. Holotype, USNM 874678; 

paratypes, USNM 860725.

Common name: Longitudinal gland pyrg.

Distribution: Snake Valley (Bonneville basin), Nevada-Utah.
32 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis archimedis Berry, 1947

Pyrgulopsis archimedis Berry, 1947: 76-77, pl. 7, fig. 6. Upper Klamath Lake near 

Algoma, Klamath County, Oregon. Holotype, USNM 739417; paratypes, 

USNM 613965.

Common name: Archimedes pyrg.

Distribution: Upper Klamath Lake and the uppermost reach of its outflow (Link 

River), Oregon; Hat Creek and Fall River drainages, lower Pit River basin, 

California (Hershler 1994; Hershler et al. 2003a).

Taxonomy: Hershler (1994), Hershler et al. (2003a).

Pyrgulopsis arizonae (Taylor, 1987)

Apachecoccus arizonae Taylor, 1987: 32, fig. 15. Unnamed spring on north side of Gila 

River about 2 mi [3.2 km] north of Bylas, Graham County, Arizona. Holotype, 

LACM 2203; paratypes, ANSP 376020, UF 160939, USNM 854090, UTEP 

10050.

Pyrgulopsis sancarlosensis Hershler in Hershler and Landye, 1988: 35, 39-40, 

figs. 13e, 26e, 31a-g, 32-33. Springs west of Tom Niece Springs, Graham 

County, Arizona. Holotype, USNM 859051; paratypes, USNM 859052.

Common name: Apache springsnail.

Distribution: Springs along north side of Gila River between Bylas and Pima, 

southern Arizona.

Taxonomy: Hershler (1994).

Pyrgulopsis augustae Hershler, 1998

Pyrgulopsis augustae Hershler, 1998: 89, figs. 9B, 22A, 40A-B. Cain Spring, Antelope 

Valley, Lander County, Nevada. Holotype, USNM 874402; paratypes, 

USNM 860687.

Common name: Elongate Cain Spring pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis aurata Hershler, 1998

Pyrgulopsis aurata Hershler, 1998: 74-75, figs. 8E, 20K-L, 36E-G. Coyote Spring, 

Pershing County, Nevada. Holotype, USNM 874393; paratypes, USNM 

860696.

Common name: Pleasant Valley pyrg.

Distribution: Known only from the type locality.
Technical Note 449 33Species Checklist HYDROBIIDAE

Pyrgulopsis avernalis (Pilsbry, 1935)

Fluminicola avernalis Pilsbry, 1935: 92-93, fig. 1. Colorado Desert [probably in error 

as there are no other reliable records for this species from the Colorado Desert; 

Hershler 1994]. Lectotype, ANSP 27784; paralectotypes, ANSP 375737 

(mixed with Pyrgulopsis carinifera).

Common name: Moapa pebblesnail.

Distribution: Springs in Moapa Valley (lower Colorado River basin), Nevada 

(Hershler 1998).

Taxonomy: Hershler (1994).

Pyrgulopsis bacchus Hershler in Hershler and Landye, 1988

Pyrgulopsis bacchus Hershler in Hershler and Landye, 1988: 21, figs. 10d, 11e-g, 12df, 14b, 15. Grapevine Spring, Mohave County, Arizona. Holotype, USNM 

859037; paratypes, USNM 859038.

Common name: Grand Wash springsnail.

Distribution: Two closely proximal springs in Grand Wash, lower Colorado River 

basin, Arizona.

Taxonomy: Hershler (1994).

Pyrgulopsis basiglans Hershler, 1998

Pyrgulopsis basiglans Hershler, 1998: 91-93, figs. 9D, 13D, 22D, 40F-H. Spring, 

Cooks Creek, Carico Lake Basin, Lander County, Nevada. Holotype, 

USNM 874280; paratypes, USNM 860692.

Common name: Large gland Carico pyrg.

Distribution: Two springs in the Carico Lake basin, Nevada.

Pyrgulopsis bernardina (Taylor, 1987)

Yaquicoccus bernardinus Taylor, 1987: 34-35, fig. 16. Spring, 2,300 ft E, 4,600 ft S of 

NW corner, sec. 15, T24S, R30E, Cochise County, Arizona. Holotype, LACM 

2186; paratypes, ANSP 376019, UF 160934, USNM 854078.

Pyrgulopsis cochisi Hershler in Hershler and Landye, 1988: 41, figs. 25d, 30h-k, 33-34. 

Spring at San Bernardino Ranch, Cochise County, Arizona. Holotype, 859055; 

paratypes, USNM 859056.

Common name: San Bernardino springsnail.

Distribution: Several springs in the upper San Bernardino basin, southern Arizona and 

northern Sonora (Mexico) (Varela-Romero et al. 2013).

Taxonomy: Hershler (1994).
34 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis bifurcata Hershler, 1998

Pyrgulopsis bifurcata Hershler, 1998: 93-94, figs. 9E, 22E, 41A-C. Springs west of 

Carico Lake, Carico Lake Basin, Lander County, Nevada. Holotype, 

USNM 874306; paratypes, USNM 860693.

Common name: Small gland Carico pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis breviloba Hershler, 1998

Pyrgulopsis breviloba Hershler, 1998: 39, 41, figs. 6G, 11C, 14D-F, 17L-M, 28D-F. 

Flag Springs (the middle of three), White River Valley, Nye County, Nevada. 

Holotype, USNM 873174; paratypes, USNM 860689.

Common name: Flag pyrg.

Distribution: Flag Springs complex (White River Valley), and one spring in Dry Lake 

Valley, Nevada.

Pyrgulopsis bruesi Hershler and Sada, 2000

Pyrgulopsis bruesi Hershler and Sada, 2000: 367-368, 370-372, figs. 1-4. Small stream 

that enters Fly Reservoir, about 23 airline-km north-northeast of Gerlach, 

Washoe County, Nevada. Holotype, USNM 892079; paratypes, USNM 860868.

Common name: Fly Ranch pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis bruneauensis Hershler, 1990

Pyrgulopsis bruneauensis Hershler, 1990: 803, 805, 809, 811, 813-814, figs. 1-6. Spring 

along west side of Bruneau River, ca. 100 m downflow from Hot Creek’s 

confluence with the river, Owyhee County, Idaho. Holotype, USNM 860507; 

paratypes, USNM 860508.

Common name: Bruneau Hot springsnail.

Distribution: Small springs along Hot Creek and confluent Bruneau River (Idaho).

Taxonomy: Hershler (1994).
Technical Note 449 35Species Checklist HYDROBIIDAE

Pyrgulopsis bryantwalkeri Hershler, 1994

Fluminicola nevadensis Walker, 1916: 6-7, unlabeled figure. Spring in the Cortez 

foot-hills, Humboldt Valley, Elko County, Nevada. Lectotype, UMMZ 118012; 

paralectotypes, ANSP 115948, MCZ 31450.

Pyrgulopsis bryantwalkeri Hershler, 1994: 23. Replacement name for Fluminicola 

nevadensis Walker, 1916; preoccupied in Pyrgulopsis by Pyrgula nevadensis

Stearns, 1883.

Common name: Cortez Hills pebblesnail.

Distribution: Known from a single spring in the Humboldt River basin, Nevada 

(Hershler 1994).

Taxonomy: Hershler (1994).

Pyrgulopsis californiensis (Gregg and Taylor, 1965)

Fontelicella californiensis Gregg and Taylor, 1965: 109. Campo Creek, San Diego 

County, California, 0.6 mi. [1.0 km] east of Mountain Empire Dam. Holotype, 

UMMZ 220000.

Distribution: Southern California and adjacent Baja California. From the southern 

Sierra Nevada (western slope only) through the western Transverse Ranges 

and coastal plains to the Laguna Mountains.

Common name: Laguna Mountain springsnail.

Taxonomy: Hershler (1994).

Remarks: Genetic evidence suggests that P. californiensis is composed of two 

evolutionarily distinct lineages and requires taxonomic revision (Hershler and 

Liu 2010).

Pyrgulopsis carinata Hershler, 1998

Pyrgulopsis carinata Hershler, 1998: 60, figs. 7E, 19C, 32F-G. Little Warm Spring, 

Duckwater Valley, Nye County, Nevada. Holotype, USNM 883975; paratypes, 

USNM 860680.

Common name: Carinate Duckwater pyrg.

Distribution: Known only from the type locality.
36 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis carinifera (Pilsbry, 1935)

Fluminicola avernalis carinifera Pilsbry, 1935: 93, fig. 3. Colorado Desert [in error, 

probably Moapa Valley, Nevada; Hershler 1994]. Lectotype, ANSP 164091; 

paralectotypes, ANSP 375736.

Common name: Moapa Valley pyrg.

Distribution: Moapa Valley, lower Colorado River basin (Hershler 1998).

Taxonomy: Hershler (1994).

Remarks: Elevated to full species by Hershler (1994).

Pyrgulopsis castaicensis Hershler and Liu, 2010

Pyrgulopsis castaicensis Hershler and Liu, 2010: 6-10, figs. 3-5. Middle Canyon Spring, 

ca. 1.46 km southwest of Castaic Junction, Los Angeles County, California. 

Holotype, USNM 1120442; paratypes, USNM 1132532.

Common name: Middle Canyon Spring pyrg.

Distribution: Known only from the type locality in the Santa Clara River basin.

Pyrgulopsis chamberlini Hershler, 1998

Pyrgulopsis chamberlini Hershler, 1998: 122-124, figs. 10G, 25A-C, 47D-G. Spring, 

Glenwood, Sevier River drainage, Sevier County, Utah. Holotype, USNM 

883576; paratypes, USNM 860729.

Common name: Smooth Glenwood pyrg.

Distribution: Two closely proximal springs in the Sevier River drainage (Utah).

Pyrgulopsis cinerana Hershler, Frest, Liu, and Johannes, 2003

Pyrgulopsis cinerana Hershler, Frest, Liu, and Johannes, 2003a: 296-299, figs. 4D, 

18-19. Nearest spring east of Ash Valley Road (Modoc County 527) crossing 

of Ash Creek, north side of creek, JJJ Ranch, Lassen County, California. 

Holotype, USNM 1004543; paratypes, USNM 1004544.

Common name: Ash Valley pyrg.

Distribution: Cold springs associated with upper Ash Creek, upper Pit River basin.
Technical Note 449 37Species Checklist HYDROBIIDAE

Pyrgulopsis coloradensis Hershler, 1998

Pyrgulopsis coloradensis Hershler, 1998: 29, figs. 6C, 17D, 27A-B. Blue Point Spring, 

Colorado River drainage, Clark County, Nevada. Holotype, USNM 854621; 

paratypes, USNM 860677.

Common name: Blue Point pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis conica Hershler in Hershler and Landye, 1988

Pyrgulopsis conicus Hershler in Hershler and Landye, 1988: 21, figs. 10e, 16a-d, 17-

18. Dripping Springs, Mohave County, Arizona. Holotype, USNM 859039; 

paratypes, USNM 859040.

Common name: Kingman springsnail.

Distribution: Three springs in the Black Mountains, lower Colorado River basin, 

Arizona.

Taxonomy: Hershler (1994).

Pyrgulopsis cruciglans Hershler, 1998

Pyrgulopsis cruciglans Hershler, 1998: 72, figs. 8C, 20F-H, 36A-B. Flat Spring, Steptoe 

Valley, White Pine County, Nevada. Holotype, USNM 874285; paratypes, 

USNM 860709.

Common name: Transverse gland pyrg.

Distribution: Several springs in Steptoe Valley and adjacent basins, eastern Nevada.

Pyrgulopsis crystalis Hershler and Sada, 1987

Pyrgulopsis crystalis Hershler and Sada, 1987: 797, 799-802, figs. 8c,f,i, 18c, 23d, 

24b, 28. Crystal Pool, Ash Meadows, Nye County, Nevada. Holotype, USNM 

859205; paratypes, UF 93956, USNM 859206.

Common name: Crystal springsnail.

Distribution: Known only from the type locality.

Taxonomy: Hershler (1994).
38 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis cybele Hershler and Liu, 2012

Pyrgulopsis cybele Hershler and Liu, 2012a: 23, 26-27, figs. 2-4. Unnamed spring brook 

on east side of South Fork Owyhee River, Elko County, Nevada. Holotype, 

USNM 1148155; paratypes, USNM 1157696.

Common name: Nature pyrg.

Distribution: Two closely proximal springs that discharge to the “Devil’s Pinball” reach 

of the South Fork Owyhee River.

Pyrgulopsis deaconi Hershler, 1998

Pyrgulopsis deaconi Hershler, 1998: 23, 25, figs. 6B, 11A, 17B-C, 26D-G. Red Spring, 

Red Rock Canyon Recreation Lands [Red Rock Canyon National Conservation 

Area], Las Vegas Valley, Clark County, Nevada. Holotype, USNM 874454; 

paratypes, USNM 860676.

Common name: Spring Mountains pyrg.

Distribution: Spring Mountains (Las Vegas and Pahrump Valley drainages), Nevada.

Pyrgulopsis deserta (Pilsbry, 1916)

Amnicola deserta Pilsbry, 1916: 111. Washington County, Utah. Lectotype, ANSP 

12112; paralectotypes, ANSP 396958.

Common name: Desert springsnail.

Distribution: Springs in the St. George area (Utah) and below the Virgin River narrows 

near Littlefield (Arizona), lower Colorado River basin (Hershler and Landye 

1998).

Taxonomy: Hershler and Landye (1988), Hershler (1994).

Pyrgulopsis diablensis Hershler, 1995

Pyrgulopsis diablensis Hershler, 1995: 344-345, 347, 349, figs. 1-3, 5A. Unnamed creek, 

Del Puerto Canyon, Del Puerto [Canyon] Road, 20 km west of HW [Interstate] 

5, Stanislaus County, California. Holotype, USNM 860645; paratypes, 

USNM 883791.

Common name: Diablo Range pyrg.

Distribution: Known only from the type locality.
Technical Note 449 39Species Checklist HYDROBIIDAE

Pyrgulopsis dixensis Hershler, 1998

Pyrgulopsis dixensis Hershler, 1998: 73-74, figs. 8D, 13C, 20I-J, 36C-D. Springs westsouthwest of Hot Springs, Dixie Valley, Pershing County, Nevada. Holotype, 

USNM 874391; paratypes, USNM 860688.

Common name: Dixie Valley pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis eremica Hershler, 1995

Pyrgulopsis eremica Hershler, 1995: 349-351, 354, figs. 5B, 7-9. Unnamed springs, 

tributary to Willow Creek, Willow Creek Valley, Lassen County, California. 

Holotype, USNM 860644; paratypes, USNM 858264.

Common name: Smoke Creek pyrg.

Distribution: Portions of the northeastern Great Basin in California.

Pyrgulopsis erythropoma (Pilsbry, 1899)

Fluminicola fusca var. minor Stearns, 1893: 282 [nomen nudum].

Fluminicola erythropoma Pilsbry, 1899: 125-126. Ash Meadows, Nye County, Nevada 

[Probably Kings Pool at Point of Rocks; Hershler and Sada 1987]. 

Lectotype, ANSP 73607; paralectotypes, ANSP 396951.

Common name: Ash Meadows pebblesnail.

Distribution: Point of Rocks spring complex, Ash Meadows.

Taxonomy: Hershler and Sada (1987), Hershler (1994).

Remarks: Stearn’s (1893) publication of Fluminicola fusca var. minor was not 

accompanied by a description and therefore this name is not available per 

International Code of Zoological Nomenclature Article 12.1. His cited

material for this nomen nudum (USNM 123624) was from the type locality of 

P. erythropoma (per Hershler 1994: 35) and closely conforms to this species in

all respects.

Pyrgulopsis fairbanksensis Hershler and Sada, 1987

Pyrgulopsis fairbanksensis Hershler and Sada, 1987: 796-797, figs. 8d,g, 18a-b, 19a, 24d, 

26-27. Fairbanks Spring, Ash Meadows, Nye County, Nevada. Holotype, 

USNM 859203; paratypes, UF 93955, USNM 859204.

Common name: Fairbanks springsnail.

Distribution: Known only from the type locality.

Taxonomy: Hershler (1994).
40 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis falciglans Hershler, Frest, Liu, and Johannes, 2003

Pyrgulopsis falciglans Hershler, Frest, Liu, and Johannes, 2003a: 293-296, figs. 4C, 

15-16. Cold spring about 0.4 km southeast of Smokey Charlie [Charley] Spring 

and 0.32 km west of Modoc County 63, at source next to homestead cabin, 

Modoc County, California. Holotype, USNM 1004605; paratypes, 

USNM 1004606.

Common name: Likely pyrg.

Distribution: Two closely proximal springs along the South Fork Pit River.

Pyrgulopsis fausta Hershler, 1998

Pyrgulopsis fausta Hershler, 1998: 15, 23, figs. 6A, 12A, 17A, 26A-C. Corn Creek 

Springs, Las Vegas Valley, Clark County, Nevada. Holotype, USNM 874757; 

paratypes, USNM 860765.

Common name: Corn Creek pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis fresti Hershler and Liu, 2009

Pyrgulopsis fresti Hershler and Liu, 2009: 9, 11-12, figs. 3-5. Tudor Warm Springs, east 

side of Owyhee River, second spring from north, Malheur County, Oregon. 

Holotype, USNM 1102148; paratypes, USNM 1116914.

Common name: Owyhee Hot springsnail.

Distribution: Springs along a short reach of Owyhee River drainage above Three Forks 

(Oregon).

Pyrgulopsis fusca Hershler, 1998

Pyrgulopsis fusca Hershler, 1998: 122, figs. 10F, 24K-M, 47A-C. Spring brook, Otter 

Creek, ca. 1.6 km above The Narrows, Piute County, Utah. Holotype, USNM 

883439; paratypes, USNM 860728.

Common name: Otter Creek pyrg.

Distribution: Sevier River drainage, Utah.

Pyrgulopsis gibba Hershler, 1995

Pyrgulopsis gibba Hershler, 1995: 354, 357-358, figs. 5C, 10-12. Unnamed springs west 

of Fee Reservoir, Surprise Valley, Modoc County, California. Holotype, 

USNM 860643; paratypes, USNM 858275.

Distribution: Northwestern Great Basin (California, Nevada, Oregon) (Hershler 1998).
Technical Note 449 41Species Checklist HYDROBIIDAE

Pyrgulopsis gilae (Taylor, 1987)

Fontelicella gilae Taylor, 1987: 16, fig. 7. Springs on north side of East Fork of Gila 

River, Grant County, New Mexico. Holotype LACM 2214; paratypes, ANSP 

376025, UF 160936, USNM 854087, UTEP 10054.

Common name: Surprise Valley pyrg.

Distribution: Several groups of springs in lower reach of the East Fork Gila River basin 

(below the mouth of Black Canyon), and a single spring along the Gila River 

ca. 2 km below the East Fork confluence (New Mexico) (Hershler et al. 2014b).

Taxonomy: Hershler (1994), Hershler et al. (2014b).

Pyrgulopsis giulianii Hershler and Pratt, 1990

Pyrgulopsis giulianii Hershler and Pratt, 1990: 279-281, 283-285, figs. 1-3. Sand 

Canyon, Kern County, California. Holotype, USNM 860444; paratypes, 

SBMNH 35140, USNM 853519.

Common name: Southern Sierra Nevada springsnail.

Distribution: Springs along the flanks of the southern Sierra Nevada, California.

Taxonomy: Hershler (1994).

Pyrgulopsis glandulosa Hershler in Hershler and Landye, 1988

Pyrgulopsis glandulosa Hershler in Hershler and Landye, 1988: 8, 11, 15, 17, 

figs. 3a-c, 4-9, 10a-b. Nelson Place Spring, Yavapai County, Arizona. Holotype, 

USNM 859047; paratypes, USNM 859048.

Common name: Verde Rim springsnail.

Distribution: Headwaters (two closely proximal springs) of Sycamore Creek, Verde 

River drainage, central Arizona.

Taxonomy: Hershler (1994).

Pyrgulopsis gracilis Hershler, 1998

Pyrgulopsis gracilis Hershler, 1998: 43, 45, 47, figs. 6I, 11D, 18B-C, 29F-H. Emigrant 

Springs (northernmost in spring complex), White River Valley, Nye County, 

Nevada. Holotype, USNM 873158; paratypes, USNM 860698.

Common name: Emigrant pyrg.

Distribution: Emigrant Springs complex, White River Valley, Nevada.
42 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis greggi Hershler, 1995

Pyrgulopsis greggi Hershler, 1995: 358-359, figs. 5D, 14-16. Grapevine Creek, Fort 

Tejon State Historical Park, Kern County, California. Holotype, USNM 

860641; paratypes, USNM 874319.

Common name: Kern River springsnail.

Distribution: Grapevine Creek drainage, upper Kern River basin (California).

Pyrgulopsis hamlinensis Hershler, 1998

Pyrgulopsis hamlinensis Hershler, 1998: 106-108, figs. 9I, 22K, 43A-C. Springs 

0.5 km east of White Rock Cabin Springs, Hamlin Valley, Beaver County, Utah. 

Holotype, USNM 883215; paratypes, USNM 860695.

Common name: Hamlin Valley pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis hovinghi Hershler, 1998

Pyrgulopsis hovinghi Hershler, 1998: 115, 117, figs. 10B, 11I, 16D-F, 24E, 45G-I. 

Prather Springs, Thousand Springs Valley, Elko County, Nevada. Holotype, 

USNM 874075; paratypes, USNM 860720.

Common name: Upper Thousand Spring pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis hualapaiensis Hershler, Liu, and Stevens, 2016

Pyrgulopsis hualapaiensis Hershler, Liu, and Stevens, 2016b: 74-79, figs. 2-4. Upper 

Peach Springs, outflow just below concrete weir, Hualapai Indian Reservation, 

Mohave County, Arizona. Holotype, USNM 1248611; paratypes, USNM 

1266144.

Common name: Hualapai pyrg [name newly introduced herein].

Distribution: Found only at upper Peach Springs (consisting of two closely proximal 

spring complexes), lower Colorado River basin, Arizona.

Pyrgulopsis hubbsi Hershler, 1998

Pyrgulopsis hubbsi Hershler, 1998: 33, 35-36, figs. 6E, 14A-C, 17F-G, 27F-H. Hiko 

Spring, Pahranagat Valley, Lincoln County, Nevada.

Common name: Hubbs pyrg.

Distribution: Pahranagat Valley, White River drainage, Nevada.
Technical Note 449 43Species Checklist HYDROBIIDAE

Pyrgulopsis humboldtensis Hershler, 1998

Pyrgulopsis humboldtensis Hershler, 1998: 97, figs. 9H, 13E, 22H-J, 42F-I. Springs, East 

Fork Beaver Creek (above Cabin Creek confluence), North Fork Humboldt 

River, Elko County, Nevada. Holotype, USNM 874722; paratypes, USNM 

860718.

Common name: Humboldt pyrg.

Distribution: Humboldt River drainage, northern Nevada.

Pyrgulopsis imperialis Hershler, 1998

Pyrgulopsis imperialis Hershler, 1998: 86-87, figs. 8L, 21J-K, 39D-F. Spring, south 

side of road, Thacker Pass, Kings River Valley, Humboldt County, Nevada. 

Holotype, USNM 874207; paratypes, USNM 860716.

Common name: Kings River pyrg.

Distribution: Two closely proximal springs in Kings River Valley, Nevada.

Pyrgulopsis inopinata Hershler, 1998

Pyrgulopsis inopinata Hershler, 1998: 124-125, figs. 10H, 25D-F, 47H-J. Spring, 

Glenwood, Sevier River drainage, Sevier County, Utah. Holotype, USNM 

883493; paratypes, USNM 860730.

Common name: Carinate Glenwood pyrg.

Distribution: Three springs in the Sevier River drainage, Utah.

Pyrgulopsis intermedia (Tryon, 1865)

Pomatiopsis intermedia Tryon, 1865: 220, pl. 22: fig. 8. Crooked Creek, Owyhee, 

southeastern Oregon. Lectotype, ANSP 27958; paralectotypes, ANSP 396959.

Common name: Crooked Creek springsnail.

Distribution: Great Basin (Barren Valley) and Owyhee River basin, Oregon (Hershler 

and Liu, 2009).

Taxonomy: Hershler (1994), Hershler and Liu (2009).

Pyrgulopsis isolata Hershler and Sada, 1987

Pyrgulopsis isolata Hershler and Sada, 1987: 807-810, figs. 19d, 29c,f, 33d,g, 37-38. 

Spring south of Clay Pits, Ash Meadows, Nye County, Nevada. Holotype, 

USNM 859201; paratypes, UF 93959; USNM 859202.

Common name: Elongate-gland springsnail.

Distribution: Known only from the type locality.

Taxonomy: Hershler (1994).
44 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis kolobensis (Taylor, 1987)

Fontelicella kolobensis Taylor, 1987: 19, fig. 8. Toquerville Springs, Washington County, 

Utah. Holotype, LACM 2216.

Fontelicella pinetorum Taylor, 1987: 20, fig. 9. Spring tributary to Leeds Creek, 

Washington County, Utah. Holotype, LACM 2217.

Common name: Toquerville springsnail.

Distribution: Eastern Great Basin and lower Colorado River drainage (Idaho, Nevada, 

Utah) (Hershler 1998).

Taxonomy: Hershler (1994).

Pyrgulopsis landyei Hershler, 1998

Pyrgulopsis landyei Hershler, 1998: 70-71, figs. 8A, 20B, 35C-F. Spring ca. 1.6 km 

north-northwest of Steptoe Ranch, Steptoe Valley, White Pine County, Nevada. 

Holotype, USNM 892014; paratypes, USNM 860685.

Common name: Landyes pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis lasseni Hershler, Frest, Liu, and Johannes, 2003

Pyrgulopsis lasseni Hershler, Frest, Liu, and Johannes, 2003a: 299, 301-302, figs. 4E, 

20-21. Willow Creek on both sides of wooden foot bridge at a picnic area 

just downstream (west) of Willow Creek Campground, north side of CA 

139, about 0.16 rd. km west of Hayden Hill Cut Off [Cutoff] Road junction, 

Modoc National Forest, Lassen County, California. Holotype, USNM 1004532; 

paratypes, USNM 1004533.

Common name: Willow Creek pyrg.

Distribution: Upper reach of Willow Creek and an associated thermal spring complex, 

upper Pit River basin (California).

Pyrgulopsis lata Hershler, 1998

Pyrgulopsis lata Hershler, 1998: 41, 43, figs. 6H, 12D, 18A, 29A-E. Butterfield Springs, 

White River Valley, Nye County, Nevada. Holotype, USNM 874667; paratypes, 

USNM 860697.

Common name: Butterfield pyrg.

Distribution: Known only from the type locality.
Technical Note 449 45Species Checklist HYDROBIIDAE

Pyrgulopsis lentiglans Hershler, 1998

Pyrgulopsis lentiglans Hershler, 1998: 118-120, figs. 10D, 24G-H, 46C-E. Crittenden 

Springs, Thousand Springs Creek, Elko County, Nevada. Holotype, USNM 

874724; paratypes, USNM 860722.

Common name: Crittenden pyrg.

Distribution: Two springs in the Thousand Springs drainage, northeastern Nevada.

Pyrgulopsis leporina Hershler, 1998

Pyrgulopsis leporina Hershler, 1998: 96-97, figs. 9G, 22G, 42A-E. Springs, Rabbit 

Creek, Humboldt River drainage, Elko County, Nevada. Holotype, USNM 

874336; paratypes, USNM 860717.

Common name: Elko pyrg.

Distribution: Single springs in both the Humboldt River drainage and Ruby Valley, 

Nevada.

Pyrgulopsis licina Hershler, Liu, and Bradford, 2013

Pyrgulopsis licina Hershler, Liu, and Bradford, 2013: 32-34, 36, figs. 3, 4A-B. Spring 

south of Clay Pits, Ash Meadows, Nye County, Nevada. Holotype, USNM 

850347; paratypes, USNM 1204732.

Common name: Curved filament pyrg [name newly introduced herein.]

Distribution: Ash Meadows, Amargosa River basin (Nevada).

Pyrgulopsis limaria Hershler, 1998

Pyrgulopsis limaria Hershler, 1998: 82-83, figs. 8I, 21E-F, 38A-E. Spring brook, Mud 

Meadow drainage, Humboldt County, Nevada. Holotype, USNM 873232; 

paratypes, USNM 860706.

Common name: Squat Mud Meadows pyrg.

Distribution: Mud Meadow drainage, northwestern Nevada.

Pyrgulopsis lindae Hershler, Liu, Babbitt, Kellogg, and Howard, 2016

Pyrgulopsis lindae Hershler, Liu, Babbitt, Kellogg, and Howard, 2016a: 7, 9-10, 

figs. 3A, 4. San Domingo Creek, 3.8 km up flow [upflow] from Dogtown along 

San Domingo Road, Calaveras County, California. Holotype, USNM 905258; 

paratypes, USNM 1254709.

Common name: San Domingo pyrg.

Distribution: Three localities in the upper Calaveras and upper Tuolumne River basins, 

western California.
46 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis lockensis Hershler, 1998

Pyrgulopsis lockensis Hershler, 1998: 57-59, figs. 7C, 12F, 14G-I, 19A, 32A-C. Spring 

at Lockes, Duckwater Valley, Nye County, Nevada. Holotype, USNM 874779; 

paratypes, USNM 860679.

Common name: Lockes pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis longae Hershler, 1995

Pyrgulopsis longae Hershler, 1995: 359, 362, figs. 5E, 17-19. Unnamed spring about 

4.8 km west-southwest of Hallelujah Junction, Long Valley, Lassen County. 

Holotype, USNM 860642; paratypes, USNM 858262.

Common name: Long Valley pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis longiglans Hershler, 1998

Pyrgulopsis longiglans Hershler, 1998: 77-79, figs. 8F, 20M-P, 37A-C. Spring northnorthwest of Holbrook Junction, Antelope Valley, Douglas County, Nevada. 

Holotype, USNM 873409; paratypes, USNM 860701.

Common name: Western Lahontan pyrg.

Distribution: Several basins in northwestern Nevada.

Pyrgulopsis longinqua (Gould, 1855)

Amnicola longinqua Gould, 1855: 130. Colorado Desert (Cienega Grande). Types not 

located [USNM 12112 may be syntypes of this species; Hershler 1994].

Common name: Salton Sea springsnail.

Distribution: A single spring in the Salton Sea basin, eastern California (Hershler 

1994).

Taxonomy: Hershler (1994).

Pyrgulopsis marcida Hershler, 1998

Pyrgulopsis marcida Hershler, 1998: 47-50, figs. 6J, 18D-F, 30A-C. Hardy Springs, 

White River Valley, Nye County, Nevada. Holotype, USNM 873154; paratypes, 

USNM 860711.

Common name: Hardy pyrg.

Distribution: Several springs in White River Valley and one spring in Cave Valley 

(Nevada).
Technical Note 449 47Species Checklist HYDROBIIDAE

Pyrgulopsis marilynae Hershler, Ratcliffe, Liu, Lang, and Hay, 2014

Pyrgulopsis marilynae Hershler, Ratcliffe, Liu, Lang, and Hay, 2014b: 76-78, figs. 3, 

4A-B. Spring 0.48 km north, 0.48 km west of Jordan Canyon, Catron County, 

New Mexico. Holotype, USNM 1135068; paratypes, USNM 1232474.

Common name: Jordan Hot springsnail [name newly introduced herein].

Distribution: Short reach of the Middle Fork Gila River drainage just below 

Jordan Hot Springs (New Mexico).

Pyrgulopsis merriami (Pilsbry and Beecher in Pilsbry, 1892)

Fluminicola merriami Pilsbry and Beecher in Pilsbry, 1892: 143. A warm spring in 

Pahranagat Valley, Nevada. Lectotype, ANSP 67278; paralectotypes, 

ANSP 27782, USNM 123626.

Common name: Pahranagat pebblesnail.

Distribution: White River and Pahranagat Valleys, southern Nevada (Hershler 1998).

Taxonomy: Hershler (1994).

Pyrgulopsis micrococcus (Pilsbry in Stearns, 1893)

Amnicola micrococcus Pilsbry in Stearns, 1893: 277, fig. 1. Small spring in Oasis Valley, 

Nevada. Lectotype, ANSP 67279; paralectotypes, ANSP 368399, USNM 

123622.

Common name: Oasis Valley springsnail.

Distribution: Oasis Valley, upper Amargosa River basin (Nevada) (Hershler et al. 

2013).

Taxonomy: Hershler and Sada (1987), Hershler (1989, 1994), Hershler et al. (2013).

Pyrgulopsis militaris Hershler, 1998

Pyrgulopsis militaris Hershler, 1998: 79-80, figs. 8G, 15G-I, 21A-B, 37D-F. Spring 

west of Soldier Meadow Ranch, Humboldt County, Nevada. Holotype, USNM 

873203; paratypes, USNM 860704.

Common name: Northern Soldier Meadow pyrg.

Distribution: One site each in Soldier Meadow and Craine Creek drainage, 

northwestern Nevada.
48 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis millenaria Hershler, 1998

Pyrgulopsis millenaria Hershler, 1998: 117-118, figs. 10C, 24F, 46A-B. Springs below 

Twentyone Mile Dam, Thousand Springs Creek, Elko County, Nevada. 

Holotype, USNM 874720; paratypes, USNM 860721.

Common name: Twentyone Mile pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis milleri Hershler and Liu, 2010

Pyrgulopsis milleri Hershler and Liu, 2010: 10, 12-15, figs. 6-8. Creek 1.13 km east of 

Pierpoint Spring along California Highway 190, ca. 25.1 km east of Springville, 

Tulare County, California. Holotype, SBMNH 83651; paratypes, SBMNH 

74688, USNM 1132568.

Common name: Pierpoint Spring pyrg.

Distribution: Several springs along a short reach of the South Fork of the Middle Fork 

Tule River.

Pyrgulopsis montana Hershler, 1998

Pyrgulopsis montana Hershler, 1998: 31, 33, figs. 6D, 11B, 12B, 17E, 27C-E. Spring, 

upper Camp Valley, Lincoln County, Nevada. Holotype, USNM 874786; 

paratypes, USNM 860694.

Common name: Camp Valley pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis montezumensis Hershler in Hershler and Landye, 1988

Pyrgulopsis montezumensis Hershler in Hershler and Landye, 1988: 23, 28, 30, 

figs. 10g, 13a,d, 21a-c, 22-23. Montezuma Well, Yavapai County, Arizona. 

Holotype, USNM 859043; paratypes, USNM 859044.

Common name: Montezuma Well springsnail.

Distribution: Known only from the type locality and its upper 110 m of outflow, 

Verde River drainage (lower Colorado River basin).

Taxonomy: Hershler (1994).
Technical Note 449 49Species Checklist HYDROBIIDAE

Pyrgulopsis morrisoni Hershler in Hershler and Landye, 1988

Pyrgulopsis morrisoni Hershler in Hershler and Landye, 1988: 21, 23, figs. 10f, 16e-h, 

19-20. Page Springs, Yavapai County, Arizona. Holotype, USNM 859041; 

paratypes, USNM 859042.

Common name: Page springsnail.

Distribution: Verde Valley, lower Colorado River basin, Arizona.

Taxonomy: Hershler (1994).

Pyrgulopsis nanus Hershler and Sada, 1987

Pyrgulopsis nanus Hershler and Sada, 1987: 802-804, figs. 29a,d, 30-32, 33a,b. Five 

Springs, Ash Meadows, Nye County, Nevada. Holotype, USNM 859191; 

paratypes, UF 93957, USNM 859192.

Common name: Distal-gland springsnail.

Distribution: Four spring brooks in Ash Meadows.

Taxonomy: Hershler (1994).

Pyrgulopsis neritella Hershler, 1998

Pyrgulopsis neritella Hershler, 1998: 68-70, figs. 7L, 11G, 20A, 35A-B. Springs north of 

Steptoe Ranch, Steptoe Valley, White Pine County, Nevada. Holotype, 

USNM 883932; paratypes, USNM 860684.

Common name: Neritiform Steptoe Ranch pyrg.

Distribution: Two closely proximal springs in Steptoe Valley.

Pyrgulopsis nevadensis (Stearns, 1883)

Pyrgula nevadensis Stearns, 1883: 173, unlabeled figure (p. 173). Pyramid Lake, 

Washoe County, Nevada. Lectotype, ANSP 27811; paralectotypes, 

ANSP 375739.

Common name: Corded pyrg.

Distribution: Three lakes in the western Great Basin; has not been collected in many 

years and may now be extinct.

Taxonomy: Hershler and Thompson (1987), Hershler (1994).
50 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis nonaria Hershler, 1998

Pyrgulopsis nonaria Hershler, 1998: 125, 127, figs. 10I, 25G, 48A-C. Spring along east 

side of Ninemile Reservoir, San Pete [Sanpete] County, Utah. Holotype, 

USNM 883566; paratypes, USNM 860731.

Common name: Ninemile pyrg.

Distribution: Two closely proximal springs in the San Pitch River drainage, Utah.

Pyrgulopsis notidicola Hershler, 1998

Pyrgulopsis notidicola Hershler, 1998: 83-84, figs. 8J, 21G-H, 38F-H. Spring, 

Mud Meadow drainage, Humboldt County, Nevada. Holotype, USNM 873215; 

paratypes, USNM 860707.

Common name: Elongate Mud Meadows pyrg.

Distribution: Mud Meadow drainage, northwestern Nevada.

Pyrgulopsis ojaiensis Hershler, Liu, Babbitt, Kellogg, and Howard, 2016

Pyrgulopsis ojaiensis Hershler, Liu, Babbitt, Kellogg, and Howard, 2016b: 10-12, 

figs. 3B, 5. Sisar Creek, 3.4 km up flow [upflow] from Sulphur Springs, 

Ventura County, California. Holotype, SBMNH 74347; paratypes, SBMNH 

46096.

Common name: Sisar pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis orbiculata Hershler, 1998

Pyrgulopsis orbiculata Hershler, 1998: 67-68, figs. 7K, 19L, 34F-G. Spring at Steptoe 

Ranch, Steptoe Valley, White Pine County, Nevada. Holotype, USNM 873196; 

paratypes, 860682.

Common name: Sub-globose Steptoe Ranch pyrg.

Distribution: Several springs in Steptoe Valley.

Pyrgulopsis owensensis Hershler, 1989

Pyrgulopsis owensensis Hershler, 1989: 187-189, figs. 26a-d, 27-31. Unnamed spring 

in canyon south of Piute Creek, Owens Valley, Mono County, California. 

Holotype, USNM 860404; paratypes, USNM 857955.

Common name: Owens Valley springsnail.

Distribution: Springs in northeastern Owens Valley; also found in one spring along the 

East Fork Walker River (Hershler and Pratt 1990).

Taxonomy: Hershler (1994).
Technical Note 449 51Species Checklist HYDROBIIDAE

Pyrgulopsis owyheensis Hershler and Liu, 2009

Pyrgulopsis owyheensis Hershler and Liu, 2009: 13-14, 16-19, figs. 6-9. Owyhee Spring, 

Malheur County, Oregon. Holotype, USNM 883435; paratypes, USNM 

1116915.

Common name: Owyhee upland pyrg.

Distribution: Owyhee and Malheur River basins (Oregon).

Pyrgulopsis papillata Hershler, 1998

Pyrgulopsis papillata Hershler, 1998: 59-60, figs. 7D, 11E, 19B, 32D-E. Big Warm 

Spring, Duckwater Valley, Nye County, Nevada. Holotype, USNM 873185; 

paratypes, USNM 860678.

Common name: Big Warm Spring pyrg.

Distribution: Two springs in Duckwater Valley, Nevada.

Pyrgulopsis peculiaris Hershler, 1998

Pyrgulopsis peculiaris Hershler, 1998: 108-110, figs. 9J, 23A-G, 43D-I. Spring, Maple 

Grove, Round Valley, Millard County, Utah. Holotype, USNM 883933; 

paratypes, USNM 860703.

Common name: Bifid duct pyrg.

Distribution: East-central portion of the Bonneville basin (Nevada, Utah).

Pyrgulopsis pellita Hershler, 1998

Pyrgulopsis pellita Hershler, 1998: 94-96, figs. 9F, 22F, 41D-F. Sullivan Spring, 

Antelope Valley, Eureka County, Nevada. Holotype, USNM 883850; 

paratypes, USNM 860715.

Common name: Antelope Valley pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis perforata Hershler, Liu, and Bradford, 2013

Pyrgulopsis perforata Hershler, Liu, and Bradford, 2013: 36-37, 39. Easternmost spring 

from Scotty’s [Scottys] Castle along California Highway 72, Grapevine Canyon, 

Death Valley, Inyo County, California. Holotype, USNM 853507; paratypes, 

USNM 1204734.

Common name: Scottys Castle pyrg [name newly introduced herein].

Distribution: Lower portion of Grapevine Canyon and Grapevine Mountains 

(lower Amargosa River basin, California).
52 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis perturbata Hershler, 1989

Pyrgulopsis perturbata Hershler, 1989: 189-190, 192, 194, figs. 26e-g, 33-35. Southern 

of two “Northwest Springs,” Fish Slough, Mono County, California. Holotype, 

USNM 860407; paratypes, USNM 853546.

Common name: Fish Slough springsnail.

Distribution: Fish Slough, Owens River drainage.

Taxonomy: Hershler (1994).

Pyrgulopsis pictilis Hershler, 1998

Pyrgulopsis pictilis Hershler, 1998:89-91, figs. 9C, 22B-C, 40C-E. Cain Spring, 

Antelope Valley, Lander County, Nevada. Holotype, USNM 874401; 

paratypes, USNM 860713.

Common name: Ovate Cain Spring pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis pilsbryana (Baily and Baily, 1952)

Amnicola pilsbryi Baily and Baily, 1951: 50, pl. 4: fig. 3. Lifton, Ideal Beach, Bear Lake, 

Idaho. Holotype, ANSP 187691; paratypes, ANSP 368401.

Amnicola pilsbryana Baily and Baily, 1952: 144. Replacement name for 

Amnicola pilsbryi Baily and Baily, 1951; preoccupied in Amnicola by 

Amnicola pilsbryi Walker, 1906.

Common name: Bear Lake springsnail.

Distribution: Bear River basin (Idaho, Utah) (Hershler 1998).

Taxonomy: Hershler (1994).

Pyrgulopsis pisteri Hershler and Sada, 1987

Pyrgulopsis pisteri Hershler and Sada, 1987: 804-807, figs. 29b,e, 33c, 34a-e, 35-36. 

Marsh Spring, Ash Meadows, Nye County, Nevada. Holotype, USNM 859197; 

paratypes, UF 94958, USNM 859198.

Common name: Median-gland springsnail.

Distribution: Three sites in Ash Meadows.

Taxonomy: Hershler (1994).
Technical Note 449 53Species Checklist HYDROBIIDAE

Pyrgulopsis planulata Hershler, 1998

Pyrgulopsis planulata Hershler, 1998: 64-66, figs. 7I, 13B, 15A-C, 19J, 34A-C. 

Spring northwest of Clark Spring, Steptoe Valley, White Pine County, Nevada. 

Holotype, USNM 892023; paratypes, USNM 860686.

Common name: Flat-topped Steptoe pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis plicata Hershler, 1998

Pyrgulopsis plicata Hershler, 1998: 120-121, figs. 10E, 13F, 24I-J, 46F-G. Spring, 

Black Canyon, East Fork Sevier River, Garfield County, Utah. Holotype, 

USNM 883594; paratypes, USNM 860727.

Common name: Black Canyon pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis robusta (Walker, 1908)

Pomatiopsis robusta Walker, 1908: 97, unlabeled figure. Jackson Lake, Wyoming. 

Types have not been located.

Amnicola hendersoni Pilsbry, 1933: 10, pl. 2: figs. 2, 9 10. South of Burns, Oregon. 

Holotype, ANSP 145951; paratypes, ANSP 396668.

Amnicola idahoensis Pilsbry, 1933: 11, pl. 2, figs. 3-5. [Snake River] at Homedale, 

Owyhee County, Idaho. Lectotype, ANSP 152677; paralectotype, ANSP 

396960.

Common name: Jackson Lake springsnail.

Distribution: Portions of the Snake-Columbia River basin and Oregon Great Basin 

(Hershler and Liu 2004a; Lysne et al. 2007).

Taxonomy: Hershler (1994), Hershler and Liu (2004a).

Remarks: Microsatellite evidence suggests that P. robusta may be a polyploid 

(Liu and Hershler 2014).

Pyrgulopsis ruinosa Hershler, 1998

Pyrgulopsis ruinosa Hershler, 1998: 54-56, figs. 7A, 18M, 31A-C. Spring southwest of 

The Crossing, Fish Lake Valley, Esmeralda County, Nevada. Holotype, 

USNM 874307; paratypes, USNM 860700.

Common name: Fish Lake Valley pyrg.

Distribution: Known only from the type locality and may now be extinct (Hershler 

1998).
54 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis rupinicola Hershler, Frest, Liu, and Johannes, 2003

Pyrgulopsis rupinicola Hershler, Frest, Liu, and Johannes, 2003a: 288-292, figs. 4B, 

11-12. Sucker Springs Creek east of California Fish and Game [California 

Fish and Wildlife] Pit River Hatchery on northwest side of access road above 

intake for the hatchery, northwest of Pit River, over 1.6 km southwest of 

Pit 1 Powerhouse (Pacific Gas and Electric), Shasta County, California. 

Holotype, USNM 892187; paratypes, USNM 1004526, USNM 1004527.

Common name: Sucker Springs pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis sadai Hershler, 1998

Pyrgulopsis sadai Hershler, 1998: 87-89, figs. 9A, 21L-N, 39G-J. Spring, Moss Creek, 

Reese River Valley, Lander County, Nevada. Holotype, USNM 874397; 

paratypes, USNM 860702.

Common name: Sadas pyrg.

Distribution: Lahontan basin and Owyhee River drainage, north-central Nevada.

Pyrgulopsis sanchezi Hershler, Liu, and Bradford, 2013

Pyrgulopsis sanchezi Hershler, Liu, and Bradford, 2013: 39-42, figs. 4E-F, 7. Purgatory 

Spring, Ash Meadows, Nye County, Nevada. Holotype, USNM 850333; 

paratypes, USNM 1204735.

Common name: Sanchez pyrg [name newly introduced herein].

Distribution: Five groundwater discharge areas in the Amargosa River basin 

(California, Nevada).

Pyrgulopsis sathos Hershler, 1998

Pyrgulopsis sathos Hershler, 1998: 37, 39, figs. 6F, 12C, 17H-K, 28A-C. Flag Springs, 

White River Valley, Nye County, Nevada. Holotype, USNM 874664; paratypes, 

USNM 860691.

Common name: White River Valley pyrg.

Distribution: White River Valley, Colorado River basin, Nevada.
Technical Note 449 55Species Checklist HYDROBIIDAE

Pyrgulopsis saxatilis Hershler, 1998

Pyrgulopsis saxatilis Hershler, 1998: 111, 113, figs. 9L, 11H, 16A-C, 23K-L, 44F-H. 

Warm Springs, Snake Valley, Millard County, Utah. Holotype, USNM 883237; 

paratypes, USNM 860726.

Common name: Sub-globose Snake pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis serrata Hershler, 1998

Pyrgulopsis serrata Hershler, 1998: 71-72, figs. 8B, 15D-F, 20C-E, 35G-J. Indian Ranch 

Spring, Steptoe Valley, White Pine County, Nevada. Holotype, USNM 874314; 

paratypes, USNM 860719.

Common name: Northern Steptoe pyrg.

Distribution: Several springs in Steptoe Valley.

Pyrgulopsis similis Hershler, Ratcliffe, Liu, Lang, and Hay, 2014

Pyrgulopsis similis Hershler, Ratcliffe, Liu, Lang, and Hay, 2014b: 78-81, figs. 4C-D, 5. 

Spring along Beaver Creek, ca. 0.29 km north and 0.4 km west of confluence 

with Taylor Creek, Catron County, New Mexico. Holotype, USNM 1135064; 

paratypes, USNM 1135065.

Common name: Catron pyrg [name newly introduced herein].

Distribution: Short reach of the East Fork Gila River drainage from just above Wall 

Lake to slightly above the mouth of Burnt Corral Canyon (New Mexico).

Pyrgulopsis simplex Hershler in Hershler and Landye, 1988

Pyrgulopsis simplex Hershler in Hershler and Landye, 1988: 32, figs. 21g-j, 26b, 

27-28. Spring near Strawberry, Gila County, Arizona. Holotype, USNM 

859049; paratypes, USNM 859050.

Common name: Fossil springsnail.

Distribution: Fossil Creek drainage, Verde River drainage, Arizona.

Taxonomy: Hershler (1994).
56 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis sola Hershler in Hershler and Landye, 1988

Pyrgulopsis solus Hershler in Hershler and Landye, 1988: 30, 32, figs. 21d-f, 24, 25, 

26a. Brown Spring, in southern end of Verde Valley, Yavapai County, Arizona. 

Holotype, 859045; paratypes, USNM 859046.

Common name: Brown springsnail.

Distribution: Known only from the type locality in the lower Colorado River basin.

Taxonomy: Hershler (1994).

Pyrgulopsis stearnsiana (Pilsbry, 1899)

Pyrgulopsis stearnsiana Pilsbry, 1899: 124-125. Near Oakland (California). 

Lectotype, ANSP 27961; paralectotypes, ANSP 396667.

Distribution: Central California coastal drainages (Hershler et al. 2016a).

Common name: Yaqui springsnail.

Taxonomy: Hershler (1994).

Remarks: Genetic evidence suggests that P. stearnsiana is paraphyletic relative to 

P. diablensis and P. giulianii, and may require taxonomic revision (Hershler 

et al. 2016a).

Pyrgulopsis sterilis Hershler, 1998

Pyrgulopsis sterilis Hershler, 1998: 53-54, figs. 6L, 18K-L, 30G-I. Spring, 

Hunts Canyon Ranch, Ralston Valley, Nye County, Nevada. Holotype,

USNM 874876; paratypes, USNM 860714.

Common name: Sterile basin pyrg.

Distribution: Three springs in Ralston and Stone Cabin Valleys, southwestern Nevada.

Pyrgulopsis sublata Hershler, 1998

Pyrgulopsis sublata Hershler, 1998: 56-57, figs. 7B, 12E, 18N-O, 31D-F. Wambolt 

Springs, Lake Valley, Lincoln County, Nevada. Holotype, USNM 874681; 

paratypes, USNM 860724.

Common name: Lake Valley pyrg.

Distribution: Known only from the type locality.
Technical Note 449 57Species Checklist HYDROBIIDAE

Pyrgulopsis sulcata Hershler, 1998

Pyrgulopsis sulcata Hershler, 1998: 66-67, figs. 7J, 11F, 19K, 34D-E. Springs north of 

Grass Springs, Steptoe Valley, White Pine County, Nevada. Holotype, 

USNM 874326; paratypes, USNM 860683.

Common name: Southern Steptoe pyrg.

Distribution: Two springs in Steptoe Valley, Nevada.

Pyrgulopsis taylori Hershler, 1995

Pyrgulopsis taylori Hershler, 1995: 362-363, 365, 367, figs. 5F, 20-22. Unnamed spring 

tributary to San Luis Obispo Creek, 4.8 km north of San Luis Obispo, east of 

HW 101, San Luis Obispo County, California. Holotype, USNM 860646; 

paratypes, USNM 883792.

Common name: San Luis Obispo pyrg.

Distribution: San Luis Obispo Creek drainage (California).

Pyrgulopsis thermalis (Taylor, 1987)

Fontelicella thermalis Taylor, 1987: 28, fig. 13. Hot spring on east side of Gila River, 

NE1/4 SW1/4 sec. 17 T13S R13W, Grant County, New Mexico. Holotype, 

LACM 2224; paratypes, ANSP 376026, UF 160941, USNM 954086, UTEP 

10058.

Common name: New Mexico hot springsnail.

Distribution: Thermal springs along the East Fork Gila River, New Mexico.

Taxonomy: Hershler (1994).

Pyrgulopsis thompsoni Hershler in Hershler and Landye, 1988

Pyrgulopsis thompsoni Hershler in Hershler and Landye, 1988: 41-42, figs. 26f, 36-

38. Peterson Ranch Springs, Santa Cruz County, Arizona. Holotype, USNM 

859057; paratypes, USNM 859058.

Common name: Huachuca springsnail.

Distribution: Lower Gila River drainage, southern Arizona; one site in the Rio 

Magdalena basin (Gulf of California drainage), northern Sonora (Mexico)

(Hershler and Landye 1988; Tsai et al. 2007).

Taxonomy: Hershler (1994).

Remarks: Genetic evidence suggests that P. thompsoni is paraphyletic relative to 

P. conica and may require taxonomic revision (Hurt 2004).
58 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis torrida Hershler, Liu, Babbitt, Kellogg, and Howard, 2016

Pyrgulopsis torrida Hershler, Liu, Babbitt, Kellogg, and Howard, 2016b: 12-14, figs. 3C, 

6. Little Sycamore Canyon, creek 3.2 km up flow [upflow] from Hwy 1, 

Ventura County, California. Holotype, SBMNH 74238; paratypes, SBMNH 

46092.

Common name: Little Sycamore pyrg.

Distribution: Known only from the type locality.

Pyrgulopsis transversa Hershler, 1998

Pyrgulopsis transversa Hershler, 1998: 129-130, figs. 10J, 16G-I, 25H-K, 48D-H. 

Sixmile Springs, Simpson Mountains, Old River Bed, Tooele County, Utah. 

Holotype, USNM 883221; paratypes, USNM 860732.

Common name: Southern Bonneville pyrg.

Distribution: West-central Utah (Great Basin).

Pyrgulopsis trivialis (Taylor, 1987)

Fontelicella trivialis Taylor, 1987: 30-32, fig. 14. Spring-fed pond 1,000 ft [0.305 km] N 

of SW corner sec. 5, T5N, R29E, Apache County, Arizona. Holotype, LACM 

2225.

Pyrgulopsis confluentis Hershler in Hershler and Landye, 1988: 32, 35, figs. 3d-g, 26c, 

29-30. Spring on north side of Blanket Creek at Three Forks, Apache County, 

Arizona. Holotype, USNM 859053.

Common name: Black River springsnail.

Distribution: Two spring complexes in the Black River drainage, lower Colorado River 

basin, Arizona (Hurt 2004; Martinez and Myers 2008).

Taxonomy: Hershler (1994).

Pyrgulopsis turbatrix Hershler, 1998

Pyrgulopsis turbatrix Hershler, 1998: 50-53, figs. 6K, 18G-J, 30D-F. Horseshutem 

Springs, Pahrump Valley, Nye County, Nevada. Holotype, USNM 883978; 

paratypes, USNM 860699.

Common name: Southwest Nevada pyrg [original name incorrectly given as Southeast 

Nevada pyrg.]

Distribution: Spring Mountains region, San Bernardino Mountains, central Death 

Valley region (California, Nevada) (Hershler et al. 2013).

Taxonomy: Hershler et al. (2013).
Technical Note 449 59Species Checklist HYDROBIIDAE

Pyrgulopsis umbilicata Hershler, 1998

Pyrgulopsis umbilicata Hershler, 1998: 81, figs. 8H, 21C-D, 37G-I. Spring near mouth 

of Warm Springs canyon, Soldier Meadow, Humboldt County, Nevada. 

Holotype, USNM 873208; paratypes, USNM 860705.

Common name: Southern Soldier Meadow pyrg.

Distribution: Soldier Meadow, Nevada.

Pyrgulopsis variegata Hershler, 1998

Pyrgulopsis variegata Hershler, 1998: 113-115, figs. 10A, 24A-D, 45A-F. Spring ca. 

2.5 km south of South Patterson Spring, Pilot Valley, Box Elder County, Utah. 

Holotype, USNM 883627; paratypes, USNM 860723.

Common name: Northwest Bonneville pyrg.

Distribution: North-central Great Basin and one site in the Snake River drainage 

(Nevada, Utah).

Pyrgulopsis varneri Hershler, Liu, and Sada 2007

Pyrgulopsis varneri Hershler, Liu, and Sada, 2007b: 176-777, 180, figs. 7, 11. Spring 

brook north of Mud Meadow Reservoir, Humboldt County, Nevada. Holotype, 

USNM 1083246; paratypes, USNM 1096917.

Common name: Varners pyrg.

Distribution: Three spring complexes in Soldier Meadow, northeastern Great Basin 

(Nevada).

Pyrgulopsis ventricosa Hershler, 1995

Pyrgulopsis ventricosa Hershler, 1995: 367, 369, 372-373, figs. 5G, 23-25. Unnamed 

creek, Seigler Canyon, 7.4 km south of HW 29 along Seigler Canyon Road, 

Lake County, California. Holotype, USNM 860647; paratypes, USNM 883790.

Common name: Clear Lake pyrg.

Distribution: Seigler Creek drainage, south end of the Clear Lake basin (California).

Pyrgulopsis villacampae Hershler, 1998

Pyrgulopsis villacampae Hershler, 1998: 62-63, figs. 7G, 13A, 19F-G, 33C-E. Little 

Warm Spring, Duckwater Valley, Nye County, Nevada. Holotype, USNM 

873191; paratypes, USNM 860712.

Common name: Duckwater Warm Springs pyrg.

Distribution: Two springs in Duckwater Valley, Nevada.
60 Technical Note 449 HYDROBIIDAE Species Checklist

Pyrgulopsis vinyardi Hershler, 1998

Pyrgulopsis vinyardi Hershler, 1998: 84-86, figs, 8K, 21I, 39A-C. Spring, Willow Creek, 

1.6 km southwest of Willow Creek Reservoir, Squaw Valley drainage, Elko 

County, Nevada. Holotype, USNM 874740; paratypes, USNM 860708.

Common name: Vinyards pyrg.

Distribution: Two springs in the Squaw Valley drainage, north-central Nevada.

Pyrgulopsis wongi Hershler, 1989

Pyrgulopsis wongi Hershler, 1989: 196, 198-202, figs. 41-46. Unnamed western 

spring tributary to Pine Creek, Birchim Canyon, Owens Valley, Inyo County, 

California. Holotype, USNM 860403; paratypes, USNM 857941.

Common name: Wong’s springsnail.

Distribution: Several valleys in the southwestern Great Basin (California, Nevada) 

(Hershler and Pratt 1990; Hershler 1998).

Taxonomy: Hershler (1994).

Remarks: Genetic evidence suggests that P. wongi is composed of two evolutionarily 

distinct lineages and requires taxonomic revision (Liu and Hershler 2007).
Technical Note 449 61

 Family Lithoglyphidae Troschel, 1857

Genus Fluminicola Carpenter, 1864

Figure 21. Fluminicola. A. F. nuttallianus, Willamette River, Oregon (USNM 121467 [lectotype]). 

B. F. coloradoense, Snake River at Clear Lake bridge, Gooding County, Idaho (USNM 905345). 

C. F. virens, Willamette River, French Prairie State Park, Clackamas County, Oregon (USNM 

883183). D. F. warnerensis, Soup Spring, Pit River drainage, Modoc County, California (USNM 

1020654). E. F. virginius, spring source of Hardscrabble Creek, Pyramid Lake basin, Washoe 

County, Nevada (USNM 874105). F. F. neritoides, Springs at Willow Creek Campground, Pit River 

drainage, Lassen County, California (USNM 883563, holotype). Scale bars, 1.0 mm.

Species Checklist

LITHOGLYPHIDAE
62 Technical Note 449 LITHOGLYPHIDAE Species Checklist

Fluminicola Carpenter, 1864: 676 [type species: Paludina nuttalliana Lea, 1838;

original designation.]

Heathilla Hannibal, 1912: 186-187 [type species: Paludina seminalis Hinds, 1842; 

original designation].

Remarks: Fluminicola is widely distributed in the northwestern United States and 

also present in portions of western Canada. Members of this genus typically 

live in rivers, springs, and large streams and are occasionally found in “spring 

influenced” lacustrine habitats. Fluminicola is composed of two anatomically 

distinctive subgroups (Hershler and Frest 1996; Hershler and Liu 2012b), 

which form evolutionarily distinct and genetically divergent lineages 

evidenced by mitochondrial DNA data (Hershler and Liu 2012b). One of 

these is composed of two species from the Snake-Columbia River drainage 

(F. gustafsoni, F. virens), while the other contains the remaining congeners 

(for which data are available). This polyphyletic assemblage continues to be 

classified as a single genus pending resolution of the phylogenetic relationships 

of the poorly known type species.

Fluminicola ahjumawi Hershler, Liu, Frest, and Johannes, 2007

Fluminicola ahjumawi Hershler, Liu, Frest, and Johannes, 2007c: 402, 404, figs. 7H, 16. 

Spring south of Sam Wolfin Spring, south of Pit River and powerlines,

1.93 km southwest of Pit 1 Powerhouse, Pacific Gas and Electric land,

Shasta County, California. Holotype, USNM 1020691;

paratypes, USNM 1020699.

Common name: Sculpin snail.

Distribution: Widely ranging in the lower Pit River basin and also present at one site in 

the upper Pit River basin, California.

Fluminicola anserinus Hershler, Liu, Frest, and Johannes, 2007

Fluminicola anserinus Hershler, Liu, Frest, and Johannes, 2007c: 409-410, 412, 

figs. 6F, 7J, 21. Unnamed spring on west side of Goose Valley Road and Goose 

Valley, north of Goose Creek ca. 0.8 km, Shasta County, California. Holotype, 

USNM 1020727; paratypes, USNM 1020728.

Common name: Striated hydrobe.

Distribution: Lower Pit River basin from Lake Britton (including Burney Creek 

drainage) to vicinity of Chalk Mountain, California.
Technical Note 449 63Species Checklist LITHOGLYPHIDAE

Fluminicola caballensis Hershler, Liu, Frest, and Johannes, 2007

Fluminicola caballensis Hershler, Liu, Frest, and Johannes, 2007c: 398, 400, figs. 7F, 

14. Davis Creek at crossing of road 0.32 km south of junction with FS22, 

Lassen National Forest, Lassen County, California. Holotype, USNM 1020675; 

paratypes, USNM 1020676.

Common name: Horse Creek pebblesnail.

Distribution: Horse Creek and Bob Creek drainages (lower Pit River basin), California.

Fluminicola coloradoense Morrison, 1940

Fluminicola coloradoense Morrison, 1940: 125-126. Green River, Wyoming. 

Holotype, USNM 526631; paratypes, USNM 526576.

Fluminicola coloradensis.—Hershler and Frest, 1996: 8 [unjustified emendation of 

coloradoense].

Common name: Green River pebblesnail.

Distribution: Upper Green River basin (Wyoming), northeastern Great Basin 

(Idaho, Utah), upper Snake River basin (Idaho, Oregon) (Liu et al. 2013a).

Taxonomy: Hershler and Frest (1996), Liu et al. (2013a).

Fluminicola dalli (Call, 1884)

Amnicola dalli Call, 1884: 21, 45-47, figs. 2-3, pl. VI: figs. 4-6. Small tributary to 

Pyramid Lake, near the north end, at Symons Ranch [Nevada]. Lectotype, 

MCZ 2087; paralectotypes, MCZ 2088.

Common name: Pyramid Lake pebblesnail.

Distribution: Northern portion of the Pyramid Lake basin, Nevada (Hershler and 

Frest 1996)

Taxonomy: Hershler and Frest (1996).

Fluminicola erosus Hershler, Liu, Frest, and Johannes, 2007

Fluminicola erosus Hershler, Liu, Frest, and Johannes, 2007c: 394-396, fig. 6C, 7D, 

12. Moderate-sized unnamed spring, ca. 0.4 km southeast of Smokey Charlie 

[Charley] Spring and 0.32 km west of Modoc County 63, at source next to 

homestead cabin, Modoc County, California. Holotype, 1020663; paratypes, 

USNM 1020664.

Common name: Smokey Charlie [Charley] pebblesnail.

Distribution: Two closely proximal springs in the upper Pit River basin, California.
64 Technical Note 449 LITHOGLYPHIDAE Species Checklist

Fluminicola favillaceus Hershler, Liu, Frest, and Johannes, 2007

Fluminicola favillaceus Hershler, Liu, Frest, and Johannes, 2007c: 396-398, figs. 6D, 

7E, 13. Ash Creek south culvert channel on west side of Ash Valley Road 

(Modoc County [Highway] 527), Crown D Ranch, Lassen County, California. 

Holotype, USNM 1020669; paratypes, USNM 1020670.

Common name: Ash Valley pebblesnail.

Distribution: Ash Creek, upper Pit River basin, California.

Fluminicola fremonti Hershler, Liu, Frest, and Johannes, 2007

Fluminicola fremonti Hershler, Liu, Frest, and Johannes, 2007c: 386, figs. 6A, 7A, 8. 

Hunters Spring on the north side of FS28, Fremont National Forest 

[Fremont-Winema National Forest], Lake County, Oregon. Holotype, 

USNM 1020662; paratypes, USNM 1020661.

Common name: Fremont pebblesnail.

Distribution: Found only at the type locality in the Thomas Creek drainage, 

Goose Lake basin, Oregon.

Fluminicola fuscus (Haldeman, 1841)

Anculosa fusca Haldeman, 1841: cover. Oregon. Type has not been located.

Amnicola hindsi Baird, 1863: 63. River Kootanie [sic, misspelling of Kootenai]. 

Lectotype, NHMUK 1863.2.4.17A; paralectotypes, NHMUK 1863.2.4.17.

Fluminicola columbiana Pilsbry, 1899: 125. Columbia River, near Wallula. 

Lectotype, ANSP 27767; paralectotypes, ANSP 398349.

Common name: Ashy pebblesnail.

Distribution: Lower Snake River, Columbia River, and large tributaries (Idaho, Oregon, 

Washington, British Columbia [historically] (Hershler and Frest 1996; 

Liu et al. 2013a).

Taxonomy: Hershler and Frest (1996).

Fluminicola gustafsoni Hershler and Liu, 2012

Fluminicola gustafsoni Hershler and Liu, 2012b: 324-325, 327-328, figs. 2-4. 

Salmon River at Pine Bar Rapids, Idaho County, Idaho. Holotype, 

USNM 905409; paratypes, USNM 1175400.

Common name: Gustafson pebblesnail [name newly introduced herein].

Distribution: Clearwater River, lower Salmon River, and the short reach of the 

Snake River between the mouths of these streams (Idaho, Washington).
Technical Note 449 65Species Checklist LITHOGLYPHIDAE

Fluminicola insolitus Hershler, 1999

Fluminicola insolitus Hershler, 1999: 319, 321, figs. 8D, 9, 10A-C. Page Springs, 

Donner und Blitzen River drainage, Harney County, Oregon. Holotype, 

USNM 883466; paratypes, USNM 860757.

Common name: Donner und Blitzen pebblesnail.

Distribution: Found only at the type locality.

Fluminicola lunsfordensis Hershler, Liu, Frest, and Johannes, 2007

Fluminicola lunsfordensis Hershler, Liu, Frest, and Johannes, 2007c: 393-394, figs. 7C, 

11. Lunsford Spring above Lunsford Springs Road, source of Canyon Creek, 

Modoc County, California. Holotype, USNM 1020688; paratypes, 

USNM 1020689.

Common name: Lunsford pebblesnail.

Distribution: Known only at the type locality in the upper Pit River basin.

Fluminicola minutissimus Pilsbry, 1907

Fluminicola minutissimus Pilsbry, 1907: 76, pl. XI; fig 4. Price Valley, Weiser Canyon, 

Washington County, Idaho. Lectotype, ANSP 94273; paralectotypes, 

ANSP 398350.

Common name: Pixie pebblesnail.

Distribution: Known only from type locality, lower Snake River basin.

Remarks: Fluminicola minutissimus is known only from shells. It has not been 

collected subsequent to its first discovery and may be extinct (Hershler and 

Frest 1996).

Fluminicola modoci Hannibal, 1912

Fluminicola modoci Hannibal, 1912: 187, pl. 8: fig. 30. Fletchers [Fletcher] Spring, 

south end, Goose Lake, California. Lectotype (and paralectotypes), CAS 

60798.

Common name: Modoc pebblesnail.

Distribution: Two springs in the Goose Lake basin (Hershler 1999).

Taxonomy: Hershler and Frest (1996).
66 Technical Note 449 LITHOGLYPHIDAE Species Checklist

Fluminicola multifarius, Hershler, Liu, Frest, and Johannes, 2007

Fluminicola multifarius, Hershler, Liu, Frest, and Johannes, 2007c: 415, 417-419, figs. 

7M, 24-25. Big Springs (source) at Big Springs City Park northwest of the city 

of Mount Shasta, south of Spring Hill, Siskiyou County, California. Holotype, 

USNM 883782; paratypes, USNM 1020753.

Common name: Shasta pebblesnail.

Distribution: Sacramento River headwater region and upper reaches of the McCloud 

River drainage, California.

Fluminicola neritoides Hershler, Liu, Frest, and Johannes, 2007

Fluminicola neritoides Hershler, Liu, Frest, and Johannes, 2007c: 400-402, figs. 6E, 7G, 

15. Springs at Willow Creek Campground along CA [Highway] 139, Lassen 

County, California. Holotype, USNM 883563; paratypes, USNM 1020682.

Common name: Willow Creek pebblesnail.

Distribution: Willow Creek and associated thermal springs (upper Pit River basin), 

California.

Fluminicola nuttallianus (Lea, 1838)

Paludina nuttalliana Lea, 1838: 101, pl. XXIII: fig. 89. Wahlamat [sic, misspelling of 

Willamette] River, near its junction with the Columbia River [Oregon]. 

Lectotype, USNM 121467; paralectotypes, USNM 860648.

Common name: Dusky pebblesnail.

Distribution: Restricted to the type locality near the mouth of the Willamette River 

(Oregon).

Taxonomy: Hershler and Frest (1996).

Remarks: This species is known only from shells. It has not been collected for more 

than 100 years and may be extinct (Hershler and Frest 1996).

Fluminicola potemicus, Hershler, Liu, Frest, and Johannes, 2007

Fluminicola potemicus, Hershler, Liu, Frest, and Johannes, 2007c: 412-414, figs. 7K, 22. 

Unnamed spring pool on the west side of FS27 (Fenders Ferry Road), ca. 

10.8 rd km north of CA 299 junction, 4.18 km north of FS27 bridge over 

Potem Creek, west side of Potem Creek, inholding in Shasta National Forest 

[Shasta-Trinity National Forest], Shasta County, California. Holotype, 

USNM 1020718; paratypes, USNM 1020719.

Common name: Potem Creek pebblesnail.

Distribution: Found only at the type locality.
Technical Note 449 67Species Checklist LITHOGLYPHIDAE

Fluminicola scopulinus, Hershler, Liu, Frest, and Johannes, 2007

Fluminicola scopulinus, Hershler, Liu, Frest, and Johannes, 2007c: 414, figs. 7L, 23. 

Northernmost of three springs and runs southwest of Popcorn Spring, west 

side of North Fork Castle Creek, 0.35 km from FS25 along FS38N35Y (west 

side), Shasta National Forest, Shasta County, California. Holotype, USNM 

1020721; paratypes, USNM 1020722.

Common name: Castle Creek pebblesnail.

Distribution: Three closely proximal springs in the Castle Creek drainage, Sacramento 

River headwaters, California.

Fluminicola seminalis (Hinds, 1842)

Paludina seminalis Hinds, 1842: 83-84. Rio Sacramento, California. Type has not been 

located.

Lithoglyphus cumingii Frauenfeld, 1863: 195. California. Lectotype, NHMUK 1993047; 

paralectotypes, NMW.

Common name: Nugget pebblesnail.

Distribution: Sacramento River basin from near its mouth (historically) upstream into 

the Pit River basin, California (Taylor 1981; Hershler and Frest 1996; Hershler 

et al. 2007c).

Taxonomy: Hershler and Frest (1996).

Fluminicola turbiniformis (Tryon, 1865)

Amnicola turbiniformis Tryon, 1865: 219, pl. 22: fig. 5. West side of Steens Mountains, 

Harney County, Oregon. Lectotype, ANSP 27779; paralectotypes, ANSP 

398352.

Common name: Turban pebblesnail.

Distribution: Northwestern Great Basin (California, Nevada, Oregon) (Taylor 1966b; 

Hershler, 1999).

Taxonomy: Hershler and Frest (1996).
68 Technical Note 449 LITHOGLYPHIDAE Species Checklist

Fluminicola umbilicatus Hershler, Liu, Frest, and Johannes, 2007

Fluminicola umbilicatus Hershler, Liu, Frest, and Johannes, 2007c: 407, 409, figs. 7I, 19. 

Big Spring, tributary of Hat Creek, south of Old Station (PO [Post Office]) at 

the south end of Hat Creek Hill, flowing from beneath CA [Highway] 89/44 on 

the south side, Lassen National Forest, Shasta County, California. Holotype, 

USNM 1020705; paratypes, USNM 1020706.

Common name: Goose Valley pebblesnail.

Distribution: Lost Creek and upper section of Hat Creek drainage (lower Pit River 

basin), California.

Fluminicola virens (Lea, 1838)

Paludina virens Lea, 1838: 91, pl. XXIII: fig. 93. Wahlamat [sic, misspelling of 

Willamette] (River), near its junction with the Columbia River [Oregon]. 

Lectotype, USNM 121431; paralectotypes, USNM 860649.

Paludina nuclea Lea, 1838: 91, pl. XXIII: fig. 103. Wahlamat [sic] (River), near its 

junction with the Columbia River. Type has not been located.

Common name: Olympia pebblesnail.

Distribution: Willamette River (and large tributary streams), from Corvallis to its 

mouth; lower Columbia River below Portland (Oregon, Washington) (Hershler 

and Frest, 1996).

Taxonomy: Hershler and Frest (1996).

Fluminicola virginius Hershler, 1999

Fluminicola virginius Hershler, 1999: 323-324, figs. 8H, 10D-F, 11. Unnamed 

(“waterfall”) spring, source of Hardscrabble Creek, Pyramid Lake basin, 

Washoe County, Nevada. Holotype, USNM 874902; paratypes, USNM 860758.

Common name: Virginia Mountains pebblesnail.

Distribution: Found only at the type locality.

Fluminicola warnerensis Hershler, Liu, Frest, and Johannes, 2007

Fluminicola warnerensis Hershler, Liu, Frest, and Johannes, 2007c: 388, 389, 392-393, 

figs. 6B, 7B, 10. Parsnip Springs, south of Southern Warner Road (FS64, also 

designated as 39NO1), in wet, open meadow, Modoc National Forest, Lassen 

County, California. Holotype, USNM 1020652; paratypes, USNM 1020653.

Common name: Topaz pebblesnail.

Distribution: South Fork Pit River drainage, along Pit River just above Big Valley 

(California).
Technical Note 449 69

 Family Tateidae Thiele, 1925

Genus Potamopyrgus Stimpson, 1865

Figure 22. Potamopyrgus antipodarum. A, B. Cassia Creek at Conner Bridge, Snake River 

drainage, Cassia County, Idaho (USNM 894703). C. Snake River, opposite Pilgrim Spring, 

Gooding County, Idaho (USNM 1097249). Scale bars, 1.0 mm.

Pomatopyrgus Stimpson, 1865: 53-54 [type species: Melania corolla Gould, 1847; 

= Amnicola antipodarum Gray, 1843].

Potamopyrgus antipodarum (Gray, 1843)

Amnicola antipodanum [sic] Gray, 1843: 241. New Zealand, in fresh water. Syntypes, 

NHMUK (per Ponder 1988).

Amnicola antipodarum Gray, 1844: 356 [correction of original spelling].

Common name: New Zealand mudsnail.

Distribution: Native to New Zealand (Ponder 1988), widely introduced in Australia, 

Europe, North America, and parts of Asia (Alonso and Castro-Díez 2012). 

Recorded from all of the western United States except New Mexico 

(USGS 2016).

Remarks: See Ponder (1988) for a list of the numerous (extralimital) junior synonyms 

of this species.

Species Checklist

TATEIDAE
70 Technical Note 449

 Family uncertain

Genus Pristinicola Hershler, Frest, Johannes, Bowler, and Thompson, 1994

Figure 23. Pristinicola hemphilli. A. Mountain springs, Silvies River drainage, 

Grant County, Oregon (USNM 883878). B. Spring, Price Valley, 4.8 km northwest 

of Hwy 95, Adams County, Idaho (USNM 874184). Scale bar, 1.0 mm.

Pristinicola Hershler, Frest, Johannes, Bowler, and Thompson, 1994: 222-225 

[type species: Bythinella hemphilli Pilsbry, 1890; original designation].

Remarks: Pristinicola is a monotypic genus that is distributed in the northwestern 

United States. It cannot be assigned to any currently recognized 

truncatelloidean family and does not appear to be closely related to other 

North American taxa (Hershler et al. 1994; Hershler and Liu 2012b).

Pristinicola hemphilli (Pilsbry, 1890)

Bythinella hemphilli Pilsbry, 1890: 63-64. Near Kentucky Ferry, Snake River, 

Washington [precise location unknown]. Lectotype, ANSP 32276; 

paralectotypes, 

ANSP 368405.

Common name: Pristine pyrg.

Distribution: Lower Snake River and Columbia River basins (Idaho, Oregon, 

Washington), minor Pacific Coastal drainages (Washington) (Hershler et al. 

1994).

Taxonomy: Hershler et al. (1994).

UNCERTAIN

Species Checklist
Technical Note 449 71

Figure 24. Taylorconcha. A. T. serpenticola, Snake River at Clover Creek confluence, Elmore 

County, Idaho (USNM 874590). B. T. serpenticola, Thousand Springs (north springs), Snake River 

drainage, Gooding County, Idaho (USNM 874588 [paratype]). C. T. insperata, Owyhee River at the 

Cave, Malheur County, Oregon (USNM 1018182 [holotype]). Scale bars, 1.0 mm.

Taylorconcha Hershler, Frest, Johannes, Bowler, and Thompson, 1994: 233 

[type species: Taylorconcha serpenticola Hershler, Frest, Johannes, Bowler 

and Thompson, 1994; original designation.]

Remarks: Taylorconcha is distributed in the Owyhee River and in the Snake River and 

associated springs. This genus is morphologically similar to the lithoglyphids 

(Hershler et al. 1994) and was delineated as sister to the lithoglyphid clade in 

a recent analysis of COI sequence data (Hershler and Liu 2012b); additional 

studies that include a broader range of truncatelloidean taxa are needed to 

confidently ascertain its relationships.

Taylorconcha insperata Hershler, Liu, Frest, Johannes, and Clark, 2006

Taylorconcha insperata Hershler, Liu, Frest, Johannes, and Clark, 2006: 175, figs. 6B, 

7-8. Owyhee River at the Cave, 1.6 river km below the mouth of Big Antelope 

Creek at river km 269.4, Malheur County, Oregon. Holotype, USNM 1018182; 

paratypes, USNM 1071513.

Common name: Unexpected pebblesnail.

Distribution: Middle Owyhee River and lower Snake River (Hells Canyon) (Idaho, 

Oregon).

Genus Taylorconcha Hershler, Frest, Johannes, Bowler, and Thompson, 1994

Species Checklist

UNCERTAIN
72 Technical Note 449 UNCERTAIN Species Checklist

Taylorconcha serpenticola Hershler, Frest, Johannes,

Bowler, and Thompson, 1994

Taylorconcha serpenticola Hershler, Frest, Johannes, Bowler, and Thompson, 1994: 

233, 235, 237, 239, 240-242, figs. 1 (bottom row), 8-12. Thousand Springs 

(north springs), Gooding County, Idaho. Holotype, USNM 860583; paratypes, 

UF 194616, USNM 874588.

Common name: Bliss Rapids snail.

Distribution: Snake River and associated springs between Twin Falls and the 

Indian Cove Bridge, Idaho (Hershler et al. 1994).
Technical Note 449 73

Literature Cited

Alonso, A., Castro-Díez, P. 2012. The exotic aquatic mud snail Potamopyrgus 

antipodarum (Hydrobiidae, Mollusca): state of the art of a worldwide invasion. 

Aquatic Sciences 74:375-383.

ANS [Aquatic Nuisance Species] Task Force. 2005. New Zealand mudsnail 

(Potamopyrgus antipodarum). Available from http://www.anstaskforce.gov/

spoc/nzms.php (accessed 3/2/2016).

Arnold, R. 1903. The paleontology and stratigraphy of the marine Pliocene and 

Pleistocene of San Pedro, California. Memoirs of the California Academy of 

Sciences 3:1-420, 37 plates.

Baily, J.L. Jr., Baily, R.I. 1951. Further observations on the Mollusca of the relict lakes 

in the Great Basin. Nautilus 65(2):46-53, pl. 4.

Baily, J.L. Jr., Baily, R.I. 1952. Amnicola pilsbryana, new name. Nautilus 65(4):144.

Baird, W. 1863. Descriptions of some new species of shells, collected at Vancouver 

Island and in British Columbia, by J.K. Lord, Esq., naturalist to the British 

North-American Boundary Commission, in the years 1858-1862. Proceedings 

of the Zoological Society of London 31:66-70.

Berry, S.S. 1947. A new Pyrgulopsis from Oregon. Nautilus 60:76-78, pl. 7: figs. 6-7.

Burch, J.B. 1982. North American freshwater snails. Identification keys, generic 

synonymy, supplemental notes, glossary, references, index. Walkerana 1:217-

365.

Burch, J.B., Tottenham, J.L. 1980. North American freshwater snails. Species list, 

ranges and illustrations. Walkerana 1:81-215.

Calkins, W.W. 1880. New freshwater Mollusca. Valley Naturalist 2:6.

Call, R.E. 1884. On the Quaternary and Recent Mollusca of the Great Basin with 

descriptions of new forms; introduced by a sketch of the Quaternary lakes 

of the Great Basin by G.K. Gilbert. United States Geological Survey 

Bulletin 11:1-66.

Call, R.E., Pilsbry, H.A. 1886. On Pyrgulopsis, a new genus of rissoid mollusk, with 

descriptions of two new forms. Proceedings of the Davenport Academy of 

Natural Sciences 5:9-14.

Literature Cited
74 Technical Note 449

Carpenter, P.P. 1864. Supplementary 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 33:517-686.

Chamberlain, R.V., Berry, E. 1933. Mollusks of the Pliocene deposits at Collinston, 

Utah. Nautilus 47:25-29. 

Clarke, A.H. 1981. The freshwater molluscs of Canada. Ottawa, Canada: National 

Museum of Natural Sciences, National Museums of Canada. 446 p.

Conrad, T.A. 1855. Description of a new species of Melania. Proceedings of the 

Academy of Natural Sciences of Philadelphia 7:269.

Cowie, R.H. 1997. Catalog and bibliography of the nonindigeneous nonmarine snails 

and slugs of the Hawaiian Islands. Bishop Museum Occasional Papers 50:1-66.

Davis, G.M., Mazurkiewicz, M., Mandracchia, M. 1982. Spurwinkia: morphology, 

systematics, and ecology of a new genus of North American marshland 

Hydrobiidae (Mollusca: Gastropoda). Proceedings of the Academy of Natural 

Sciences of Philadelphia 134:143-177.

Frauenfeld, G.R. von. 1863. Die Arten der Gattung Lithoglyphus Mhlf., Paludinella Pf., 

Assiminea Gray, in der Kaiserlichen und Cuming’s Sammlung. Verhandlungen 

der Kaiserlich-Königlichen Zoologische-Botanischen Gesellschaft in 

Wien 13:193-212.

Gill, T. 1863. Systematic arrangement of the mollusks of the family Viviparidae, and 

others, inhabiting the United States. Proceedings of the Academy of Natural 

Sciences of Philadelphia 15:33-40.

Gould, A.A. 1839. [Descriptions of thirteen new species of shells.] Silliman’s American 

Journal of Science and Arts 38:196-197.

Gould, A.A. 1847. [Shells collected by the United States Exploring Expedition under 

the command of Charles Wilkes.] Proceedings of the Boston Society of Natural 

History 2:222-225.

Gould, A.A. 1855. New species of land and fresh-water shells from western (N.) 

America. Proceedings of the Boston Society of Natural History 5:127-130.

Gray, J.E. 1840. Mollusks. In: British Museum, Synopsis of the contents of the British 

Museum. 42nd edition. London: G. Woodfall and Son:86-89, 106-156.

Literature Cited
Technical Note 449 75

Gray, J.E. 1843. Catalogue of the species of Mollusca and their shells, which have 

hitherto been recorded as found at New Zealand, with the description of some 

lately discovered species. In: E. Dieffenbach, Travels in New Zealand; with 

contributions to the geography, geology, botany, and natural history of that 

country. Volume II. London: John Murray:228-265.

Gray, J.E. 1844. Catalogue des Mollusques et do leurs Coquilles, trouvés jusqu’à 

présent à la Nouvelle-Zélande, avec la description des espèces récemment 

découvertes. Revue Zoologique 7:346-358.

Gregg, W.O., Taylor, D.W. 1965. Fontelicella (Prosobranchia: Hydrobiidae), a new 

genus of west American freshwater snails. Malacologia 3:103-110.

Haldeman, S.S. 1840. A monograph of the Limniades and other freshwater univalve 

shells of North America. No. 1. Philadelphia: J. Dobson: 3.

Haldeman, S.S. 1841. A monograph of the Limniades and other freshwater univalve 

shells of North America. No. 3. Philadelphia: J. Dobson: cover.

Haldeman, S.S. 1844. A monograph of the freshwater univalve mollusca of the United 

States, including notices of species in other parts of North America. No. 7. 

Philadelphia: Carey and Hart: cover.

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 10:112-211.

Herrmannsen, A.N. 1846. Indicis generum malacozoorum primordia. Nomina 

subgenerum, generum, familiarum, tribuum, ordinum, classium; adjectis 

auctoribus, temporibus, locis systematicis atque literariis, etymis, synonymis. 

Praetermittuntur Cirripedia, Tunicata et Rhizopoda. Volume 1. Cassellis: 

Sumptibus and Typis Theodori Fischeri. 637 p.

Hershler, R. 1985. Systematic revision of the Hydrobiidae (Gastropoda: Rissoacea) of 

the Cuatro Cienegas basin, Coahuila, Mexico. Malacologia 26:31-123.

Hershler, R. 1989. Springsnails (Gastropoda: Hydrobiidae) of Owens and Amargosa 

River (exclusive of Ash Meadows) drainages, Death Valley system, CaliforniaNevada. Proceedings of the Biological Society of Washington 102:176-248.

Hershler, R. 1990. Pyrgulopsis bruneauensis, a new springsnail (Gastropoda: 

Hydrobiidae) from the Snake River Plain, southern Idaho. Proceedings of the 

Biological Society of Washington 103:803-814.

Literature Cited
76 Technical Note 449

Hershler, R. 1994. A review of the North American freshwater snail genus Pyrgulopsis

(Hydrobiidae). Smithsonian Contributions to Zoology 554:1-115.

Hershler, R. 1995. New freshwater snails of the genus Pyrgulopsis (Rissooidea: 

Hydrobiidae) from California. Veliger 38:343-373.

Hershler, R. 1998. A systematic review of the hydrobiid snails (Gastropoda: 

Rissooidea) of the Great Basin, western United States. Part I. Genus 

Pyrgulopsis. Veliger 41:1-132.

Hershler, R. 1999. A systematic review of the hydrobiid snails (Gastropoda: 

Rissooidea) of the Great Basin, western United States. Part II. Genera 

Colligyrus, Eremopyrgus, Fluminicola, Pristinicola, and Tryonia. 

Veliger 42:306-337.

Hershler, R. 2001. Systematics of the North and Central American aquatic snail genus 

Tryonia (Rissooidea: Hydrobiidae). Smithsonian Contributions to 

Zoology 612:1-53.

Hershler, R., Davis, C.L., Kitting, C.L., Liu, H.-P. 2007a. Discovery of introduced and 

cryptogenic cochliopid gastropods in the San Francisco Estuary, California. 

Journal of Molluscan Studies 73:323-332.

Hershler, R., Frest, T.J. 1996. A review of the North American freshwater snail genus 

Fluminicola (Hydrobiidae). Smithsonian Contributions to Zoology 583:1-41.

Hershler, R., Frest, T.J., Johannes, E.J., Bowler, P.A., Thompson, F.G. 1994. Two new 

genera of hydrobiid snails (Prosobranchia: Rissooidea) from the northwestern 

United States. Veliger 37:221-243.

Hershler, R., Frest, T.J., Liu, H.-P., Johannes, E.J. 2003a. Rissooidean snails from the Pit 

River basin, California. Veliger 46:275-304.

Hershler, R., Landye, J.J. 1988. Arizona Hydrobiidae. Smithsonian Contributions to 

Zoology 459:1-63.

Hershler, R., Liu, H.-P. 2004a. Taxonomic reappraisal of species assigned to the North 

American freshwater gastropod subgenus Natricola (Rissooidea: Hydrobiidae). 

Veliger 47:66-81.

Hershler, R., Liu, H.-P. 2004b. A molecular phylogeny of aquatic gastropods provides a 

new perspective on biogeographic history of the Snake River region. Molecular 

Phylogenetics and Evolution 32:927-937.

Literature Cited
Technical Note 449 77

Hershler, R., Liu, H.-P. 2008. Ancient vicariance and recent dispersal of springsnails 

(Hydrobiidae: Pyrgulopsis) in the Death Valley system, California-Nevada. In: 

Reheis, M.C., Hershler, R., Miller, D.M., eds. Late Cenozoic drainage history of 

the southwestern Great Basin and lower Colorado River region: geologic and 

biotic perspectives. Geological Society of America Special Paper 439:91-101.

Hershler, R., Liu, H.-P. 2009. New species and records of Pyrgulopsis (Gastropoda: 

Hydrobiidae) from the Snake River basin, southeastern Oregon: further 

delineation of a highly imperiled fauna. Zootaxa 2006:1-22.

Hershler, R., Liu, H.-P. 2010. Two new, possibly threatened species of Pyrgulopsis

(Gastropoda: Hydrobiidae) from southwestern California. Zootaxa 2343:1-17.

Hershler, R., Liu, H.-P. 2012a. A new species of springsnail (Pyrgulopsis) from the 

Owyhee River basin, Nevada. Western North American Naturalist 72:21-31.

Hershler, R., Liu, H.-P. 2012b. Molecular phylogeny of the western North American 

pebblesnails, genus Fluminicola (Rissooidea: Lithoglyphidae), with description 

of a new species. Journal of Molluscan Studies 78:321-329.

Hershler, R., Liu, H.-P., Babbitt, C., Kellogg, M.G., Howard, J.K. 2016a. Three new 

species of western California springsnails previously confused with Pyrgulopsis

stearnsiana (Caenogastropoda, Hydrobiidae). ZooKeys 601:1-19.

Hershler, R., Liu, H.-P., Bradford, C. 2013. Systematics of a widely distributed western 

North American springsnail, Pyrgulopsis micrococcus (Caenogastropoda, 

Hydrobiidae), with descriptions of three new congeners. Zookeys 330:27-52.

Hershler, R., Liu, H.-P., Carlton, J.T., Cohen, A.N., Davis, C.B., Sorensen, J., Weedman, 

D. 2015a. New discoveries of introduced and cryptogenic fresh and brackish 

water gastropods (Caenogastropoda: Cochliopidae) in the western United 

States. Aquatic Invasions 10:147-156.

Hershler, R., Liu, H.-P., Frest, T.J., Johannes, E.J. 2007c. Extensive diversification 

of pebblesnails (Lithoglyphidae: Fluminicola) in the upper Sacramento 

River basin, northwestern USA. Zoological Journal of the Linnean 

Society 149:371-422.

Hershler, R., Liu, H.-P., Frest, T.J., Johannes, E.J., Clark, W.H. 2006. Genetic structure 

of the western North American aquatic gastropod genus Taylorconcha and 

description of a second species. Journal of Molluscan Studies 72:167-177.

Literature Cited
78 Technical Note 449

Hershler, R., Liu, H.-P., Gustafson, D.L. 2008. A second species of Pyrgulopsis

(Hydrobiidae) from the Missouri River basin, with molecular evidence 

supporting faunal origin through Pliocene stream capture across the northern 

continental divide. Journal of Molluscan Studies 74:403-413.

Hershler, R., Liu, H.-P., Howard, J. 2014a. Springsnails: a new conservation focus in 

western North America. Bioscience 64:693-700.

Hershler, R., Liu, H.-P., Landye, J.J. 2002. A new species of Eremopyrgus (Hydrobiidae: 

Cochliopinae) from the Chihuahuan Desert, Mexico: phylogenetic 

relationships and biogeography. Journal of Molluscan Studies 68:7-15.

Hershler, R., Liu, H.-P., Landye, J.J. 2011. Two new genera and four new species of 

freshwater cochliopid gastropods (Rissooidea) from northeastern Mexico. 

Journal of Molluscan Studies 77:8-23.

Hershler, R., Liu, H.-P., Mulvey, M. 1999b. Phylogenetic relationships within the 

aquatic snail genus Tryonia: implications for biogeography of the North 

American Southwest. Molecular Phylogenetics and Evolution 13:377-391.

Hershler, R., Liu, H.-P., Sada, D.W. 2007b. Origin and diversification of the 

Soldier Meadow springsnails (Hydrobiidae: Pyrgulopsis), a species flock 

in the northwestern Great Basin, United States. Journal of Molluscan

Studies 73:167-183.

Hershler, R., Liu, H.-P., Simpson, J.S. 2015b. Assembly of a micro-hotspot of 

caenogastropod endemism in the southern Nevada desert, with a description 

of a new species of Tryonia (Truncatelloidea, Cochliopidae). 

ZooKeys 492:107-122.

Hershler, R., Liu, H.-P., Stevens, L.E. 2016b. A new springsnail (Hydrobiidae: 

Pyrgulopsis) from the lower Colorado River basin, northwestern Arizona. 

Western North American Naturalist 76:72-81.

Hershler, R., Liu, H.-P., Thompson, F.G. 2003b. Phylogenetic relationships of North 

American nymphophiline gastropods based on mitochondrial DNA sequences. 

Zoologica Scripta 32:357-366.

Hershler, R., Mulvey, M., Liu, H.-P. 1999a. Biogeography in the Death Valley region: 

evidence from springsnails (Hydrobiidae: Tryonia). Zoological Journal of the 

Linnean Society 126:335-354.

Literature Cited
Technical Note 449 79

Hershler, R., Mulvey, M., Liu, H.-P. 2005. Genetic variation in the desert springsnail 

(Tryonia porrecta): implications for reproductive mode and dispersal. 

Molecular Ecology 14:1755-1765.

Hershler, R., Ponder, W.F. 1998. A review of morphological characters of hydrobioid 

snails. Smithsonian Contributions to Zoology 600:1-55. Available from 

https://repository.si.edu/handle/10088/5530 (accessed 4/17/2017).

Hershler, R., Pratt, W.L. 1990. A new Pyrgulopsis (Gastropoda: Hydrobiidae) from 

southeastern California, with a model for historical development of the 

Death Valley hydrographic system. Proceedings of the Biological Society of 

Washington 103:279-299.

Hershler, R., Ratcliffe, V., Liu, H.-P., Lang, B., Hay, C. 2014b. Taxonomic revision 

of the Pyrgulopsis gilae (Caenogastropoda: Hydrobiidae) species complex, 

with descriptions of two new species from the Gila River basin, New Mexico. 

ZooKeys 429:69-85.

Hershler, R., Sada, D.W. 1987. Springsnails (Gastropoda: Hydrobiidae) of Ash 

Meadows, Amargosa basin, California-Nevada. Proceedings of the Biological 

Society of Washington 100:776-843.

Hershler, R., Sada, D.W. 2000. A new species of hydrobiid snail of the genus 

Pyrgulopsis from northwestern Nevada. Veliger 43:367-375.

Hershler, R., Thompson, F.G. 1987. North American Hydrobiidae (Gastropoda: 

Rissoacea): redescription and systematic relationships of Tryonia Stimpson, 

1865 and Pyrgulopsis Call and Pilsbry, 1886. Nautilus 101:25-32.

Hershler, R., Thompson, F.G. 1992. A review of the aquatic gastropod subfamily 

Cochliopinae (Prosobranchia: Hydrobiidae). Malacological Review 

Supplement 5:1-140.

Hinds, R.B. 1842. Descriptions of new shells. Annals and Magazine of 

Natural History 10:81-84.

Hurt, C.R. 2004. Genetic divergence, population structure and historical demography 

of rare springsnails (Pyrgulopsis) in the lower Colorado River basin. Molecular 

Ecology 13:1173-1187.

Literature Cited
80 Technical Note 449

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.

Kabat, A.R., Hershler, R. 1993. The prosobranch snail family Hydrobiidae 

(Gastropoda: Rissooidea): review of classification and supraspecific taxa. 

Smithsonian Contributions to Zoology 547:1-94.

Lea, I. 1838. Description of new freshwater and land shells. Transactions of the 

American Philosophical Society 6(1):1-154.

Lea, I. 1844. On fresh water and land shells. Proceedings of the American 

Philosophical Society 2:30-35.

Liu, H.-P., Hershler, R. 2005. Molecular systematics and radiation of western 

North American nymphophiline gastropods. Molecular Phylogenetics 

and Evolution 34:284-298.

Liu, H.-P., Hershler, R. 2007. A test of the vicariance hypothesis of western North 

American freshwater biogeography. Journal of Biogeography 34:534-548.

Liu, H.-P., Hershler, R. 2012. Phylogeography of an endangered western North 

American springsnail. Conservation Genetics 13:299-305.

Liu, H.-P., Hershler, R. 2014. Microsatellite primers for a western North American 

springsnail (Pyrgulopsis robusta): evidence for polyploidy and crossamplification in P. bruneauensis. Journal of Molluscan Studies 80:107-110.

Liu, H.-P., Hershler, R., Clift, K. 2003. Mitochondrial DNA sequences reveal 

extensive cryptic diversity within a western American springsnail. 

Molecular Ecology 12:2771-2782.

Liu, H.-P., Hershler, R., Lang, B., Davies, J. 2013b. Molecular evidence for cryptic 

species in a narrowly endemic western North American springsnail 

(Pyrgulopsis gilae). Conservation Genetics 14:917-923.

Liu, H.-P., Hershler, R., Rossel, C.S. 2015a. Taxonomic status of the Columbia 

duskysnail (Truncatelloidea, Amnicolidae, Colligyrus). ZooKeys 514:1-13.

Liu, H.-P., Hovingh, P., Hershler, R. 2015b. Genetic evidence for recent spread of 

springsnails (Hydrobiidae: Pyrgulopsis) across the Wasatch Divide. Western 

North American Naturalist 75:325-331.

Literature Cited
Technical Note 449 81

Liu, H.-P., Marceau, D., Hershler, R. 2016. Taxonomic identity of two amnicolid 

gastropods of conservation concern in lakes of the Pacific Northwest of the 

USA. Journal of Molluscan Studies 82:464-471.

Liu, H.-P., Walsh, J., Hershler, R. 2013a. Taxonomic clarification and phylogeography 

of Fluminicola coloradensis Morrison, a widely ranging western North 

American pebblesnail. Monographs of the Western North American Naturalist 

6:87-110.

Lysne, S.J., Riley, L.A., Clark, W.H. 2007. The life history, ecology, and distribution 

of the Jackson Lake springsnail (Pyrgulopsis robusta Walker 1908). Journal of 

Freshwater Ecology 22:647-653.

McAlpine, D.F., Bateman, D., Davis, C.A. 2005. Spurwinkia salsa (Pilsbry 1905) 

(Gastropoda: Hydrobiid) in the Kennebecasis Estuary, New Brunswick: a 

brackish-water snail new to Canada. Journal of Conchology 38:602-604.

Martinez, M.A., Myers, T.L. 2008. Associations between aquatic habitat variables and 

Pyrgulopsis trivialis presence/absence. Journal of Freshwater Ecology 

23: 189-194.

Mighels, J.W. 1848. Description of shells from the Sandwich Islands, and other 

localities. Proceedings of the Boston Society of Natural History 2:18-25.

Morrison, J.P.E. 1940. A new species of Fluminicola with notes on “Colorado Desert” 

shells, and on the genus Clappia. Nautilus 53:124-127.

Morrison, J.P.E. 1945. Durangonella, a new hydrobiine genus from Mexico, with three 

new species. Nautilus 59:18-23; pl. 3.

Pilsbry, H.A. 1890. Notices of new Amnicolidae. Nautilus 4:63-64.

Pilsbry, H.A. 1892. Preliminary notices of new forms of fresh water mollusks. 

Nautilus 5:142-143.

Pilsbry, H.A. 1899. Catalogue of the Amnicolidae of the western United States. 

Nautilus 12:121-127.

Pilsbry, H.A. 1905. A new brackish-water snail from New England. 

Nautilus 19:90-91, pl. III: fig. 10.

Pilsbry, H.A. 1907. A new species of Fluminicola. Nautilus 21:75-76.

Literature Cited
82 Technical Note 449

Pilsbry, H.A. 1916. New species of Amnicola from New Mexico and Utah. 

Nautilus 29:111-112.

Pilsbry, H.A. 1933. Amnicolidae from Wyoming and Oregon. 

Nautilus 47:9-12, pl. 2: figs. 1-10.

Pilsbry, H.A. 1934. Pliocene fresh-water fossils of the Kettleman Hills and neighboring 

Californian oil fields. Nautilus 48:15-17.

Pilsbry, H.A. 1935. Western and southwestern Amnicolidae and a new Humboldtiana. 

Nautilus 48:91-94.

Pilsbry, H.A. 1952. Littoridina tenuipes (Couper). Nautilus 66:50-54.

Ponder, W.F. 1988. Potamopyrgus antipodarum—a molluscan colonizer of Europe and 

Australia. Journal of Molluscan Studies 54:271-285.

Say, T. 1817. Descriptions of new species of land and fresh water shells of the United 

States. Journal of the Academy of Natural Sciences of Philadelphia 1:123-126.

Say, T. 1821. Descriptions of univalve shells of the United States. Journal of the 

Academy of Natural Sciences of Philadelphia 2:149-179.

Stearns, R.E.C. 1883. Description of a new hydrobiinoid gasteropod [sic] from the 

mountain lakes of the Sierra Nevada, with remarks on allied species and the 

physiographical features of said region. Proceedings of the Academy of Natural 

Sciences of Philadelphia 35:171-176.

Stearns, R.E.C. 1893. Report on the land and fresh-water shells collected in California 

and Nevada by the Death Valley Expedition, including a few additional species 

obtained by Dr. C. Hart Merriam and assistants in parts of the southwestern 

United States. North American Fauna 7:269-283.

Stimpson, W. 1865. Diagnoses of newly discovered genera of gasteropods [sic], 

belonging to the sub-fam. Hydrobiinae, of the family Rissoidae. American 

Journal of Conchology 1:52-54.

Taylor, D.W. 1950. Three new Pyrgulopsis from the Colorado Desert, California. 

Leaflets in Malacology 1:27-33.

Taylor, D.W. 1966a. A remarkable snail fauna from Coahuila, México. 

Veliger 9:152-228.

Literature Cited
Technical Note 449 83

Taylor, D.W. 1966b. Summary of North American Blancan nonmarine mollusks. 

Malacologia 4:1-172.

Taylor, D.W. 1975. Index and bibliography of late Cenozoic freshwater Mollusca 

of western North America. University of Michigan Papers on 

Paleontology 10:1-384.

Taylor, D.W. 1981. Freshwater mollusks of California: a distributional checklist. 

California Fish and Game 67:140-163.

Taylor, D.W. 1987. Fresh-water molluscs from New Mexico and vicinity. Bulletin of the 

New Mexico Bureau of Mines and Mineral Resources 116:1-50.

Taylor, D.W., Bright, R.C. 1987. Drainage history of the Bonneville basin. In: Kopp, 

R.S., Cohenour, R.E., eds. Cenozoic geology of western Utah; sites for 

precious metal and hydrocarbon accumulations. Utah Geological Association 

Publication 16:239-256.

Thiele, J. 1925. Mollusca = Weichtiere. In: Kukenthal, W., Krumbach, T., eds. 

Handbuch der Zoologie, eine Naturgeschichte der Stämme des Tierreiches. 

Volume 5. Berlin and Leipzig: Walter de Gruyter and Co.:15-96, figures 8-109.

Thompson, F.G. 1968. The aquatic snails of the family Hydrobiidae of peninsular 

Florida. Gainesville: University of Florida Press. 268 p.

Thompson, F.G., Hershler, R. 1991. Two new hydrobiid snails (Amnicolinae) from 

Florida and Georgia, with a discussion of the biogeography of freshwater 

gastropods of south Georgia streams. Malacological Review 24:55-72.

Thompson, F.G., Hershler, R. 2002. Two genera of North American freshwater snails: 

Marstonia Baker, 1926, resurrected to generic status, and Floridobia, new genus 

(Prosobranchia: Hydrobiidae: Nymphophilinae). Veliger 45:269-271.

Troschel, F. H. 1857. Das Gebiss der Schnecken zur Begrundung Einer 

Naturlichen Classification. Erster Band, Part 1. Berlin: Nicolaische 

Verlagsbuchhandlung:73-112, plates 5-8.

Tryon, G.W. 1863. Notes on American fresh water shells, with descriptions of two 

new species. Proceedings of the Academy of the Natural Sciences of 

Philadelphia 14:451-452.

Tryon, G.W. 1865. Descriptions of new species of Amnicola, Pomatiopsis, Somatogyrus, 

Gabbia, Hydrobia, and Rissoa. American Journal of Conchology 1:219-222.

Literature Cited
84 Technical Note 449

Tryon, G.W. 1866. [Review of “Researches upon the Hydrobiinae and allied forms by 

Dr. Wm. Stimpson.”] American Journal of Conchology 2:152-158.

Tsai, Y.J., Maloney, K., Arnold, A.E. 2007. Biotic and abiotic factors influencing the 

distribution of the Huachuca springsnail (Pyrgulopsis thompsoni). Journal of 

Freshwater Ecology 22:213-218.

Turgeon, D.D., Quinn, J.F., Bogan, A.E., Coan, E.V., Hochberg, F.G., Lyons, W.G., 

Mikkelsen, P.M., Neves, R.J., Roper, C.F.E., Rosenberg, G., Roth, B., Scheltema, 

A., Thompson, F.G., Vecchione, M., Williams, J.D. 1998. Common and 

scientific names of aquatic invertebrates from the United States and Canada: 

Mollusks. Second edition. American Fisheries Society Special 

Publication 26. 526 p.

USGS (United States Geological Survey). 2016. Nonindigenous aquatic species 

database, Gainesville, FL. Available from http://nas.er.usgs.gov (accessed 

2/19/2016).

Varela-Romero, A., Myers, T.L., Sorensen, J., Abarca, F. 2013. Taxonomic status and 

phylogeny of the San Bernardino spring snail populations into the genus 

Pyrgulopsis in Sonora and Arizona. Biotecnia 15:45-50.

Vázquez, A.A., Cobian, D., Sánchez, J., Pointier, J.-P. 2012. First record of Littoridinops 

monroensis (Frauenfeld, 1863) (Gastropoda: Cochliopidae) in Cuba through a 

likely natural dispersal event. Molluscan Research 32:50-54.

Walker, B. 1908. Pomatiopsis robusta n. sp. Nautilus 21 [incorrectly listed as 22 on the 

first page]:97.

Walker, B. 1916. The Mollusca collected in northeastern Nevada by the WalkerNewcomb Expedition of the University of Michigan. Occasional Papers of the 

Museum of Zoology, University of Michigan 29:1-8.

Wilke, T., Davis, G.M., Falniowski, A., Giusti, F., Bodon, M., Szarowska, M. 2001. 

Molecular systematics of Hydrobiidae (Mollusca: Gastropoda: Rissooidea): 

testing monophyly and phylogenetic relationships. Proceedings of the 

Academy of Natural Sciences of Philadelphia 151:1-21.

Wilke, T., Haase, M., Hershler, R., Liu, H.-P., Misof, B., Ponder, W. 2013. Pushing 

short DNA fragments to the limit: phylogenetic relationships of ‘hydrobioid’ 

gastropods (Caenogastropoda: Rissooidea). Molecular Phylogenetics and 

Evolution 66:715-736.

Literature Cited
Technical Note 449 85

Appendix

Compilation of COI sequences. Amnicola: Liu et al. 2016; Colligyrus: Hershler 

et al. 2003a, Liu et al. 2015a; Lyogyrus: Liu et al. 2016; Eremopyrgus: Hershler et al. 

1999; Ipnobius: Hershler et al. 1999; Littoridinops: Hershler et al. 2007a; Spurwinkia: 

Hershler et al. 2015a; Tryonia: Hershler et al. 1999a, Hershler et al. 1999b, Hershler 

et al. 2015a, Hershler et al. 2015b; Pyrgulopsis: Hershler et al. 2003a, Hershler et al. 

2003b, Liu et al. 2003, Hershler and Liu 2004a, Hershler and Liu 2004b, Hurt 2004, 

Liu and Hershler 2005, Hershler et al. 2007b, Liu and Hershler 2007, Hershler and Liu 

2008, Hershler et al. 2008, Hershler and Liu 2009, Hershler and Liu 2010, Hershler 

and Liu 2012a, Liu and Hershler 2012, Hershler et al. 2013, Liu et al. 2013b, Hershler 

et al. 2014b, Liu et al. 2015b, Hershler et al. 2016a, Hershler et al. 2016b; Fluminicola: 

Hershler et al. 2007c, Hershler and Liu 2012b, Liu et al. 2013a; Pristinicola: Hershler 

et al. 2003b; Taylorconcha: Hershler et al. 2006.

Appendix
86 Technical Note 449

AMNICOLIDAESpecies

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Amnicola

Amnicola 

limosa AM5, AM11 KU991117, KU991123 6 0% 0.5%with AM15

Pattison Lake, east 

side at Washington 

Department of Fish 

and Wildlife boat 

ramp, Thurston 

Cso., WA

AM6 KU991118, 

KU991119 4 0-0.20% 0.5%with AM15

Bonaparte Lake, 

south end at 

campground, 

Okanogan Co., WA

AM7 KU991120 4 0% 0.5%

with AM15

Curlew Lake at 

state park west 

of Hwy 21, Ferry 

Co., WA

AM8 KU991121 4 0% 0.5%

with AM15

Spectacle Lake, 

south side, near 

resort, Okanogan 

Co., WA

AM9 KU991122 4 0% 0.5%

with AM15

Leader Lake, north 

side near dam, 

Okanogan Co., WA

AM12 KU991124 4 0-0.20% 0.6%

with AM15

McWenneger 

Slough, south 

side near Hwy 35, 

Flathead Co., MT

AM15 KU991126, 

KU991127 4 0-0.50% 0.6%with AM12

Browns Lake, 

cutoff pond along 

northeast side, 

Powell Co., MT 

AM16 KU991128, 

KU991129 4 0-0.30% 0.5% with AM12, AM15

Georgetown 

Lake, east side 

north of Stuart 

Campground, 

Granite Co., MT 

Colligyrus

Colligyrus 

convexus BL AY196166 1 0% 0.3% with MB

Colligyrus sp. 

KL, differ by 

3.9%

Baum Lake, Pit 

River basin, Shasta 

Co., CA

TS AY196167 1 0% 0.5% with MB

Colligyrus sp. 

KL, differ by 

4.0%

Fall River (spring 

source), Pit River 

basin, Shasta Co., 

CA

MB AY196168, 

AY196169 2 0% 0.5% with TS

Colligyrus sp. 

KL, differ by 

3.8%

Burney Creek, Pit 

River basin, Shasta 

Co., CA

Colligyrus 

depressus SRa AY196170 1 0% 0% with SRb

C. convexus, 

differ by 

7.8-8.2%, 

Colligyrus sp. 

KL, differ by 

7.8%

Second spring 

south of Turner 

Ranch, Silvies 

River basin, Harney 

Co., OR

SRb AY196171 1 0% 0% with SRa

C. convexus, 

differ by 

7.8-8.2%, 

Colligyrus sp. 

KL, differ by 

7.8%

Third spring south 

of Turner Ranch, 

Silvies River basin, 

Harney Co., OR
Technical Note 449 87

AMNICOLIDAE

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Colligyrus 

greggi COL1 KT248021 4 0% 2.2% with SN C. depressus, differ by 9.3%

Small spring, 

Brooks Meadow, 

middle Columbia 

River basin, Hood 

River Co., OR

COL2 KT248022 4 0% 2.5% with SN C. depressus, 

differ by 9.3%

Spring tributary, 

Tony Creek, middle 

Columbia River 

basin, Hood River 

Co., OR

COL3 KT248023 4 0% 2.2% with SN C. depressus, 

differ by 9.3%

Bottle Prairie, 

middle Columbia 

River basin, Hood 

River Co., OR

COL4 KT248024 4 0% 2.2% with SN C. depressus, 

differ by 9.3%

Spring tributary, 

Ramsey Creek, 

middle Columbia 

River basin, Wasco 

Co., OR

COL5

KT248025 

(N=3), 

KT248026 

(N=1)

4 0-0.30% 2.2% with SN

C. depressus, 

differ by

9.2-9.3%

Spring tributary, 

Clear Creek, 

Deschutes River 

basin, Wasco Co., 

OR

COL6 KT248027 

(N=3), 4 0-0.20% 2.7% with SN

C. depressus, 

differ by

9.3-9.5%

Bear Creek, Hood 

River Co., OR

CL AY196174 1 0% 2.3% with SN C. depressus, 

differ by 9.2%

Oak Grove Fork 

[Clackamas River], 

Willamette River 

basin, Clackamas 

Co., OR

SN AY196172 1 0% 2.7% with 

COL6

C. depressus, 

differ by 9.8%

Springs along 

Cliff Creek, upper 

Snake River basin, 

Sublette Co., WY

BR AY196173 1 0% 2.3% with 

COL6

C. depressus, 

differ by 10.2%

Spring at 

Saint Charles 

Campground, Bear 

Lake basin, Bear 

Lake Co., ID

AM17 KT248030 2 0% 2.3% with 

COL6

C. depressus, 

differ by 10.3%

Spring at Porcupine 

Campground, Bear 

Lake basin, Bear 

Lake Co, ID

AM20 KT248031 2 0% 2.3% with 

COL6

C. depressus, 

differ by 10.1%

Springs along 

Trail Creek, upper 

Snake River basin, 

Caribou Co., ID

Colligyrus sp. KL AY196175 1 0%

C. convexus, 

differ by

3.8-4.0%

Link River, Klamath 

River basin, 

Klamath Co., OR

Colligyrus sp. COL7 KT248029 1 0%

C. greggi, differ 

by 10.3-11.2%, 

Colligyrus sp. 

KL, differ by 

10.6%

Allison Springs, 

Puget Sound 

drainage, Thurston 

Co., WA
88 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Lyogyrus

Lyogyrus sp. AM10 KU991131 4 0% 0.5%

with AM13

Fish Lake, south 

side at Cove 

Resort, Chelan 

Co., WA

AM22 KU991131 3 0% 0.5%

with AM13

Curlew Lake at 

Camp Curlew Road 

terminus, Ferry 

Co., WA

AM13

KU991132, 

KU991133, 

KU991134 

4 0-0.20% 0.8%

with AM13

Smith Lake, south 

side at public 

access boat ramp, 

Flathead Co., MT

AM14 KU991134, 

KU991136 4 0-0.20% 0.6%with AM13

Upsata Lake, east 

side near outlet, 

Powell Co., MT

Eremopyrgus

Eremopyrgus 

eganensis AF129317 1 0.00%

Spring, southeast 

Steptoe Valley, 

north of Ely, White 

Pine Co., NV

Tryonia sp. 

in GenBank; 

Tryonia n. sp. 

2 in Hershler 

et al. (1999b)

Ipnobius

Ipnobius 

robustus AF061774 1

Travertine Springs, 

Death Valley, Inyo 

Co., CA

Tryonia 

robusta in 

GenBank

Littoridinops

Littoridinops 

monroensis MISC1A EF490565 1 0.00%

Restored tidal 

marsh on north 

side of Waterfront 

Road, Point Edith 

Wildlife Area, 

Contra Costa Co., 

CA

Spurwinkia

Spurwinkia 

salsa KP057916 4 0.00%

Salt marsh, Eden 

Landing, south 

San Francisco Bay, 

Alameda Co., CA

Tryonia

Tryonia 

angulata AF061764 1 0.00% T. variegata SS, differ by 0.5%

Big Spring, Ash 

Meadows, Nye 

Co., NV

Tryonia 

clathrata AF061767 1 0.00%

T. infernalis, 

differ by 

3.4-3.9%

Warm Spring, 

Moapa Valley, Clark 

Co., NV

Tryonia

elata AF61766 1 0.00% T. ericae, differ by 2.3%

Spring tributary to 

Kings Pool, Point 

of Rocks, Ash 

Meadows, Nye 

Co., NV

Tryonia

ericae AF061768 1 0.00% T. elata, differ by 2.3%

Spring north of 

Collins Ranch, Ash 

Meadows, Nye 

Co., NV

AMNICOLIDAE CHOCHLIOPIDAE
Technical Note 449 89

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Tryonia

gilae AF129309 1 0.00% T. angulata, differ by 4.2%

Tom Niece Spring, 

Graham Co., AZ

Tryonia

imitator M, MO AF061770 1 0.00% 0.2%with S/PE

T. salina, differ 

by 1.8%

Moro Cojo Slough, 

Moss Landing, 

Monterey Co., CA

S, PE AF061769 1 0.00% 0.2%

with M/MO

T. salina, differ 

by 1.6%

Los Penasquitos 

Lagoon, San Diego 

Co., CA

Tryonia 

infernalis BP2 KP899916-18 6 0.0-1.1%

T. clathrata, 

differ by

3.4-3.9%

Blue Point Spring, 

just below source, 

Clark Co., NV

Tryonia

margae

AF061771 1 0.00% T. monitorae, 

differ by 1.9%

Grapevine Springs 

(upper warm 

spring), Death 

Valley, Inyo Co., CA

Tryonia 

monitorae AF129316 1 0.00% T. margae,differ by 1.9%

Spring, Potts 

Ranch, Monitor 

Valley, Nye Co., NV

Tryonia sp. 

in GenBank, 

Tryonia n. sp. 

1 in Hershler 

et al. (1999b)

Tryonia

porrecta WH AF061773 (HI) 3 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator,

differ by 

4.9-5.0%

Whitmore Hot 

Springs, Long 

Valley, Mono Co., 

CA

Tryonia protea

in GenBank

HC AY803024 

(HI) 3 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator,

differ by 

4.9-5.0%

Hot Creek, Long 

Valley, Owens 

River basin, Mono 

Co., CA

OA AF061772 

(HI) 1 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator,

differ by 

4.8-4.9%

Oasis Spring, 

Salt Creek, Salton 

Trough, Riverside 

Co., CA

Tryonia protea

in GenBank

HU AY803025 

(HI) 4 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator, 

differ by 

4.9-5.0%

Hunters Spring, 

Salton Sea basin, 

Riverside Co., CA

SS

AY803023 

(HI), 

AY803029 

(HIII), 

AY803030 

(HX), 

AY803031 

(HIX)

6 0.0-2.5%

HI: 2.1% with 

SS (HX) and 

HF (HVIII);

HIII: 2.5% with 

SS (HX), HF 

(HVIII), Misc4A, 

and Misc5A; 

HX: 2.6% with 

HS (HV);

HIX: 2.5% with 

HS (HV)

T. imitator,

differ by 

4.8-5.1%

South Springs, 

Fish Springs Flat, 

Bonneville basin, 

Juab Co., UT

CHOCHLIOPIDAE
90 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Tryonia

porrecta

(continued)

HS

AF129312 

(HIV), 

AY803027 

(HII), 

AY803028 

(HV), 

AY803032 

(HVI)

4 0.8-2.3%

HIV: 2.4% with 

SS (HX), HF 

(HVIII), Misc4A, 

and Misc5A; 

HII: 2.5% with 

SS (HX), HF 

(HVIII), Misc4A, 

and Misc5A; 

HV: 2.6% with 

SS (HX), HF 

(HVIII), Misc4A, 

and Misc5A; 

HVI: 2.3% with 

HS (HV)

T. imitator,

differ by 

4.6-5.1%

Horseshoe Springs, 

Skull Valley, Tooele 

Co., UT

Tryonia protea

in GenBank, 

collection 

locality given 

as Warm 

Spring in 

GenBank

WS AF129313 

(HI) 2 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator,

differ by 

4.9-5.1%

Warm Springs, 

Tooele Valley, 

Tooele Co., UT

Tryonia protea

in GenBank

BL AF129314 

(HI) 2 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator,

differ by 

4.7-4.9%

Spring, Blue Lake, 

Great Salt Lake 

Desert, Tooele 

Co., UT

Tryonia protea

in GenBank

HF

AY803033 

(HVIII), 

AY803034 

(HVII)

5 0.0-0.2% 2.5-2.6% with 

HS (HV)

T. imitator,

differ by 

4.9-5.3%

Spring, Hualapai 

Flat, Lahontan 

basin, Washoe 

Co., NV

ED AY803026 

(HI) 3 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator,

differ by 

4.9-5.0%

Spring at El Doctor, 

Colorado River 

basin, Sonora, 

Mexico

FS AF129311 

(HI) 1 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator,

differ by 

4.9-5.0%

Spring, south end 

of Fish Springs 

National Wildlife 

Refuge, Juab Co., 

UT

Tryonia protea

in GenBank

KP057915 

(HI) 1 0.00%

2.1% with SS 

(HX) and HF 

(HVIII)

T. imitator,

differ by 

4.9-5.0%

Garden Lakes, 

Avondale, Maricopa 

Co., AZ

Misc4A EF490563 1 0.00% 2.6% with HS 

(HV)

T. imitator,

differ by 

4.6-5.0%

Restored tidal 

marsh on north 

side of Waterfront 

Road, Point Edith 

Wildlife Area, 

Contra Costa Co., 

CA

Misc5A EF490564 1 0.00% 2.6% with HS 

(HV)

T. imitator,

differ by 

4.6-5.0%

Restored tidal 

marsh, southeast 

of Point Edith 

Wildlife Area, 

Contra Costa Co., 

CA

Tryonia 

quitobaquitae AF129315 1 0.00% T. angulata, differ by 5.0%

Quitobaquito 

Springs, Pima 

Co., AZ

Tryonia 

rowlandsi AF061775 1 0.00% T. angulata, differ by 3.8%

Grapevine Springs 

(lower warm 

spring), Death 

Valley, Inyo Co., CA

CHOCHLIOPIDAE
Technical Note 449 91

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Tryonia

salina AF061776 1 0.00%

T. imitator,

differ by 

1.6-1.8%

Spring, Cottonball 

Marsh, Death 

Valley, Inyo Co., CA

Tryonia 

variegata DV, DH AF061778 1 0.00% T. ericae, differ by 2.9%

Devils Hole, Ash 

Meadows, Nye 

Co., NV

SA, SS AF061777 1 0.00% T. angulata, 

differ by 0.5%

Saratoga Spring, 

Amargosa River 

drainage, San 

Bernardino Co., CA

Pyrgulopsis

Pyrgulopsis 

amargosae

P. amer 2 AY367479 1 0.00% 7.3-7.4% with P. 

"amargosae" (2)

P. anatina, 

differ by 4.8%, 

P. bacchus, 

differ by 

4.0-5.1%, 

P. perforata, 

differ by 

4.5-4.8%

Saratoga Spring, 

Death Valley, San 

Bernardino Co., CA

P.

“amargosae” 

(1)

DQ364002 1 0.00% 6.9% with P. 

amargosae

P. bifurcata, 

differ by 4.9%, 

P. intermedia, 

differ by

4.8-5.4%, 

P. thompsoni, 

differ by 

4.8-5.9%

Third spring along 

trail (from north), 

Amargosa Canyon, 

Inyo Co., CA

Pyrgulopsis sp. 

1 HPL-2003 in 

GenBank

P.

“amargosae” 

(2)

AF520924, 

KF559193 2 1.30% 7.3-7.4% withP. amargosae

P. intermedia, 

differ by

5.1-5.8%,

P. micrococcus, 

differ by

5.3-5.6%, 

P. milleri, differ

by 5.4-5.7%,

P. thompsoni, 

differ by

5.0-6.3%

Spring, Amargosa 

Canyon, Inyo, Co., 

CA

Pyrgulopsis 

anatina P58 AY627926 1 0.00%

P. bifurcata, 

differ by 4.5%, 

P. torrida, differ 

by 4.5%

Spring southeast of 

Old Collins Spring, 

Railroad Valley, Nye 

Co., NV

Pyrgulopsis 

anguina P94B EU700466 1 0.00%

P. kolobensis, 

differ by

3.2-4.4%, 

P. marcida, 

differ by 3.4%, 

P. nonaria, 

differ by 3.6%, 

P. variegata, 

differ by 3.6%

Big Springs, Snake 

Valley, White Pine 

Co., NV

Pyrgulopsis 

archimedis AH AY197583 1 0.00% 0.2-0.4% with UK/P50

P. rupinicola, 

differ by 

2.7-2.9%,

P. sp. A, differ 

by 2.4-2.9%

Big Lake Springs, 

Pit River basin, 

Shasta Co., CA

BL, D8B AY197579 1 0.00% 0.2-0.4% with 

UK/P50

P. rupinicola, 

differ by

2.7-2.9%, 

P. sp. A, differ 

by 2.4-2.9%

Baum Lake, Pit 

River basin, Shasta 

Co., CA

CHOCHLIOPIDAE HYDROBIIDAE
92 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

archimedis

(continued)

FR_a/D1A, 

D1D

AY197577 

(FR_a/D1A), 

AY426357 

(D1D)

2 0.00% 0.2-0.4% with 

UK/P50

P. rupinicola, 

differ by

2.7-2.9%, 

P. sp. A, differ 

by 2.4-2.9%

Fall River (CalTrout 

Access), Pit River 

basin, Shasta Co., 

CA

AY426357/

D1D listed as 

P. intermedia

in GenBank

FR_b AY197581 1 0.00% 0.2-0.4% with 

UK/P50

P. rupinicola, 

differ by

2.7-2.9%,

P. sp. A, differ 

by 2.4-2.9%

Spring source, 

Fall River, Pit River 

basin, Shasta Co., 

CA

JS AY197578 1 0.00% 0.3-0.6% with 

UK/P50

P. rupinicola, 

differ by

2.9-3.1%,

P. sp. A, differ 

by 2.6-3.1%

Ja She Creek, Pit 

River basin, Shasta 

Co., CA

KL_a/D31A,

D31B

AY197586 

(KL_a/D31A), 

AY426356 

(D31B)

2 0.00% 0.3-0.6% with 

UK/P50

P. rupinicola, 

differ by

2.6-2.7%, 

P. sp. A, differ

by 2.4-2.9%

Fifth Link River 

spring, Klamath 

basin, Klamath 

Co., OR

AY426356/

D31B 

listed as P. 

intermedia in 

GenBank

LO1, LO2

AY197584 

(LO1),

AY197585 

(LO2)

2 0.00% 0.3% with JS, 

KL_a/D31

P. rupinicola, 

differ by 

2.9-3.1%, 

P. sp. A, differ

by 2.6-2.9%

Lost River near 

Horsefly Irrigation 

District, Klamath 

basin, Klamath 

Co., OR

PR/D7A, D7C

AY197580

(PR/D7A),

AY426358 

(D7C)

2 0.00% 0.2-0.4% with 

UK/P50

P. rupinicola, 

differ by

2.7-2.9%,

P. sp. A, differ

by 2.4-2.9%

Pit River at 

confluence with

Hat Creek,

Shasta Co., CA

AY426358/

D7C listed as 

P. intermedia

in GenBank

TH AY197582 1 0.00% 0.2-0.4% with 

UK/P50

P. rupinicola, 

differ by

2.7-2.9%,

P. sp. A, differ

by 2.4-2.9%

Three unnamed 

springs, Pit River 

basin, Shasta Co., 

CA

UK/P50A, 

P50C

AF520950 

(UK/P50A), 

AY426355 

(P50C)

2 0.20%

0.3-0.6%

with JS,

KL_a/D31

P. rupinicola, 

differ by

2.9-3.2%,

P. sp. A, differ

by 2.6-3.6%

Upper Klamath

Lake at Hagelstein 

Park outlet,

Klamath Co., OR

Pyrgulopsis 

arizonae P148 AY627948 1 0.00% 1.5-1.7% with Psan6-9

P. coloradensis, 

differ by 5.0%, 

P. sanchezi,

differ by

4.1-5.2%

Medicine Spring, 

Bylas, Gila River 

basin, Graham Co., 

AZ

Psan1,

Psan2,

Psan3,

Psan4,

Psan5

AY485561 

(Psan1, N=26), 

AY485562 

(Psan2, N=1),

AY485563 

(Psan3, N=1),

AY485564 

(Psan4, N=1),

AY485565 

(Psan5, N=1) 

30 0.2-0.6% 1.1-1.5% with 

Psan6-9

P. coloradensis, 

differ by

5.6-5.9%,

P. sanchezi,

differ by

4.6-5.9%

Cold

Hurt (2004), 

listed as P. 

sancarlosensis

in GenBank

HYDROBIIDAE
Technical Note 449 93

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

arizonae

(continued)

Psan6,

Psan7,

Psan8,

Psan9

AY485566 

(Psan1, N=30),

AY485567 

(Psan2, N=1),

AY485568 

(Psan3, N=1),

AY485569 

(Psan4, N=1)

34 0.2-1.2% 1.5-1.7% with 

P148

P. coloradensis, 

differ by

6.1-6.4%,

P. sanchezi,

differ by

5.2-6.2%

Tom Niece

Hurt (2004), 

listed as P. 

sancarlosensis

in GenBank

Pyrgulopsis 

aurata P144A EU700473 1 0.00%

P. pictilis,

differ by 0.0%, 

P. bryantwalkeri, 

differ by 0.3%,

P. kolobensis, 

differ by

1.7-2.9%,

P. nonaria,

differ by 2.0%,

P. variegata,

differ by 2.0%

Coyote Spring, 

Pleasant Valley, 

Pershing Co., NV

Pyrgulopsis 

avernalis AF520930 1 0.00% P. perforata,differ by 8.6%

Muddy Spring, 

Moapa Valley, Clark 

Co., NV

Pyrgulopsis 

bacchus P191 DQ364005 1 0.00% P. montana,differ by 2.2%

Tassi Spring,Grand 

Wash, Colorado 

River basin,

Mohave Co., AZ

Pbac1,

Pbac2

AY485540 

(N=17), 

AY485541 

(N=13)

30 0.20%

P. montana,

differ by

2.2-2.4%

Grapevine Hurt (2004)

Pbac1,

Pbac6,

Pbac8 

AY485540 

(N=5), 

AY485544 

(N=27), 

AY485545 

(N=1)

33 0.5-0.6%

P. montana,

differ by

2.4-2.6%

Whiskey Hurt (2004)

Pbac2,

Pbac3,

Pbac5

AY485541 

(N=1), 

AY485542 

(N=32), 

AY485543 

(N=1)

34 0.2-0.5%

P. montana,

differ by

2.2-2.6%

Tassi Hurt (2004)

Pyrgulopsis 

bernardina Pber

AY485595 

(N=23, Pber1), 

AY485596 

(N=7, Pber2), 

AY485597 

(N=1, Pber3)

31 0.7-4.9%

P. kolobensis, 

differ by

5.5-7.5%,

P. owyheensis, 

differ by

5.4-7.1% 

P. pilsbryana, 

differ by

4.8-6.1%, 

P. rupinicola,

differ by

5.1-7.0%, 

P. sp. C, differ

by 4.7-6.0%, 

P. sp. D, differ

by 5.2-6.1%,

P. sp. H, differ

by 4.8-6.1%

San Bernardino

Hurt (2004), 

listed as 

P. cochisi, 

P. bernadensis

in GenBank

HYDROBIIDAE
94 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

bifurcata P56 AY627925 1 0.00%

P. pellita,

differ by 3.3%,

P. perforata,

differ by

3.3-3.5%

Springs west of 

Carico Lake, Carico 

Lake basin, Lander 

Co., NV

Pyrgulopsis 

breviloba P63 AY627928 1 0.00%

P. lata, differ

by 0.5%,

P. kolobensis, 

differ by

3.4-4.8%,

P. marcida,

differ by 3.7%,

P. nonaria,

differ by 3.7%

Flag Springs 

(middle), White 

River Valley, Nye 

Co., NV

Pyrgulopsis

bruesi AF520926 1 0.00%

P. dixensis,

differ by 0.3%,

P. conica,

differ by 3.4%,

P. glandulosa, 

differ by

3.3-3.8%,

P. montana,

differ by 3.3%,

P. thompsoni, 

differ by

3.2-4.4%

Fly Geyser, Black 

Rock Desert, 

Washoe Co., NV

Pyrgulopsis 

bruneauensis bruneauensis AF520941 1 0.00% 1.8% with Pbr1

P. kolobensis, 

differ by

5.3-6.1%, 

P. marcida,

differ by 5.3%,

P. pilsbryana, 

differ by 5.3%, 

P. variegata,

differ by 5.3%, 

P. sp. B,

differ by 5.3%,

P. sp. C, differ

by 5.2-5.3%,

P. sp. D,

differ by 5.3%,

P. sp. H,

differ by 5.3%,

P. sp. I,

differ by 5.1%

Bruneau Hot 

Springs, Owyhee 

Co., ID

Pbr1 JN184636-645 10 0.00% 1.8% with 

bruneauensis

P. kolobensis, 

differ by

5.2-6.5%, 

P. marcida, differ 

by 5.3%,

P. pilsbryana, 

differ by 5.7%, 

P. variegata, 

differ by 5.3%, 

P. sp. B, differ

by 5.3%, 

P. sp. C, differ

by 5.6-5.8%,

P. sp. D, differ

by 5.7%,

P. sp. H, differ

by 5.8%,

P. sp. I, differ

by 5.1%

Pool at mouth of 

Hot Creek, Owyhee 

Co., ID

HYDROBIIDAE
Technical Note 449 95

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

bruneauensis

(continued)

Pbr2 JN184646-655 10 0-1.5% 0.3-1.8% with 

bruneauensis

P. kolobensis, 

differ by

4.4-6.5%, 

P. marcida, differ 

by 4.5-5.3%, 

P. pilsbryana, 

differ by

4.7-5.7%, 

P. variegata,

differ by

4.3-5.3%, 

P. sp. B, differ

by 4.5-5.3%, 

P. sp. C, differ

by 4.6-5.8%, 

P. sp. D, differ

by 4.8-5.7%,

P. sp. H, differ

by 4.7-5.8%, 

P. sp. I, differ

by 4.3-5.1%

Spring along 

Bruneau River 

below Hot Creek 

confluence,

Owyhee Co., ID

Pbr3 JN184656-665 10 0-1.4% 0.7-1.8% with 

bruneauensis

P. kolobensis, 

differ by

4.4-6.5%, 

P. marcida, differ 

by 4.5-5.3%,

P. pilsbryana, 

differ by

4.7-5.7%, 

P. variegata,

differ by

4.3-5.3%,

P. sp. B, differ

by 4.5-5.3%,

P. sp. C, differ

by 4.6-5.8%,

P. sp. D, differ

by 4.8-5.7%,

P. sp. H, differ

by 4.7-5.8%,

P. sp. I, differ

by 4.3-5.1%

Spring along 

Bruneau River 

above Hot Creek 

confluence,

Owyhee Co., ID

Pbr4 JN184666-672 7 0-1.4% 0.7-1.8% with 

bruneauensis

P. kolobensis, 

differ by

4.4-6.5%, 

P. marcida, differ 

by 4.5-5.3%, 

P. pilsbryana, 

differ by

4.7-5.7%, 

P. variegata,

differ by

4.3-5.3%, 

P. sp. B, differ

by 4.5-5.3%, 

P. sp. C, differ

by 4.6-5.8%, 

P. sp. D, differ

by 4.8-5.7%, 

P. sp. H, differ

by 4.7-5.8%, 

P. sp. I, differ

by 4.3-5.1%

Hot Creek

ca. 100 m upflow 

from mouth, 

Owyhee Co., ID

HYDROBIIDAE
96 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

bruneauensis

(continued)

Pbr5 JN184673-679 7 0-1.5% 0.7-2.0% with 

bruneauensis

P. kolobensis, 

differ by

4.7-6.7%, 

P. marcida, differ 

by 4.8-5.5%, 

P. pilsbryana, 

differ by

5.0-5.9%, 

P. variegata,

differ by

4.7-5.4%,

P. sp. B, differ

by 4.8-5.5%, 

P. sp. C, differ

by 4.9-6.0%, 

P. sp. D, differ

by 5.2-5.8%,

P. sp. H, differ

by 5.0-5.9%, 

P. sp. I, differ

by 4.6-5.3%

Spring along 

Bruneau River 

below Hot Creek 

confluence,

Owyhee Co., ID

Pbr6 JN184680-686 7 0-1.4% 0.7-1.8% with 

bruneauensis

P. kolobensis, 

differ by

4.4-6.5%, 

P. marcida, differ 

by 4.5-5.3%,

P. pilsbryana, 

differ by

4.7-5.7%,

P. variegata,

differ by

4.3-5.3%,

P. sp. B, differ

by 4.5-5.3%,

P. sp. C, differ

by 4.6-5.8%,

P. sp. D, differ

by 4.8-5.7%,

P. sp. H, differ

by 4.7-5.8%, 

P. sp. I, differ

by 4.3-5.1%

Bruneau River 

below Hot Creek 

confluence,

Owyhee Co., ID

Pyrgulopsis 

bryantwalkeri P121A AY627942 1 0.00%

P. aurata, differ

by 0.3%, 

P. pictilis, differ

by 0.3%, 

P. kolobensis, 

differ by

1.7-2.7%,

P. nonaria, differ 

by 2.0%,

P. variegata,

differ by 2.0%, 

P. sp. B, differ

by 1.9%

Warm Spring, 

Humboldt River 

drainage, Elko Co., 

NV

Pyrgulopsis 

californiensis P30 DQ364019 3 0.00% 3.4% with P31

P. kolobensis, 

differ by

4.1-5.3%, 

P. variegata,

differ by 4.3%,

Spring tributary to 

Lytle Creek, Santa 

Ana River basin,

San Bernardino 

Co., CA

P31 AY367481 1 0.00% 3.4% with P30

P. chamberlini, 

differ by 5.1%, 

P. leporina, differ 

by 5.1%

Spring tributary 

to Snow Creek, 

Whitewater River 

basin, Riverside 

Co., CA

HYDROBIIDAE
Technical Note 449 97

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

californiensis

(continued)

P33 AY627924 2 0.00% 7.0% with P30 P. longinqua,

differ by 2.4%

Spring tributary 

to Campo Creek, 

Southern California 

coastal drainage, 

San Diego Co., CA

Pyrgulopsis 

carinifera P9 AY627920 1 0.00%

P. archimedis, 

differ by

6.3-7.5%, 

P. bruneauensis, 

differ by

6.9-7.5%,

P. lentiglans,

differ by 6.9%, 

P. thermalis,

differ by

6.0-12.1%

Muddy Spring, 

Moapa Valley,

Clark Co., NV

Pyrgulopsis 

castaicensis GQ275097 3 0.00% P. hualapaiensis, differ by 3.3%

Middle Canyon 

Spring, Southern 

California coastal 

drainage, Los 

Angeles Co., CA

Pyrgulopsis

sp. 1 HPL2009 in 

GenBank

Pyrgulopsis 

chamberlini P101A EU700468 1 0.00%

P. variegata,

differ by 1.9%, 

P. sp. B, differ

by 1.8%, 

P. sp. I, differ

by 1.9%

Spring at

Glenwood, Sevier 

River drainage, 

Sevier Co., UT

Prygulopsis 

cinerana AC AY197602 1 0.00% P. falciglans,differ by 4.8%

Ash Creek, 

Pit River basin, 

Lassen Co., CA

Pyrgulopsis 

coloradensis BP1A KP899919 1 0.00%

P. sanchezi,

differ by

3.8-5.0%

Blue Point Spring, 

Clark Co., NV

Pyrgulopsis 

conica DS AY627958 1 0.00% 0.0% with Pcon1

P. glandulosa, 

differ by

2.6-3.0%, 

P. perforata,

differ by

2.6-2.7%

Dripping Spring, 

Sacramento Wash, 

Mohave Co., AZ

Pcon1 AY485546 30 0.00% 0.0% with DS

P. glandulosa, 

differ by

2.6-3.0%, 

P. perforata,

differ by

2.6-2.7%

Dripping Hurt (2004)

Pyrgulopsis 

cruciglans P70A AY627931 1 0.00%

P. kolobensis, 

differ by

3.0-4.2%,

P. marcida, 

differ by 3.3%, 

P. variegata, 

differ by 3.3%

Flat Spring, 

Steptoe Valley, 

White Pine Co., NV

HYDROBIIDAE
98 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

crystalis AY367482 1 0.00%

P. pisteri, differ

by 0.6%, 

P. erythropoma, 

differ by 4.5%

Crystal Pool, Ash 

Meadows, Nye Co., 

NV

Pyrgulopsis 

cybele P237 JN255353-58 6 0.0-0.5% 0.0-0.5% with P252

P. glandulosa, 

differ by

2.8-3.2%, 

P. intermedia, 

differ by 2.9-

3.6%

Spring brook 

entering north 

side of South Fork 

Owyhee River, Elko 

Co., NV

Pyrgulopsis

sp. HPL2011a 

isolate P237A, 

P237B, P237C, 

P237D, P237E, 

P237F in 

GenBank

P252 JN255350-52 3 0.0-0.2% 0.0-0.5% with 

P237

P. glandulosa, 

differ by

2.8-3.0%,

P. intermedia, 

differ by

2.9-3.5%

Spring brook 

entering north 

side of South Fork 

Owyhee River, Elko 

Co., NV

Pyrgulopsis

sp. HPL2011a 

isolate P252A, 

P252B, P252C 

in GenBank

Pyrgulopsis 

deaconi AY367483 1 0.00%

P. eremica, differ 

by 6.6-10%, 

P. kolobensis, 

differ by

6.8-8.6%,

P. variegata,

differ by 6.8%,

P. sp. A, differ

by 6.3-7.3%

Red Spring, Las 

Vegas Valley, Clark 

Co., NV

Pyrgulopsis 

deserta P115 DQ251077 1 0.00%

0.8-4.4% with 

Pdes1/2, 

1.1-3.6% with 

Pdes6/7

P. kolobensis, 

differ by

5.9-7.0%,

P. pilsbryana, 

differ by

6.2-6.4%, 

P. sp. D, differ

by 6.2%

Springs tributary 

to Virgin River, 

Littlefield, Mohave 

Co., AZ

Pdes1,

Pdes2

AY485534 

(N=15),

AY485535 

(N=13)

28 3.60%

0.8-4.4%

with P115,

0.6-4.4%

with Pdes4/5,

0.6-4.1%

with Pdes5,

0.8-5.7% with 

Pdes6/7

P. kolobensis, 

differ by

5.5-7.8%,

P. pilsbryana, 

differ by

5.7-7.0%,

P. sp. C, differ

by 5.9-6.9%,

P. sp. D, differ

by 5.7-7.0%,

P. sp. H, differ

by 5.9-6.9%

Green Hurt (2004)

Pdes4, 

Pdes5

AY485536 

(N=6),

AY485537 

(N=25)

31 0.20%

0.8-4.4% with 

Pdes1/2, 

1.0-4.0% with 

Pdes6/7

P. kolobensis, 

differ by

5.6-7.0%,

P. pilsbryana, 

differ by

5.8-6.5%, 

P. sp. D, differ

by 5.8-6.1%

Littlefield I Hurt (2004)

Pdes5 AY485537 33 0.00%

0.6-4.1% with 

Pdes1/2, 

1.0-3.9% with 

Pdes6/7

P. kolobensis, 

differ by

5.6-6.6%,

P. pilsbryana, 

differ by

5.8-6.0%, 

P. sp. D, differ

by 5.8%

Littlefield II

Hurt (2004). 

In Hurt (2004) 

table 3 listed 

haplotypes 

Pdes5 and 

Pdes6 are 

both at 100% 

frequency.

HYDROBIIDAE
Technical Note 449 99

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

deserta

(continued)

Pdes6, 

Pdes7

AY485536 

(N=17), 

AY485537 

(N=14)

31 2.70%

1.1-3.6%

with P115,

0.8-5.7%

with Pdes1/2, 

1.0-4.0%

with Pdes4/5, 

1.0-3.9%

with Pdes5

P. kolobensis, 

differ by

6.0-7.7%,

P. pilsbryana, 

differ by

6.3-6.6%,

P. sp. C, differ

by 6.4-6.7%, 

P. sp. D, differ

by 6.3-6.5%,

P. sp. H, differ

by 6.4-6.6%

Red Hill Hurt (2004)

Pyrgulopsis 

diablensis P25 AY627922 2 0.00% P. giulianii,differ by 1.1%

Stream, Del Puerto 

Canyon, San 

Joaquin River basin, 

Stanislaus Co., CA

Pyrgulopsis 

dixensis P143 AY627946 1 0.00%

P. bruesi,

differ by 0.3%,

P. montana,

differ by 3.1%

Spring west of Cain 

Mountain, Dixie 

Valley, Pershing 

Co., NV

Pyrgulopsis 

eremica P145 AY627947 1 0.00%

7.1% with EL_a, 

7.0-7.1% with 

EL_b

P. lasseni, differ 

by 0.3%, 

P. kolobensis, 

differ by

4.9-6.1%,

P. pilsbryana, 

differ by

5.1-5.4%,

P. sp. D, differ

by 5.1%

Spring, tributary to 

Willow Creek, Eagle 

Lake basin, Lassen 

Co., CA

This

sequence 

does not 

appear to 

be that of P. 

eremica (per 

Hershler et 

al. 2003a) 

and was 

apparently 

submitted to 

GenBank in 

error

HC AY197597 1 0.00% 7% with P145

P. kolobensis, 

differ by

3.7-5.3%,

P. sp. D, differ

by 3.7%

Spring west of 

Russell Dairy

Spring, Pit River 

basin, Lassen Co., 

CA

EL_a AY197598 1 0.00% 7.1% with P145

P. kolobensis, 

differ by

3.9-5.5%, 

P. sp. D, differ

by 3.9%

Troxel Point Spring, 

Honey-Eagle Lakes 

basin, Lassen Co., 

CA

EL_b AY197599-600 2 0.20% 7.0-7.1% with 

P145

P. kolobensis, 

differ by

3.9-5.5%, 

P. sp. D, differ

by 3.9-4.1%

Murrers Upper 

Meadow, HoneyEagle Lakes basin, 

Lassen Co., CA

Pyrgulopsis 

erythropoma AY367484 4 0.00%

P. crystalis,

differ by 4.5%, 

P. pisteri, differ

by 4.5%

Kings Pool 

(outflow), Point 

of Rocks, Ash 

Meadows, Nye Co., 

NV

Pyrgulopsis 

fairbanksensis P19 AY627921 1 0.00%

P. isolata, differ

by 0.5%, 

P. nanus, differ

by 1.3%

Fairbanks Spring, 

Ash Meadows, Nye 

Co., NV

HYDROBIIDAE
100 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

falciglans SC AY197601 1 P. cinerana,differ by 4.8%

Smokey Charley 

Spring, Modoc Co., 

CA

Pyrgulopsis 

fausta AY367485 1 0.00%

P. longiglans, 

differ by 6.3%, 

P. trivialis, differ 

by 6.3-6.4%

Corn Creek Springs, 

Las Vegas Valley, 

Clark Co., NV

Pyrgulopsis

fresti IP68

FJ172470 

(IP68A), 

FJ172471 

(IP68B), 

FJ172472 

(IP68C)

3 0.2-0.3% 2.1-2.3%

with IP69B

P. kolobensis, 

differ by

6.2-7.6%,

P. marcida, differ 

by 6.5-6.7%,

P. plicata, differ

by 6.5-6.7%,

P. serrata, differ 

by 6.4-6.6%,

P. variegata,

differ by

6.5-6.7%,

P. sp. C, differ

by 6.5-6.9%

Tudor Warm

Springs, east

side of Owyhee 

River, fourth spring 

from north,

Malheur Co., OR

Pyrgulopsis

sp. 2

HPL2008 

isolate IP68A, 

IP68B, IP68C 

in GenBank

IP69B FJ172473 1 0.00% 2.5%

with P207A

P. kolobensis, 

differ by

6.3-7.3%, 

P. marcida,

differ by 6.4%,

P. plicata,

differ by 6.4%,

P. rupinicola,

differ by

6.4-6.6%,

P. variegata,

differ by 6.4%

Tudor Warm

Springs, west side 

of Owyhee River, 

first spring north 

of mouth of Warm 

Springs Canyon, 

Malheur Co., OR

Pyrgulopsis

sp. 2

HPL2008 

isolate IP69B 

in GenBank

P176B FJ172474 1 0.00% 2.1%

with IP69B

P. kolobensis, 

differ by

6.2-7.3%, 

P. neritella,

differ by 6.3%

Tudor Warm

Springs, east side 

of Owyhee River, 

second spring from 

north, Malheur Co., 

OR

Pyrgulopsis

sp. 2

HPL2008 

isolate

IP176B in 

GenBank

P207A FJ172476 2 0.00% 2.5%

with IP69B

P. kolobensis, 

differ by

5.9-7.4%,

P. neritella,

differ by 6.1%

Spring tributary to 

Owyhee River

upflow from Tudor 

Warm Springs, 

Malheur Co., OR

Pyrgulopsis

sp. 2

HPL2008 

isolate

IP207A in 

GenBank

P208

FJ172477 

(P208A), 

FJ172478 

(P208E)

5 0.80% 1.6-2.1%

with IP69B

P. kolobensis, 

differ by

6.1-7.3%, 

P. serrata, differ 

by 6.3-6.4%

Spring complex 

along Owyhee River 

upflow from Tudor 

Warm Springs, 

Malheur Co., OR

Pyrgulopsis

sp. 2

HPL2008 

isolate

IP208A, 

IP208E in 

GenBank

P210 FJ172480 3 0.00% 2.1%

with IP69B

P. kolobensis, 

differ by

6.2-7.4%, 

P. serrata,

differ by 6.4%

Tudor Warm

Springs, east side of 

Owyhee River, third 

spring from north, 

Malheur Co., OR

Pyrgulopsis

sp. 2

HPL2008 

isolate

IP210A in 

GenBank

P212 FJ172483 2 0.00% 2.1%

with IP69B

P. kolobensis, 

differ by

6.2-7.4%, 

P. serrata,

differ by 6.4%

Tudor Warm

Springs, east side 

of Owyhee River, 

second spring from 

north, Malheur Co., 

OR

Pyrgulopsis

sp. 2

HPL2008 

isolate

IP212A in 

GenBank0

HYDROBIIDAE
Technical Note 449 101

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

gibba P134/SV AY197603, AY426359 2 0.00% 1.0%with P192

P. vinyardi, differ 

by 0.5%, 

P. kolobensis, 

differ by

4.6-5.9%,

P. nonaria, differ 

by 4.4-4.8%

Spring west of Fee 

Reservoir, Surprise 

Valley, Modoc Co., 

CA

P192 DQ364016 1 0.00% 1.0%

with P134

P. vinyardi, differ 

by 1.1%, 

P. kolobensis, 

differ by

4.6-5.5%,

P. nonaria, differ 

by 4.6%

Spring along 

Hwy 395 below 

Bridgeport 

Reservoir, Walker 

River basin, Mono 

Co., CA

Pyrgulopsis

gilae G1

KC571284 

(N=2), 

KC571285 

(N=1)

3 0-1.2% 0.6-1.5% with 

Pgil3 and Pgil4

P. marilynae,

differ by

3.8-4.0%

Spring along East 

Fork Gila River, ca. 

1.53 km north, 2.9 

km east of State 

Route 527 bridge 

crossing, Grant Co., 

NM

G2

KC571286 

(N=9), 

KC571287 

(N=1)

10 0-0.2% 0.6-1.0% with 

Pgil3 and Pgil4

P. marilynae,

differ by

3.5-3.8%

Spring along East 

Fork Gila River, ca. 

1.29 km north, 0.56 

km west of Black 

Canyon confluence, 

Grant Co., NM

G3

KC571288 

(N=4), 

KC571289 

(N=1),

KC571290 

(N=1)

6 0-0.3% 0.5-1.2%

with G1

P. marilynae,

differ by

3.6-4.0%

Spring along East 

Fork Gila River, ca. 

1.53 km north, 2.38 

km east of State 

Route 527 bridge 

crossing, Grant Co., 

NM

G14

KC571288 

(N=2), 

KC571306 

(N=1)

3 0-0.2% 0.5-1.2%

with G1

P. marilynae,

differ by

3.6-4.0%

Alum Hot Spring, 

ca. 1.93 km south, 

0.16 km west of 

State Route 527 

bridge crossing, 

Grant Co., NM

Pgilae AY627952 1 0.00% 0.0-1.3%

with G1

P. marilynae,

differ by

3.8-4.0%

Spring tributary to 

East Fork Gila River, 

Grant Co., NM

Pgil1, 

Pgil2

AY485570 

(N=18),

AY485571 

(N=14)

32 0-0.2% 0.0-1.4%

with G1

P. marilynae,

differ by

3.2-3.6%

Gila I Hurt (2004)

Pgil3, 

Pgil4

AY485572 

(N=28), 

AY485573 

(N=1)

29 0-0.4% 0.6-1.5%

with G1

P. marilynae,

differ by

3.5-3.8%

Gila II Hurt (2004)

Pgil3 AY485572 29 0.00% 0.6-1.2%

with G1

P. marilynae,

differ by

3.5-3.6%

Gila III Hurt (2004)

HYDROBIIDAE
102 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

giulianii M23 AF520937 1 0.00% 0% with P21 P. diablensis, differ by 1.1%

Stream, Sand 

Canyon, Indian

Wells Valley 

(Northern Mojave 

basin), Kern Co., CA

P. guiliani in 

GenBank

P21 DQ364018 1 0.00% 0% with M23 P. diablensis, 

differ by 1.1%

Dougherty Creek, 

Hwy 178 crossing, 

Kern River basin, 

Kern Co., CA

Pyrgulopsis 

glandulosa NP2 AY627959 1 0.00% 0% with Pgla1

P. intermedia, 

differ by

2.1-2.6%

Nelson Place 

Spring, Verde River 

basin, Yavapai Co., 

AZ

Pgla1 AY485557 25 0.00% 0% with NP2

P. intermedia, 

differ by

2.2-2.4%

Nelson Place Hurt (2004)

Pyrgulopsis 

gracilis P60 DQ364011 1 0.00%

P. hubbsi,

differ by 3.2%, 

P. sathos,

differ by 3.2%

Emigrant Springs 

(north), White River 

Valley, Nye Co., NV

Pyrgulopsis

greggi P22

AF520943 

(N=3), 

GQ275088 

(N=2)

5 0-0.3%

P. kolobensis, 

differ by

4.3-6.5%,

P. pilsbryana, 

differ by

4.8-5.7%,

P. transversa, 

differ by

4.6-5.3%,

P. sp. C, differ

by 4.8-5.5%,

P. sp. H, differ

by 4.8-5.3%

Grapevine Creek, 

Fort Tejon Historic 

State Park, Kern 

Co., CA

Pyrgulopsis 

hualapaiensis KU720383 1 0.00%

P. conica, differ

by 2.9%, 

P. milleri, differ

by 2.9%, 

P. thompsoni, 

differ by

2.5-4.0%

Upper Peach 

Springs, outflow

just below concrete 

weir, Hualapai

Indian Reservation, 

Mohave Co., AZ

Pyrgulopsis 

hubbsi P7B AY627918 1 0.00% P. sathos,differ by 1%

Crystal Spring, 

Pahranagat Valley, 

Lincoln Co., NV

Pyrgulopsis 

imperialis P140

AY379450 

(P140A), 

AY426350 

(P140C)

2 0.30%

P. cybele, differ

by 3.1-3.7%, 

P. intermedia, 

differ by

3.3-4.0%,

P. perforata,

differ by

3.2-3.7%, 

P. robusta, differ 

by 3.4-4.1%

Spring, Thacker 

Pass, Kings River 

Valley, Humboldt 

Co., NV

HYDROBIIDAE
Technical Note 449 103

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

inopinata P100

AY426360 

(P100A), 

AY426361 

(P100C)

2 0.00%

P. kolobensis, 

differ by

2.6-3.6%,

P. marcida,

differ by 2.6%, 

P. plicata,

differ by 2.6%

Spring, Glenwood, 

Sevier River 

drainage Sevier 

Co., UT

Pyrgulopsis 

intermedia

P. intermedia, 

OW_b 

AF520928, 

AY197593 2 0.20% 0.2-0.7%with P223

P. glandulosa, 

differ by

2.2-2.6%

Crooked Creek 

Spring State 

Wayside, Harney 

Co., OR

OW_a, 

P1B/

intermedia1,

P1C/

intermedia1,

P1E

AY197592,

AY379442,

AY379443,

AY426351

4 0-0.2% 0.2-0.6%

with P223

P. glandulosa, 

differ by

2.2-2.6%

Crooked Creek, 

Hwy 95 crossing, 

middle Snake River 

basin, Malheur 

Co., OR

P2 AY426352

AY426353 2 0.00% 0.3-0.6%with P223

P. glandulosa, 

differ by

2.2-2.4%

Crooked Creek 

Spring, Owyhee 

River drainage, 

Malheur Co., OR

P4/

intermedia2

AY379444, 

AY379445 2 0.20% 0.2-0.8%with P223

P. glandulosa, 

differ by

2.2-2.6%

Skylight Spring, 

Barren Valley, 

Malheur Co., OR

IP60 FJ172460-63 4 0-0.2% 0.3-0.8%

with P223

P. glandulosa, 

differ by

2.2-2.6%

Crooked Creek at 

Crooked Creek

State Wayside, 

Harney Co., OR

IP67 FJ172468-69 2 0.00% 0.2-0.5%

with P223

P. glandulosa, 

differ by

2.2-2.3%

Spring tributary to 

Birch Creek,

Malheur Co., OR

P217 FJ172488 1 0.00% 0.2-0.5%

with P223

P. glandulosa, 

differ by

2.2-2.3%

Spring on Owyhee 

River, above Long 

Sweetwater rapids, 

Malheur Co., OR

P222 FJ172496-497 2 0.20% 0.2-0.6%

with P223

P. glandulosa, 

differ by

2.2-2.4%

Spring on hillside, 

Jackson Hole, 

Malheur Co., OR

P223 FJ172498-499 2 0.30% 0.3-0.8%

with IP60

P. glandulosa, 

differ by

2.0-2.4%

Mouth of Rinehart 

Creek, Malheur 

Co., OR

P224 FJ172500 1 0.00% 0.2-0.5%

with P223

P. glandulosa, 

differ by

2.2-2.3%

Spring west of 

Twomile Spring, 

Malheur Co., OR

Pyrgulopsis 

isolata AY367486 1 0.00%

P. fairbanksensis, 

differ by 0.5%, 

P. nanus, differ

by 1.4%

Spring south of

Clay Pits, Ash 

Meadows, Nye Co., 

NV

HYDROBIIDAE
104 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

kolobensis P162

AY379448 

(P162A), 

AY379449 

(P162B),

AY379449 

(P162B),

AY426354 

(P162D)

3 0.00% 3.1%

with P193

P. pilsbryana, 

differ by

0.2-0.3%,

P. transversa, 

differ by 1.1%, 

P. sp. C, differ

by 0.3%, 

P. sp. D, differ

by 0.2%, 

P. sp. H, differ

by 0.3%

Big Malad Spring, 

Malad Valley,

Oneida Co., ID

P117 AY627939

(Tq3, P117A) 1 0.00% 2.9% with P162 P. variegata,differ by 1.9%

Toquerville Springs, 

Washington Co., UT

P193 DQ364008 1 0.00% 3.1% with P162 P. variegata,

differ by 1.9%

Spring southwest

of Pinto, 

Washington Co., UT

P194 DQ364009 1 0.00%

1.9% with Kol3, 

Kol4, P162, 1.7-

1.9% with Kol5

P. variegata,

differ by 1.1%

Spring, Left Fork 

Santa Clara 

River, Pine Valley 

campground, 

Washington Co., UT

P195 DQ364010 1 0.00% 2.6%

with P162

P. variegata,

differ by 1.6%

Spring, Meadow 

Valley Wash, 

Kershaw-Ryan 

State Park, Lincoln 

Co., NV

Kol1

KT248032 

(Kol1A, N=4),

KT248033 

(Kol1B, N=1)

5 0-0.8% 2.3-2.8%

with P193

P. nonaria, differ 

by 0.6-1.1%, 

P. pilsbryana, 

differ by

0.0-0.9%,

P. transversa, 

differ by

0.8-1.0%,

P. sp. C, differ

by 0.2-1.0%,

P. sp. D, differ

by 0.0-0.8%, 

P. sp. H, differ

by 0.2-0.9%

Spring, Right Fork 

Hobble Creek, Utah 

Co., UT

Kol2

KT248034 

(Kol2A, N=1),

KT248035 

(Kol2B, N=3),

KT248036 

(Kol2E, N=1)

5 0-0.3%

2.0-2.2% with 

P117, 2.0-2.3% 

with P193

P. nonaria, differ 

by 0.5-0.6%,

P. pilsbryana, 

differ by

0.6-0.9%, 

P. transversa, 

differ by

0.6-0.8%,

P. sp. C, differ

by 0.8-1.0%, 

P. sp. D, differ

by 0.6-0.8%,

P. sp. H, differ

by 0.-8-0.9%

Spring Creek, 

Wallsburg, 

Wasatch Co., UT

Kol3 KT248037 

(Kol3A, N=5) 5 0.00% 3.0%with P193

P. pilsbryana, 

differ by

0.2-0.3%, 

P. transversa, 

differ by 1.1%, 

P. sp. C, differ

by 0.3%,

P. sp. D, differ

by 0.2%,

P. sp. H, differ

by 0.3%

Willow Creek, 

Wasatch Co., UT

HYDROBIIDAE
Technical Note 449 105

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

kolobensis

(continued)

Kol4 KT248038 

(Kol4A, N=5) 5 0.00%

2.2-2.9%

with

Pkol1/Pkol2, 

2.6%

with

P117,

P193

P. pilsbryana, 

differ by

0.2-0.3%, 

P. transversa, 

differ by 1.1%, 

P. sp. C, differ

by 0.3%, 

P. sp. D, differ

by 0.2%, 

P. sp. H, differ

by 0.3%

Spring at Peoa, 

Summit Co., UT

Kol5

KT248039 

(Kol5A, N=2), 

KT248040 

(Kol5B, N=1),

KT248041 

(Kol5C, N=1),

KT248042 

(Kol5D, N=1)

5 0-0.5%

2.1-3.1%

with

Pkol1/Pkol2,

2.6-2.8%

with P117,

2.5-2.8%

with P193

P. pilsbryana, 

differ by

0.0-0.5%, 

P. transversa, 

differ by

1.0-1.3%,

P. sp. C, differ

by 0.2-0.5%,

P. sp. D, differ

by 0.0-0.3%, 

P. sp. H, differ

by 0.2-0.5%

Spring southwest 

of Francis, Summit 

Co., UT

Pkol1,

Pkol2

AY485532 

(Pkol1, N=29), 

AY485533 

(Pkol2, N=4) 

33 0-0.2% 2.5-3.1%

with P162

P. variegata,

differ by

1.6-2.1%

Toquerville Hurt (2004)

Pyrgulopsis 

landyei P68A AY627930 1 0.00% P. sulcata,differ by 3.7%

Middle Spring, 

Steptoe Ranch, 

Steptoe Valley, 

White Pine Co., NV

Pyrgulopsis 

lasseni WC_a AY197604 1 0.00% 0.3%with WC_a

P. eremica, differ 

by 0.3-6.8%, 

P. kolobensis, 

differ by

4.4-5.7%,

P. pilsbryana, 

differ by

4.8-4.9%,

P. transversa, 

differ by 4.8%,

P. sp. D, differ

by 4.8%

Willow Creek west 

of Lower McBride 

Springs, Lassen 

Co., CA

WC_b AY197605 1 0.00% 0.3%

with WC_b

P. eremica, differ 

by 0.3-6.8%, 

P. kolobensis, 

differ by

4.8-6.0%,

P. pilsbryana, 

differ by

5.1-5.3%,

P. transversa, 

differ by 5.1%,

P. sp. D,

differ by 5.1%

Willow Creek 

downstream from 

Willow Creek 

Campground, 

Lassen Co., CA

HYDROBIIDAE
106 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

lata P61B AY627927 1 0.00%

P. breviloba,

differ by 0.5%, 

P. kolobensis, 

differ by

3.9-5.2%,

P. marcida,

differ by 4.2%,

P. nonaria,

differ by 4.2%

Butterfield Springs, 

White River Valley, 

Nye Co., NV

Pyrgulopsis 

lentiglans P103A AY627936 1 0.00% P. millenaria,differ by 1.6%

Crittenden Springs, 

Thousand Springs 

Creek drainage, 

Elko Co., NV

Pyrgulopsis 

leporina P123A EU700471 1 0.00% P. sublata, differ by 1.9%

Springs along 

Rabbit Creek, 

Humboldt River 

drainage, Elko Co., 

NV

Pyrgulopsis

licina M7

AY367438 

(M7A), 

AY367439 

(M7B)

2 0.20% 0.6-0.8%

with M29

P. bifurcata,

differ by

3.5-3.7%

Last Chance Spring, 

Ash Meadows, Nye 

Co., NV

P. micrococcus

in GenBank

M29 AY367476 1 0.00% 0.6-0.8%

with M7

P. bifurcata,

differ by 3.8%

Spring east of 

Crystal Reservoir, 

Ash Meadows, Nye 

Co., NV

P. micrococcus

in GenBank

M30 AY367477 

(M30A, M30C) 2 0.00% 0.5-0.6%with M7

P. bifurcata,

differ by 3.7%

Spring south of 

Clay Pits, Ash 

Meadows, Nye Co., 

NV

P. micrococcus

in GenBank

M52 KF559189 2 0.00% 0.5-0.6%

with M7

P. bifurcata,

differ by 3.7%

Spring ca. 100 m 

north of Collins 

Ranch, Ash 

Meadows, Nye Co., 

NV

P. perforata in 

GenBank

M54 KF559190 3 0.00% 0.5-0.6%

with M7

P. bifurcata,

differ by 3.7%

Spring south of 

Rogers Spring, Ash 

Meadows, Nye Co., 

NV

P. perforata in 

GenBank

M58 KF559191 2 0.00% 0.5-0.6%

with M7

P. bifurcata,

differ by 3.7%

Spring east of 

Crystal Reservoir, 

Ash Meadows, Nye 

Co., NV

P. perforata in 

GenBank

Pyrgulopsis 

limaria SM2 EF119088-92 6 0-0.9%

0.3-1.5% with 

SM10, 0.2-1.4% 

with SM22

P. umbilicata, 

differ by

0.2-0.9%

Spring brook, 

western Soldier 

Meadow, Humboldt 

Co., NV

SM7 EF119095-97 5 0-0.3% 0.5-1.1% with 

SM10

P. umbilicata, 

differ by

0.2-0.5%

Spring 150 m W 

of Satellite Spring, 

Humboldt Co., NV

SM10 EF119098-100 5 0-0.3%

0.3-1.2%

with SM23, 

SM36, 0.3-1.1% 

with SM24, 

SM26,

SM29/31, 

0.3-1.4% with 

SM2, 0.5-1.1% 

with SM7

P. umbilicata, 

differ by

0.6-1.2%

Tole Spring,

western Soldier 

Meadow,

Humboldt Co., NV

HYDROBIIDAE
Technical Note 449 107

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

limaria

(continued)

SM22 EF119106-110 5 0.2-0.6%

0.2-1.1% with 

SM23, SM36,

0.3-1.1%

with SM26,

0.2-1.4%

with SM2

P. umbilicata, 

differ by

0.5-1.1%

Tole Spring 30 m 

downflow from 

source, western 

Soldier Meadow, 

Humboldt Co., NV

SM23 EF119111-114 4 0.2-0.6%

0.3-1.2% with 

SM10, 0.2-1.1% 

with SM22

P. umbilicata, 

differ by

0.2-0.6%

Satellite Spring 

complex, spring "A," 

60+ m downflow 

from source, 

Humboldt Co., NV

SM24 EF119115-116 4 0.50% 0.3-1.1% with 

SM10

P. umbilicata, 

differ by

0.2-0.6%

Satellite Spring 

complex, spring "B," 

Humboldt Co., NV

SM26 EF119117-120 4 0.3-0.6%

0.0-1.1% with 

SM2, 0.3-1.1% 

with SM10, 

SM22

P. umbilicata, 

differ by

0.2-0.8%

Satellite Spring 

complex, spring "F," 

Humboldt Co., NV

SM29,

31 EF119121-123 7 0-0.5% 0.3-1.1% with SM10

P. umbilicata, 

differ by

0.2-0.6%

Satellite Spring 

complex, spring "H," 

Humboldt Co., NV

SM34 EF1190124 2 0.00% 0.0-0.9% with 

SM2

P. umbilicata, 

differ by

0.3-0.6%

Satellite Spring 

complex, spring "J," 

Humboldt Co., NV

SM36 EF119125-126 5 0-0.6%

0.3-1.2% with 

SM10, 0.2-1.1% 

with SM22

P. umbilicata, 

differ by

0.3-0.6%

Satellite Spring 200 

m downflow from 

source, Humboldt 

Co., NV

Pyrgulopsis

lindae 17/P24 GQ275091 4 0.00%

P. kolobensis, 

differ by

5.6-7.8%, 

P. pilsbryana, 

differ by 6.0%,

P. sp. C, differ

by 5.9-6.1%, 

P. sp. C, differ

by 5.9-6.1%, 

P. sp. D, differ

by 6.0%

San Domingo Creek, 

San Joaquin River 

basin, Calaveras 

Co., CA

P. sp. 

USNM905258 

isolate

P24AA in 

GenBank, 

described as 

P. lindae in 

Hershler et al. 

(2016b)

Pyrgulopsis 

lockensis P82D AY627932 1 0.00% P. villacampae, differ by 2.6%

Big Spring, Lockes 

Ranch, Railroad 

Valley, Nye Co., NV

Pyrgulopsis 

longiglans P133A EF119141, DQ364017 1 0.00%

P. kolobensis, 

differ by

4.5-5.9%, 

P. variegata,

differ by 5.0%

Spring NNW of 

Holbrook Junction, 

Antelope Valley, 

Douglas Co., NV

DQ364017: 

P. sp. Virginia

HPL-2006 in 

GenBank

Pyrgulopsis 

longinqua P32B DQ364006 1 0.00%

P. californiensis, 

differ by

2.4-7.9%, 

P. montana,

differ by 2.9%

Spring westsouthwest of 

Hunters Spring, 

Salton Sea basin, 

Riverside Co., CA

HYDROBIIDAE
108 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

marcida P59A DQ364012 1 0.00%

P. kolobensis, 

differ by

1.1-2.7%,

P. serrata,

differ by 1.4%,

P. sterilis, differ

by 1.4%,

P. variegata,

differ by 1.4%

Hardy Springs,

White River Valley, 

Nye Co., NV

Pyrgulopsis 

marilynae G8

KC571299 

(N=1), 

KC571300 

(N=1),

KC571301 

(N=3),

KC571302 

(N=1)

6 0-0.3% 0.2-0.3%

with G11

P. gilae,

differ by

3.2-4.0%

Spring along

Middle Fork Gila 

River, ca. 0.97 km 

north, 0.64 km

west of Jordan 

Canyon, Grant Co., 

NM

P. gilae in 

GenBank

G11 KC571305 2 0.00% 0.2-0.3%

with G8

P. gilae,

differ by

3.4-4.0%

Spring along Middle 

Fork Gila River, ca. 

0.48 km north, 

0.48 km west of 

Jordan Canyon, 

Grant Co., NM

P. gilae in 

GenBank

Pyrgulopsis 

merriami P8B AY627919 1 0.00%

P. notidicola,

differ by

8.4-9.7%, 

P. trivialis, differ 

by 8.6-8.9%,

Ash Spring, 

Pahranagat Valley 

Lincoln Co., NV

Pyrgulopsis 

micrococcus M1

AF520944/

AY367415-417/

AY367420/

AY367424/

AY367427-28 

(N=8, M1A-1D, 

G, K, N-O), 

AY367418-19 

(N=2, M1E-F), 

AY367421-23/

AY367425-26/

AY367429 

(N=6, M1H-J, 

L-M, P)

16 0-0.6% 0.0-0.3%

with M31

P. perforata,

differ by

3.9-4.5%

AF520944:

Spring just S of 

Springdale, Nye 

Co., NV

AY367418-29: 

Spring, Fleur de 

Lis Ranch, Oasis 

Valley, Nye Co., NV

M31 AY367478 1 0.00% 0.0-0.3%

with M1

P. perforata,

differ by

3.9-4.3%

Goss Springs, 

Oasis Valley, Nye 

Co., NV

Pyrgulopsis 

militaris

SM/P147A, 

P147C,

P147D,

P147E,

P147F

AY197596,

AY426362,

EF119077,

EF119078,

EF119079

5 0.0-0.9% 1.5-2.2%

with SM1

P. varneri,

differ by

2.0-2.7%

Spring, Soldier 

Meadow, Humboldt 

Co., NV

SM1 EF119076 5 0.00% 1.5-2.2%

with P147

P. varneri, differ 

by 2.3-2.7%

West Spring, Bog 

Hot Valley,

Humboldt Co., NV

HYDROBIIDAE
Technical Note 449 109

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

millenaria P104A EU700469 1 0.00% P. lentiglans,differ by 1.6%

Spring below 

Twentyone Mile 

Dam, Thousand 

Springs Creek 

drainage, Elko Co., 

NV

Pyrgulopsis

milleri P23 GQ275096 3 0.00%

P. glandulosa, 

differ by

2.8-3.3%,

P. hualapaiensis, 

differ by 2.9%,

P. perforata,

differ by

2.8-2.9%, 

P. thompsoni, 

differ by

2.7-3.4%,

P. torrida, differ

by 2.8%

Pierpoint Spring, 

Tulare-Buena Vista 

Lakes drainage, 

Tulare Co., CA

Pyrgulopsis

sp. 2

HPL2009 

isolate P23B

in GenBank

Pyrgulopsis 

montana P118A AY627940 1 0.00%

P. bacchus,

differ by

2.2-2.6%

Spring, upper Camp 

Valley, Meadow 

Valley Wash, Lincoln 

Co., NV

Pyrgulopsis 

montezumensis

Pmon1, 

Pmon2

AY485552 

(Pmon1, N=33), 

AY485553 

(Pmon2, N=1)

34 0.0-0.2%

P. glandulosa, 

differ by

3.3-3.8%,

P. intermedia, 

differ by

3.6-4.1%,

P. perforata,

differ by

3.6-3.9%,

P. robusta, differ 

by 3.7-4.4%

Montezuma Well Hurt (2004)

Pyrgulopsis 

morrisoni P150 DQ364007 1 0.00%

0.0-1.7% with 

Pmor1/2/3/4, 

0.5% with

Pmor5

P. glandulosa, 

differ by

3.1-3.5%

Spring, Bubbling 

Pond Hatchery, 

Verde River basin, 

Yavapai Co., AZ

Pmor1, 

Pmor2,

Pmor3, 

Pmor4

AY485547 

(Pmor1, N=12), 

AY485548 

(Pmor2, N=4), 

AY485549 

(Pmor3, N=3), 

AY485550 

(Pmor4, N=6)

26 0-1.7%

0.0-1.7% with 

P150, 0.2-1.5% 

with Pmor5

P. glandulosa, 

differ by

2.8-3.6%

Bubbling

Hurt (2004), 

listed 

frequency

only added

to 0.961

Pmor5 AY185551 30 0.00%

0.5% with P150, 

0.2-1.5% with 

Pmor1/2/3/4

P. glandulosa, 

differ by

3.3-3.6%

Page Hurt (2004)

Pyrgulopsis

nanus AY367487 1 0.00%

P. fairbanksensis, 

differ by 1.3%, 

P. isolata,

differ by 1.4%

Five Springs, Ash 

Meadows, Nye Co., 

NV

Pyrgulopsis 

neritella AF520951 1 0.00% P. sulcata, differ by 3.2%

Big Spring, Steptoe 

Ranch, White Pine 

Co., NV

HYDROBIIDAE
110 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

nonaria P97B EU700467 1 0.00%

P. kolobensis, 

differ by

0.5-2.5%, 

P. pilsbryana, 

differ by

1.1-1.2%,

P. transversa, 

differ by 1.1%,

P. sp. D,

differ by 1.1%

Spring east of 

Ninemile Reservoir, 

Sevier River 

drainage, Sanpete 

Co., UT

Pyrgulopsis 

notidicola SM6 EF119128 4 0.00%

0-2.6%

with SM35,

1.1%

with SM9,

SM30

P. limaria, differ 

by 6.2-7.0%, 

P. umbilicata, 

differ by

6.4-6.5%

Satellite Spring, 

western Soldier 

Meadow, Humboldt 

Co., NV

SM9 EF119132-33 5 0-0.3% 1.1-2.1%

with SM35

P. limaria, differ 

by 5.8-7.1%, 

P. umbilicata, 

differ by

5.9-6.4%

Satellite Spring 

complex, spring 

brook, Humboldt 

Co., NV

SM27 EF119134-35 4 0-0.5% 0.9-2.3%

with SM35

P. limaria, differ 

by 5.9-7.1%, 

P. umbilicata, 

differ by

6.1-6.4%

Satellite Spring 

complex, spring 

brook, Humboldt 

Co., NV

SM30 EF119136 3 0.00% 1.1-1.8%

with SM35

P. limaria, differ 

by 6.1-7.1%, 

P. umbilicata, 

differ by

6.2-6.4%

Satellite Spring 

complex, spring “H,” 

Humboldt Co., NV

SM32 EF119137-38 3 0-0.2% 0.9-2.0%

with SM35

P. limaria, differ 

by 5.9-7.1%, 

P. umbilicata, 

differ by

6.1-6.4%

Satellite Spring 

complex, spring “I,” 

Humboldt Co., NV

SM33 EF119139 5 0.00% 0.9-2.0%

with SM35

P. limaria, differ 

by 5.9-7.0%, 

P. umbilicata, 

differ by

6.1-6.2%

Satellite Spring 

complex, spring “J,” 

Humboldt Co., NV

SM35 EF119129-30, 

EF119140 5 0-2.7%

0.9-2.3% with 

SM27, 1.1-2.1% 

with SM9,

0-2.6% with

SM6, 0-2.2%

with P146

P. limaria, differ 

by 6.1-7.1%, 

P. umbilicata, 

differ by

6.2-6.7%

Satellite Spring, 

western Soldier 

Meadow, Humboldt 

Co., NV

P146 EF119131 1 0.00%

0-2.2% with 

SM35, 1.0% with 

SM9, SM30

P. limaria,

differ by

6.1-6.9%, 

P. umbilicata, 

differ by

6.3-6.4%

Northernmost 

spring of large 

complex, western 

Soldier Meadow, 

Humboldt Co., NV

Pyrgulopsis 

ojaiensis M56 KF559192 3 0.00% 0.8%with P29

P. castaicensis, 

differ by 3.6%, 

P. glandulosa, 

differ by

3.7-4.4% 

Grapevine Springs, 

Death Valley, Inyo 

Co., CA

P. sp. M56 in 

GenBank

HYDROBIIDAE
Technical Note 449 111

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

ojaiensis

(continued)

P29 GQ275093 4 0.00% 0.8%

with M56

P. castaicensis, 

differ by 3.8%, 

P. glandulosa, 

differ by

3.5-4.3%

Spring tributary 

to Sisar Creek, 

Southern California 

coastal drainage, 

Ventura Co., CA

P. sp. 

USNM905259 

isolate P29A 

in GenBank, 

placed under 

P. stearnsiana

in Hershler 

and Liu

(2010),

placed under 

clade F in 

Hershler et 

al. (2013), 

described as 

P. ojaiensis in 

Hershler et al. 

(2016b)

Pyrgulopsis 

owensensis P34

AF520922 

(P34C, N=1), 

GQ275089 

(P34AA, N=2), 

GQ275090 

(P34AB, N=2)

5 0-0.5%

P. perturbata, 

differ by

1.0-1.3%

Spring, Graham 

Ranch, ca. 8.0 km 

E of Big Pine, Inyo 

Co., CA

Pyrgulopsis 

owyheensis IP59

FJ172457 

(IP59A, N=1), 

FJ172458 

(IP59C, N=1),

FJ172459 

(IP59D, N=1)

3 0.2-0.3%

1.8-2.1%

with P211,

1.8-2.0%

with P213

P. aurata,

differ by

4.7-4.9%, 

P. kolobensis, 

differ by

4.2-5.9%,

P. pictilis, differ

by 4.7-4.8%, 

P. plicata, differ

by 4.7-4.8%,

P. sublata, differ 

by 4.6-4.8%

Owyhee Spring, 

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

IP59A,

IP59C,

IP59D in 

GenBank

IP62

FJ172464 

(IP62A, N=1), 

FJ172465 

(IP62B, N=1),

FJ172466 

(IP62C, N=1),

FJ172467 

(IP62D, N=1)

4 0.00% 1.5-1.7%

with IP59

P. kolobensis, 

differ by

4.0-5.7%,

P. plicata,

differ by 4.7%

Tree Spring,

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

IP62A, IP62B, 

IP62C, IP62D 

in GenBank

P177B FJ172475 1 0.00% 2.0-2.1%

with IP59

P. bryanwalkeri, 

differ by 5.0%, 

P. kolobensis, 

differ by

4.2-5.7%,

P. plicata,

differ by 5.0%,

Tudor Warm

Springs, east side

of river, second 

spring from north, 

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

P177B in 

GenBank

P209 FJ172479 5 0.00% 1.4-1.5% with 

IP59

P. lentiglans,

differ by 4.8%, 

P. kolobensis, 

differ by

4.2-5.7%,

P. pictilis,

differ by 4.8%,

P. plicata,

differ by 4.8%

Kane Springs, 

Malheur River

basin, Malheur 

Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

P209A in 

GenBank

HYDROBIIDAE
112 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

owyheensis

(continued)

P211

FJ172481 

(P211A, N=1), 

FJ172482 

(P211B, N=1)

2 0.20% 1.8-2.1%

with IP59

P. kolobensis, 

differ by

4.4-6.4%, 

P. plicata,

differ by

4.7-4.8%

Tudor Warm 

Springs, east side 

of river, third spring 

from north, Malheur 

Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

P21AA,

P211B in 

GenBank

P213A FJ172484 1 0.00% 1.8-2.0%

with IP59

P. kolobensis, 

differ by

4.4-5.9%, 

P. plicata,

differ by 4.7% 

Tudor Warm 

Springs, east side

of river, second 

spring from north, 

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate P213A 

in GenBank

P215 FJ172485 2 0.00% 1.5-1.7%

with IP59

P. kolobensis, 

differ by

4.4-5.7%,

P. lentiglans,

differ by 4.8%

Spring tributary 

to Owyhee River 

upflow from Tudor 

Warm Springs, 

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate P215A 

in GenBank

P216

FJ172486 

(P216A), 

FJ172487 

(P216B)

4 0.30% 1.4-1.5%

with IP59

P. kolobensis, 

differ by

3.9-5.5%,

P. lentiglans,

differ by

4.8-4.9%

Spring on Owyhee 

River, below mouth 

of Crooked Creek, 

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

P216A, P216B 

in GenBank

P218

FJ172489 

(P218A), 

FJ172490 

(P218B),

FJ172491 

(P218C)

4 0-0.5% 1.4-1.7%

with IP59

P. kolobensis, 

differ by

3.9-5.5%,

P. plicata, differ

by 4.5-4.7%, 

P. sublata, differ 

by 4.5-4.8%

Spring on Owyhee 

River, above Long 

Sweetwater rapids, 

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

P218A,

P218B, P218C 

in GenBank

P219,

P220

FJ172492 

(P219A), 

FJ172493 

(P220B),

FJ172494 

(P220C)

7 0-0.3% 1.4-1.5%

with IP59

P. kolobensis, 

differ by

3.9-5.7%,

P. plicata, differ

by 4.5-4.8%

Weeping Wall 

springs, just below 

mouth of Granite 

Creek, Malheur 

Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

P219A,

P220B, P220C 

in GenBank

P221 FJ172495 3 0.00% 1.4-1.5%

with IP59

P. kolobensis, 

differ by

3.9-5.5%, 

P. plicata,

differ by 4.5%

Spring on Owyhee 

River, above

Artillery rapids, 

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate P221B 

in GenBank

P225,

P226

FJ172501 

(P225A), 

FJ172502 

(P226A)

4 0.30% 1.4-1.5%

with IP59

P. kolobensis, 

differ by

3.9-5.5%,

P. plicata, differ

by 4.5-4.7%,

P. sublata, differ 

by 4.6-4.8%

Seep wall on 

Owyhee River, just 

above mouth of 

Crooked Creek, 

Malheur Co., OR

Pyrgulopsis

sp. 3

HPL2008 

isolate

P225A,

P226A in 

GenBank

Pyrgulopsis 

peculiaris AF520912 1 0.00%

P. kolobensis, 

differ by

5.9-6.9%, 

P. pilsbryana, 

differ by

5.9-6.1%,

P. sp. D, differ

by 5.9%

Spring above 

Swasey Spring, 

Whirlwind Valley, 

Millard Co., UT

Pyrgulopsis

pellita P125A AY627943 1 0.00% P. bifurcata,differ by 3.3%

Sullivan Spring, 

Antelope Valley, 

Eureka Co., NV

HYDROBIIDAE
Technical Note 449 113

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

perforata M3

AY367433 

(M3A), 

AY367434 

(M3B, M3O)

3 0.00% 0.2-0.3%

with M4

P. conica, differ

by 2.6%, 

P. diablensis, 

differ by 2.6%,

P. glandulosa, 

differ by

2.6-2.7%,

P. simplex, differ 

by 2.7%

Spring east of 

Scottys Castle, 

Grapevine Canyon, 

Death Valley, Inyo 

Co., CA

P. micrococcus

in GenBank

M4

AY367435 

(M4A), 

AY367436 

(M4B)

2 0.50%

0.2-0.3%

with M3,

0-0.5%

with M5

P. conica, differ

by 2.6-2.7%, 

P. diablensis, 

differ by

2.6-2.7%,

P. glandulosa, 

differ by

2.4-2.7%,

P. simplex, differ 

by 2.6-2.7%

Spring west of 

Scottys Castle, 

Grapevine Canyon, 

Death Valley, Inyo 

Co., CA

P. micrococcus

in GenBank

M5 AY367437 

(M5A) 1 0.00%

0.2%

with M3,

0-0.5%

with M4

P. conica, differ

by 2.6%, 

P. diablensis, 

differ by 2.7%,

P. glandulosa, 

differ by

2.4-2.6%,

P. simplex, differ 

by 2.6%

Surprise Spring, 

Death Valley, Inyo 

Co., CA

P. micrococcus

in GenBank

Pyrgulopsis 

perturbata AY367488 1 0.00%

P. owensensis, 

differ by

1.0-1.3%

Northeast Spring, 

Fish Slough, Owens 

River basin, Mono 

Co., CA

Pyrgulopsis 

pictilis P126A AY627944 1 0.00%

P. aurata, differ 

by 0.0%, 

P. bryantwalkeri, 

differ by 0.3%,

P. kolobensis, 

differ by

1.7-2.9%,

P. sp. B, differ

by 1.9%

Cain Spring, 

Antelope Valley, 

Lander Co., NV

Pyrgulopsis 

pilsbryana P137 AY426363-64 2 0.00%

0.2%

with

P159A

P. kolobensis, 

differ by

0.2-2.9%, 

P. transversa, 

differ by 1.0%,

P. sp. C, differ

by 0.2%, 

P. sp. D, differ

by 0.0%, 

P. sp. H, differ

by 0.2%

Spring, Saint 

Charles Creek, Bear 

Lake Valley, Bear 

Lake Co., ID

P159A EU700475 1 0.00%

0.2%

with

P137

P. kolobensis, 

differ by

0.2-2.7%, 

P. transversa, 

differ by 1.1%,

P. sp. C,

differ by 0.0%,

P. sp. D,

differ by 0.2%,

P. sp. H, differ

by 0.0%

Bear River, Black 

Canyon, Caribou 

Co., ID

Pyrgulopsis

sp. Black 

Canyon

isolate

P159A in 

GenBank

HYDROBIIDAE
114 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

pisteri P157A DQ364004 1 0.00%

P. crystalis, 

differ by 0.6%, 

P. erythropoma, 

differ by 4.5%

Marsh Spring, 

Ash Meadows, 

Amargosa River 

basin, Nye Co., NV

Pyrgulopsis 

plicata P99A AY627935 1 0.00% P. marcida,differ by 1.6%

Spring, Black 

Canyon, Sevier 

River drainage, 

Garfield Co., UT

Pyrgulopsis 

robusta

HL/

hendersoni1/

P3

AY197594/

AY379430-31 2 0.00%

0.5% with P171, 

Ida, P181, 0.5-

0.6% with P179

P. intermedia, 

differ by

2.6-3.1%

Hughet Spring, 

Harney-Malheur 

Lakes, Harney Co., 

OR

P. hendersoni/

robusta in 

GenBank

MA/

hendersoni2/

P5

AY197595/

AY379432-33 2 0.00%

0.5% with P171, 

Ida, P181, 0.5-

0.6% with P179

P. intermedia, 

differ by

2.6-3.1%

Spring at South

Fork (Malheur

River) Reservoir, 

middle Snake River 

basin, Malheur 

Co., OR

P. hendersoni/

robusta in 

GenBank

D33/

herdersoni3

AY379434-35, 

AY426348-49 4 0.2-0.3% 0.5-0.8% with P179

P. intermedia, 

differ by

2.6-3.2%

XL Spring, Abert 

Lake basin (Great 

Basin), Lake Co., 

OR

P. hendersoni/

robusta in 

GenBank

P171/

idahoensis1 AY379424-25 2 0.00% 0.5-0.6% with D33

P. intermedia, 

differ by

2.6-3.0%

Bruneau Arm of 

CJ Strike Reservoir,

(Bruneau) river mile 

3.8, Owyhee Co., ID

P179/

idahoensis2 AY379426-27 2 0.20%

0.3-0.8% with 

D30, 0.5-0.8% 

with D33

P. intermedia, 

differ by

2.6-3.2%

Snake River river 

mile 538.1, Glenns 

Ferry, Snake River 

basin, Elmore 

Co., ID

Ida AY426346-47 2 0.00%

0.3-0.6% with 

D30, 0.5-0.6% 

with D33

P. intermedia, 

differ by

2.6-3.0%

Snake River, river 

mile 446.8, Canyon 

Co., ID

P. idahoensis

in GenBank

P181/

Idahoensis3 AY379428-29 2 0.00%

0.3-0.6% with 

D30, 0.5-0.6% 

with D33

P. intermedia, 

differ by

2.6-3.0%

Snake River, river 

mile 365.9, 366.3, 

367.2, Malheur 

Co., OR

P178/

robusta AY379436-37 2 0.00% 0.3-0.5% with P179

P. intermedia, 

differ by

2.7-3.2%

Spring, tributary to 

Polecat Creek, 

Teton Co., WY

P47B AF520949 1 0.00% 0.5-0.7% with 

P179

P. intermedia, 

differ by

3.0-3.5%

Polecat Creek, west 

of Flagg Ranch, 

Teton Co., WY

D30/spA1 AY379438-39 2 0.30% 0.3-0.8% with 

P179

P. intermedia, 

differ by

2.7-3.2%

Columbia River, 

East Mayer State 

Park, Wasco 

Co., OR

P49/spA2 AY379440-41 2 0.00%

0.5% with P171, 

Ida, P181, 0.5-

0.6% with P179

P. intermedia, 

differ by

2.8-3.4%

Columbia River, 

Celilo State Park, 

Wasco Co., OR

Pyrgulopsis 

rupinicola

SS1/D3, 

SS2/D3

AY197590 

(N=3), 

AY197591 

(N=1)

4 0-0.5%

P. archimedis, 

differ by

2.6-3.2%,

P. sp. A, differ

by 2.4-3.4% 

Sucker Springs 

Creek, Shasta Co., 

CA

HYDROBIIDAE
Technical Note 449 115

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

sadai P141A AY627945 1 0.00%

P. kolobensis, 

differ by

6.2-7.8%, 

P. pilsbryana, 

differ by

6.2-6.5%,

P. sp. C, differ

by 6.2-6.3%

Spring, Moss Creek, 

Reese River Valley, 

Lander Co., NV

Pyrgulopsis 

sanchezi M2 AY367430-32 (M2A-O) 15 0.00%

2.0-2.1%

with M25,

1.7-2.1%

with M26,

1.8-2.1%

with SS

P. coloradensis, 

differ by 4.1%

Grapevine Springs, 

Death Valley, Inyo 

Co., CA

P. micrococcus

in GenBank

M8

AY367440 

(M8A), 

DQ364001 

(N=9, P197A, 

M8AA-AH)

10 0-0.2% 2.1-2.4%

with SS

P. arizonae,

differ by

4.1-5.6%,

P. coloradensis, 

differ by

4.3-4.4%

Purgatory Spring, 

Ash Meadows, Nye 

Co., NV

P. micrococcus

in GenBank

M25

AY367469 

(M25A), 

AY367470-71 

(N=8, M25B-C, 

AB-AD, AG-AI)

9 0.20% 2.0-2.1%

with M2

P. coloradensis, 

differ by

4.1-4.3%

Spring north of 

Tecopa Hot

Springs, Amargosa 

River basin, Inyo 

Co., CA

P. micrococcus

in GenBank

M26

AY367472 

(N=5, M26A, 

AD-AE, AG, AI), 

AY367473 

(N=3, M26B, 

M26C,

M26AC),

KF559187 

(N=5, M26AAAB, AF, AH, AJ),

KF559188 

(M26AK)

14 0-0.5%

1.8-2.2%

with SS,

1.7-2.1%

with M2

P. arizonae,

differ by

4.7-6.2%,

P. coloradensis, 

differ by

4.6-5.0%

Shoshone Spring, 

Amargosa River 

basin, Inyo Co., CA

P. micrococcus

in GenBank

M51 KF559184 

(M51A-B) 2 0.00% 2.1-2.2%with SS

P. coloradensis, 

differ by 3.8%

Spring ca. 100 m 

north of Collins 

Ranch, Ash 

Meadows, Nye Co., 

NV

M53 KF559185 

(M53A-C) 3 0.00% 2.1-2.2%with SS

P. coloradensis, 

differ by 4.1%

Spring south of 

Rogers Spring, 

Ash Meadows, Nye 

Co., NV

M57 KF559186 

(M57A-B) 2 0.00% 2.1-2.2%with SS

P. coloradensis, 

differ by 3.8%

Spring east of 

Crystal Reservoir, 

Ash Meadows, Nye 

Co., NV

SS

AY367480 

(SS1), 

DQ364003 

(SS2)

2 0.50% 2.1-2.4%

with M8

P. arizonae,

differ by

4.5-6.1%,

P. coloradensis, 

differ by 4.5%

Saratoga Spring, 

Death Valley, San 

Bernardino Co., CA

Pyrgulopsis

sp. L HPL2003 in 

GenBank

Pyrgulopsis 

sathos P62A DQ364013 1 0.00% P. hubbsi,differ by 1.0%

Flag Springs

(north), White River 

Valley, Nye Co., NV

HYDROBIIDAE
116 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

saxatilis P95A AY627934 1 0.00%

P. eremica,

differ by

6.6-9.4%,

P. lentiglans,

differ by 6.7%

Warm Springs, 

Snake Valley, 

Millard Co., UT

Pyrgulopsis 

serrata P69A EU700464 1 0.00% P. marcida,differ by 1.4%

Indian Ranch

Spring, Steptoe 

Valley, White Pine 

Co., NV

Pyrgulopsis 

similis G4

KC571291 

(N=4), 

KC571292 

(N=1),

KC571293 

(N=1),

KC571294 

(N=1)

7 0-1.2% 0.5-1.5%

with G6

P. marilynae,

differ by

4.0-5.1%

Seepage along 

Taylor Creek, ca. 

0.32 km south, 

0.93 km west of 

Wall Lake Dam 

(below Wall Lake), 

Catron Co., NM

P. gilae in 

GenBank

G5 KC571295 7 0.00% 1.1% with G7, 

G13, Pgil6

P. marilynae, 

differ by

3.8-4.1%

Hillside seep, 

1.61 km north, 

0.97 km east of 

Burnt Corral

Canyon, Catron

Co., NM

P. gilae in 

GenBank

G6

KC571296 

(N=2), 

KC571297 

(N=6),

KC571298 

(N=1)

9 0-0.6%

1.2-1.5%

with G7, G13, 

Pgil6

P. marilynae,

differ by

3.8-4.3%

Spring along

Beaver Creek, ca. 

0.29 km north, 

0.40 km west 

of Taylor Creek 

confluence, Catron 

Co., NM

P. gilae in 

GenBank

G7 KC571292 10 0.00%

1.2-1.5%

with G6,

1.2-1.4%

with G9

P. marilynae,

differ by

4.8-5.1%

Seepage along 

Taylor Creek, 50 m 

west of Whitetail 

Canyon, Catron Co., 

NM

P. gilae in 

GenBank

G9

KC571303 

(N=1), 

KC571304 

(N=8)

9 0-0.2%

1.2-1.4%

with G7, G13, 

Pgil6

P. marilynae,

differ by

4.0-4.5%

Fall Spring, 

1.61 km north, 

0.56 km east of 

Burnt Corral

Canyon, Catron

Co., NM

P. gilae in 

GenBank

G10 KC571304 4 0.00% 1.2% with G7, 

G13, Pgil6

P. marilynae,

differ by

4.0-4.3%

Fall Spring, 

1.61 km north, 

0.56 km east of 

Burnt Corral 

Canyon, Catron

Co., NM

P. gilae in 

GenBank

G13 KC571292 8 0.00%

1.2-1.5%

with G6,

1.2-1.4%

with G9

P. marilynae,

differ by

4.8-5.1%

Spring along Taylor 

Creek, 0.81 km 

north, 1.13 km east 

of Wall Lake Dam, 

Catron Co., NM

P. gilae in 

GenBank

Pgil6 AY485574 14 0.00%

1.2-1.5%

with G6,

1.2-1.4%

with G9

P. marilynae,

differ by

4.8-5.1%

Wall

Hurt (2004), 

P. gilae in 

GenBank

HYDROBIIDAE
Technical Note 449 117

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

simplex P151A AY627949 1 0.00%

0%

with

Psim1

P. glandulosa, 

differ by

2.6-2.7%,

P. perforata,

differ by

2.6-2.7%

Spring near 

Strawberry, Verde 

River drainage, Gila 

Co., AZ

Psim1 AY485558 31 0.00%

0%

with

P151A

P. glandulosa, 

differ by 2.6-

2.7%,

P. perforata, 

differ by

2.6-2.7%

Strawberry Hurt (2004)

Pyrgulopsis

sola BRS1 AY627957 1 0.00%

0.2-0.4%

with

Psol1/2/3

P. glandulosa, 

differ by 3.3%, 

P. intermdeia, 

differ by

3.3-3.9%

Brown Springs, 

Verde River

drainage, Yavapai 

Co., AZ

Psol1, 

Psol2, 

Psol3

AY485554 

(N=19),

AY485555 

(N=6),

AY485556 

(N=1)

26 0.2-0.4% 0.2-0.4%

with BRS

P. glandulosa, 

differ by

3.5-3.8%, 

P. intermedia, 

differ by

3.5-4.3%

Brown

Hurt (2004), 

P. solus in 

GenBank

Pyrgulopsis 

stearnsiana

16/

stearnsianaA/

WC

AF520925 2 0.00% 3.6-3.7%

with Pst8

P. diablensis, 

differ by 1.1%

Springs, Wildcat 

Canyon, El

Sobrante, Contra 

Costa Co., CA

18/PC/P26

AY367489 

(P26A, N=1), 

GQ275092 

(P26AA, N=1)

2 0.20% 3.2-3.7%

with Pst8

P. diablensis, 

differ by

1.6-1.8%

Partington Creek, 

Central California 

coastal drainage, 

Monterey Co., CA

19/CC/P28 AY367490 3 0.00% 3.2-3.4%

with Pst8

P. diablensis, 

differ by 1.1%

Stream in Colson 

Canyon, Central 

California coastal 

drainage, Santa 

Barbara Co., CA

Pst1 KX398371 4 0.00% 3.0-3.2%

with Pst8

P. diablensis, 

differ by 1.7%

Spring run, Glen 

Canyon Park, San 

Francisco Co., CA

Pst2 KX398372 2 0.00% 3.0-3.2%

with Pst8

P. diablensis, 

differ by 1.7%

Seep under the 

connector between 

Hwy 280 and Hwy 

101, San Francisco 

Co., CA

Pst3

KX398373 

(N=4), 

KX398374 

(N=1)

5 0-0.2% 2.7-3.0%

with Pst8

P. diablensis, 

differ by

0.9-1.1%

Spring north of 

Laguna Honda, San 

Francisco Co., CA

Pst4 KX398375 4 0.00% 3.6-3.8%

with Pst8

P. diablensis, 

differ by 2.0%

Lobos Creek, San 

Francisco Co., CA

Pst5 KX398376 4 0.00% 3.0-3.2%

with Pst8

P. diablensis, 

differ by 1.7%

Spring along trail 

northeast of Ft. 

Miley, San 

Francisco Co., CA

Pst6 KX398377 4 0.00% 2.8%

with WC

P. diablensis, 

differ by 0.6%

Spring north of 

Geary Road, east 

of Calaveras 

Reservoir,

Alameda Co., CA

HYDROBIIDAE
118 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

stearnsiana

(continued)

Pst7

KX398378 

(N=3), 

KX398379 

(N=1)

4 0-1.2% 2.7-3.2%

with Pst8

P. diablensis, 

differ by

0.6-1.1%

Spring north of 

Geary Road, north

of Calaveras 

Reservoir, Alameda 

Co., CA

Pst8

KX398380 

(N=3), 

KX398381 

(N=1)

4 0-0.2%

3.6-3.8%

with Pst4,

3.0-3.8%

with Pst13,

3.3-3.8%

with Pst15,

3.6-3.8%

with WC,

3.2-3.7%

with PC

P. diablensis, 

differ by

2.7-2.8%

Spring south 

of Geary Road, 

northeast of 

Calaveras

Reservoir, Alameda 

Co., CA

Pst9 KX398382 5 0.00%

3.5% with Pst4, 

2.9-3.5%

with Pst13,

3.2-3.5%

with Pst15,

3.4% with WC, 

3.0-3.4%

with PC

P. diablensis, 

differ by 2.5%

Spring along 

Weller Road, west

of Calaveras 

Reservoir, Santa 

Clara Co., CA

Pst10

KX398383 

(N=1), 

KX398384 

(N=1)

2 0.60% 2.4-2.9% with 

Pst8

P. diablensis, 

differ by

1.1-1.4%

Temescal Creek, 

Alameda Co., CA

Pst11 KX398385 4 0.00% 2.9-3.0%

with Pst8

P. diablensis, 

differ by 0.9%

Spring tributary to 

northeast corner 

of Rodeo Lagoon, 

Marin Co., CA

Pst12

KX398386 

(N=3), 

KX398387 

(N=1)

4 0-2.4%

2.6-2.9% with 

Pst8, 2.5-3.0% 

with WC

P. diablensis, 

differ by

1.0-1.1%

McWay Creek, 

Monterey Co., CA

Pst13

KX398388 

(N=1), 

KX398389 

(N=3)

4 0-0.9%

3.0-3.8% with 

Pst8, 2.9-3.5% 

with Pst9 

P. diablensis, 

differ by

0.8-1.4%

Spring at Seven 

Stairs, Monterey 

Co., CA

Pst14 KX398390 4 0.00% 3.0-3.2%

with Pst8

P. diablensis, 

differ by 0.8%

Stream south of 

Salmon Creek, 

Monterey Co., CA

Pst15

KX398391 

(N=2), 

KX398392 

(N=2)

4 0-0.3%

3.3-3.8% with 

Pst8, 3.2-3.5% 

with Pst9

P. diablensis, 

differ by

1.1-1.4%

Springs southeast 

of Leffingwell

Creek, San Luis 

Obispo Co., CA

Pst16

KX398393 

(N=2), 

KX398394 

(N=2)

4 0-1.5%

2.6-3.2% with 

Pst8, 2.4-2.9% 

with Pst9, 2.6-

3.0% with WC

P. diablensis, 

differ by

1.1-1.4%

San Miguelito

Creek, Santa 

Barbara Co., CA

Pst17 KX398395 4 0.00% 3.2-3.3%

with Pst8

P. giulianii, differ 

by 1.6-1.7%

Stream at Hilarita, 

Marin Co., CA

Pst18 KX398396 4 0.00% 3.2-3.3%

with Pst8

P. diablensis, 

differ by 1.3%

Seep south of 

Stinson Beach, 

Marin Co., CA

Pyrgulopsis 

sterilis AY367491 1 0.00% P. marcida,differ by 1.4%

Spring, Hunts 

Canyon Ranch, 

Ralston Valley, Nye 

Co., NV

HYDROBIIDAE
Technical Note 449 119

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

sublata P111A AY627938 1 0.00% P. leporina,differ by 1.9%

Wambolt Springs, 

Lake Valley, Lincoln 

Co., NV

Pyrgulopsis 

sulcata P66A AY627929 1 0.00% P. neritella,differ by 3.2%

Spring northwest 

of Clark Spring, 

Steptoe Valley, 

White Pine Co., NV

Pyrgulopsis

taylori P27

AY627923 

(P27A, N=1), 

GQ275095 

(P27AA, N=3)

4 0-0.3%

P. bacchus,

differ by

4.0-4.5%, 

P. milleri,

differ by

3.7-4.0%,

P. montana,

differ by

3.7-4.0%,

P. turbatrix,

differ by

3.0-6.9%

Spring tributary to 

San Luis Obispo 

Creek, California 

central coastal 

drainage, San Luis 

Obispo Co., CA

Pyrgulopsis 

thermalis P165A AY627953 1 0.00% 2.7-3.2% with Pthe3/4/5/6/7

P. lockensis,

differ by 7.3%

Hot Spring, Gila 

River drainage,

Grant Co., NM

Pthe1,

Pthe2

AY485575 

(N=30),

AY485576 

(N=1)

31 0-0.3% 2.1-2.7% with 

Pthe3/4/5/6/7

P. deaconi, differ 

by 7.1-7.7%,

P. lockensis, 

differ by 7.5-

7.8%,

P. trivialis, differ 

by 7.3-7.8%,

Gila I Hurt (2004)

Pthe3, 

Pthe4, 

Pthe5, 

Pthe6, 

Pthe7

AY485577 

(N=1),

AY485578 

(N=1),

AY485579 

(N=1),

AY485580 

(N=1),

AY485581 

(N=1)

5 0.2-0.9%

2.7-3.2%

with P165,

2.7-3.6% with 

Pthe8/9/10/11

P. carinifera,

differ by

6.4-7.0%

Gila II Hurt (2004)

Pthe8, 

Pthe9,

Pthe10,

Pthe11

AY485582 

(N=?),

AY485583 

(N=?), 

AY485584 

(N=?), 

AY485585 

(N=?)

31 0.0-0.8% 2.7-3.6% with 

Pthe3/4/5/6/7

P. kolobensis, 

differ by,

7.1-8.9%,

P. lockensis,

differ by

7.3-8.3%,

P. trivialis, differ 

by 7.7-9.4%,

P. wongi, differ

by 7.1-10.5%

Gila III

Hurt (2004), 

listed 

frequency 

greater than 

1.0 (1.582)

Pyrgulopsis 

thompsoni Ptho1 AY485588 32 0.00% 2.5% withPtho5

P. conica, differ

by 3.2%, 

P. glandulosa, 

differ by

3.2-3.3%,

P. morrisoni,

differ by

2.9-4.6%

Bear Hurt (2004)

Ptho1,

Ptho2

AY485588 

(N=29), 

AY485589 

(N=1)

30 0.00% 2.5% with

Ptho5

P. conica, differ

by 3.2%,

P. glandulosa, 

differ by

3.2-3.3%,

P. morrisoni,

differ by

2.9-4.6%

Canelo Hills

Hurt (2004), 

No variation 

between 

Ptho1 and 

Ptho2 in 

GenBank

HYDROBIIDAE
120 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

thompsoni

(continued)

Ptho3,

Ptho4

AY485590 

(N=28), 

AY485591 

(N=2)

30 0.0-1.9% 2.7-3.8%

with Ptho5

P. conica, differ

by 2.7-3.8%,

P. glandulosa, 

differ by

3.0-3.8%,

P. milleri, differ

by 3.0-3.4%,

P. morrisoni,

differ by

2.9-4.6%

Cottonwood Hurt (2004)

Ptho5 AY485592 32 0.00%

2.7-3.8%

with Pth3/4,

2.9-3.2%

with Ptho8/9

P. conica, differ

by 2.7%, 

P. milleri, differ

by 2.7%

McClure Hurt (2004)

Ptho5 AY485592 36 0.00%

2.7-3.8%

with Pth3/4, 

2.9-3.2%

with Ptho8/9

P. conica, differ

by 2.7%, 

P. milleri, differ

by 2.7%

Garden Hurt (2004)

Ptho5 AY485592 30 0.00%

2.7-3.8%

with Pth3/4, 

2.9-3.2%

with Ptho8/9

P. conica, differ

by 2.7%,

P. milleri, differ

by 2.7%

Cave Hurt (2004)

Ptho8,

Ptho9

AY485593 

(N=28), 

AY485594 

(N=2)

30 0.0-0.3% 2.9-3.2%

with Ptho5

P. conica, differ

by 3.2-3.8%,

P. glandulosa, 

differ by

3.2-3.8%,

P. hualapaiensis, 

differ by

2.8-3.4%,

P. milleri, differ

by 3.0-3.4%,

P. montana,

differ by

2.8-3.7%

Monkey Hurt (2004)

Ptho1,

Ptho5

AY485588 

(N=34), 

AY485592 

(N=2)

36 0.0-2.5%

0.0-2.5%

with Ptho1, 

Ptho1/2, Ptho5,

0.2-3.8%

with Ptho3/4, 

0.9-3.2%

with Ptho8/9

P. conica, differ

by 2.7-3.2%,

P. glandulosa, 

differ by

3.2-3.4%,

P. milleri, differ

by 2.7-3.4%,

P. morrisoni,

differ by

2.9-5.0%

Peterson Ranch Hurt (2004)

Ptho1 AY485588 32 0.00% 2.5%

with Ptho5

P. conica, differ

by 3.2%, 

P. glandulosa, 

differ by

3.2-3.3%,

P. morrisoni,

differ by

2.9-4.6%

Sawmill Hurt (2004)

Pyrgulopsis 

torrida 21/P250 GQ275094 4 0.00%

P. glandulosa, 

differ by

2.6-2.7%

Stream, Little 

Sycamore Canyon, 

Southern California 

coastal drainage, 

Ventura Co., CA

P. sp. 

USNM1120443 

isolate 

P250AA, 

described as 

P. torrida in 

Hershler et al. 

(2016b)

HYDROBIIDAE
Technical Note 449 121

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

transversa P112A EU700470 1 0.00%

P. kolobensis, 

differ by

0.6-2.7%,

P. nonaria, differ 

by 1.1%,

P. pilsbryana, 

differ by

1.0-1.1%,

P. sp. C, differ

by 1.1%,

P. sp. D, differ 

by 1%,

P. sp. H, differ

by 1.1%

Sixmile Springs,

Old River Bed, 

Tooele Co., UT

Pyrgulopsis 

trivialis P119A AY627941 1 0.00% 0.3% with Ptri1

P. kolobensis, 

differ by

3.1-4.4%,

P. nonaria, differ 

by 3.6%,

P. pilsbryana, 

differ by 3.6%, 

P. variegata,

differ by 3.7%,

P. sp. C, differ

by 3.6-3.7%,

P. sp. D, differ

by 3.6%, 

P. sp. H, differ

by 3.6%

Spring, Three Forks, 

Gila River drainage, 

Apache Co., AZ

Ptri1 AY485559 

(N=31) 31 0.00% 0.5% with Ptri2

P. kolobensis, 

differ by

3.3-4.8%,

P. nonaria,

differ 3.8%,

P. pilsbryana, 

differ by 3.9%,

P. variegata,

differ by 4.0%,

P. sp. C, differ

by 3.8-4.0%,

P. sp. D, differ

by 3.9%,

P. sp. H, differ

by 3.9%

Boneyard Bog Hurt (2004)

Ptri2 AY485560 

(N=31) 31 0.00% 0.5% with Ptri1

P. kolobensis, 

differ by

3.2-4.7%, 

P. nonaria,

differ by 3.6%,

P. pilsbryana, 

differ by 3.7%,

P. variegata,

differ by 3.9%,

P. sp. C, differ

by 3.6-3.9%,

P. sp. D,

differ by 3.7%,

P. sp. H,

differ by 3.7%

Three Forks Hurt (2004)

HYDROBIIDAE
122 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

turbatrix M9

AY367441 

(M9A), 

AY367442 

(M9B)

2 0.00%

1.3%

with M19,

P11

P. imperialis,

differ by 5.2%

Stream below 

Darwin Falls, 

Panamint Valley, 

Inyo Co., CA

P. micrococcus

in GenBank

M10

AY367443 

(M10A), 

AY367444 

(M10B)

2 0.00%

1.4%

with M19,

P11

P. imperialis,

differ by 5.2%

China Garden

Spring, Panamint 

Valley, Inyo Co., CA

P. micrococcus

in GenBank

M11 AY367445 1 0.00%

1.4%

with M19, P11, 

P85, P196

P. imperialis,

differ by 5.2%

Spring above 

Darwin Falls, 

Panamint Valley, 

Inyo Co., CA

P. micrococcus

in GenBank

M12

AY367446 

(M12A), 

AY367447 

(M12B)

2 0.00%

1.4%

with P11,

P196

P. imperialis,

differ by 5.0%

Lower spring, 

Johnson Canyon, 

Death Valley, Inyo 

Co., CA

P. micrococcus

in GenBank

M13

AY367448/

AY367450-52 

(N=4, 

M13A, C-E), 

AY367449 

(M13B),

AY367453 

(M13F)

6 0-0.3%

1.4-1.6%

with P11,

P196

P. imperialis,

differ by

5.0-5.2%

Stream, Hanaupah 

Canyon, Death 

Valley, Inyo Co., CA

P. micrococcus

in GenBank

M14

AY367454 

(M14A), 

AY367455 

(M14B)

2 0.20%

1.4-1.6%

with P11,

P196

P. imperialis,

differ by

5.0-5.2%

Stream, Jail

Canyon, Panamint 

Valley, Inyo Co., CA

P. micrococcus

in GenBank

M15

AY367456 

(M15A, M15M), 

AY367457 

(M15B)

3 0.20%

1.8-1.9%

with P11,

P196

P. imperialis,

differ by

5.2-5.3%

Stream, Hall

Canyon, Panamint 

Valley, Inyo Co., CA

P. micrococcus

in GenBank

M16 AY367458 

(M16A, M16M) 2 0.00%

2.1%

with P11,

P86, P196

P. imperialis,

differ by 5.5%

Spring, Snow 

Canyon, Panamint 

Valley, Inyo Co., CA

P. micrococcus

in GenBank

M17 AY367459 1 0.00%

2.1%

with P11,

P196

P. imperialis,

differ by 5.5%

Cottonwood Spring, 

Wildrose Canyon, 

Panamint Valley, 

Inyo Co., CA

P. micrococcus

in GenBank

M18 AY367460 

(M18A, M18O) 2 0.00%

2.1%

with P11,

P196

P. imperialis,

differ by

5.3-5.5%

Saline Marsh,

Saline Valley, Inyo 

Co., CA

P. micrococcus

in GenBank

M19 AY367461 

(M19A, M19O) 2 0.00% 1.8%with M16

P. imperialis,

differ by 5.5%

Cushenbury Springs, 

San Bernardino 

Mountains, San 

Bernardino Co., CA

P. micrococcus

in GenBank

M20 AY367462 

(M20A, M20N) 2 0.00%

1.6%

with P11,

P196

P. imperialis,

differ by 5.2%

Springs at Big Bear 

Ranger Station, 

San Bernardino 

Mountains, San 

Bernardino Co., CA

P. micrococcus

in GenBank

M21

AY367463 

(M21A, M21M), 

AY367464 

(M21B)

3 0.00% 1.6%

with M16

P. imperialis,

differ by 5.3%

Springs, Mill 

Creek Canyon, 

San Bernardino 

Mountains, San 

Bernardino Co., CA

P. micrococcus

in GenBank

M22

AY367465/

AY367466 

(N=2, M22A-B), 

AY367467 

(M22C)

3 0.20%

1.4-1.6%

with P11,

P196

P. imperialis,

differ by

5.0-5.2%

Spring, Knight 

Canyon, Panamint 

Valley, Inyo Co., CA

P. micrococcus

in GenBank

HYDROBIIDAE
Technical Note 449 123

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

turbatrix

(continued)

M24 AY367468 

(M24A, M24N) 2 0.00% 1.4% with P11, P196

P. imperialis,

differ by 5.0%

Tennessee Spring, 

Panamint Valley, 

Inyo Co., CA

P. micrococcus

in GenBank

M27 AY367474 1 0.00% 1.4% with P11, 

P196

P. imperialis,

differ by 5.0%

Stream below 

Limekiln Spring, 

Panamint Valley, 

Inyo Co., CA

P. micrococcus

in GenBank

M28 AY367475 1 0.00% 1.4% with P11, 

P196

P. imperialis,

differ by 5.0%

Brewery Spring, 

Surprise Canyon, 

Panamint Valley, 

Inyo Co., CA

P. micrococcus

in GenBank

P11 AF520936 

(P11A) 1 0.00% 2.1% with M16, M17, M18

P. imperialis,

differ by 5. 2%

Horseshutem 

Springs, Pahrump 

Valley, Nye Co., NV

P84 DQ364014 

(P84A) 1 0.00% 1.9% with M16, M17, M18

P. imperialis,

differ by 5.0%

Cold Creek Spring, 

Indian Springs 

Valley, Clark Co., NV

P86 DQ364015 

(P86A) 1 0.00% 2.1% with M16 P. imperialis,differ by 5.2%

Spring, Lost Creek 

Canyon, Las Vegas 

Wash, Clark Co., NV

P196

DQ363999 

(P196A), 

DQ364000 

(P196D)

2 0.00% 2.1% with M16, 

M17, M18

P. imperialis,

differ by

3.9-4.2%

Grapevine Springs, 

Amargosa River 

basin, Nye Co., NV

Pyrgulopsis 

umbilicata SM3 EF119093-94 4 0-0.2% 0.3-0.5% with SM37

P. limaria, differ 

by 0.2-1.1%

Spring, northern 

Soldier Meadow, 

Humboldt Co., NV

SM14 EF119101 5 0.00% 0.3% with SM37 P. limaria, differ 

by 0.2-0.9%

Spring, northern 

Soldier Meadow, 

Humboldt Co., NV

SM16 EF119102 5 0.00% 0.3% with SM37 P. limaria, differ 

by 0.2-0.9%

Spring ca. 40 m 

downflow from 

source, northern 

Soldier Meadow, 

Humboldt Co., NV

SM19 EF119103 5 0.00% 0.5% with SM37 P. limaria, differ 

by 0.3-1.1%

Spring, northern 

Soldier Meadow, 

Humboldt Co., NV

SM20 EF119104 4 0.00% 0.3% with SM37 P. limaria, differ 

by 0.2-0.9%

Spring, southern 

Soldier Meadow, 

Humboldt Co., NV

SM21 EF119105 4 0.00% 0.3% with SM37 P. limaria, differ 

by 0.2-0.9%

Spring 5 m 

downflow from 

SM20, southern 

Soldier Meadow, 

Humboldt Co., NV

SM37 EF119127 2 0.00%

0.3-0.5% with 

SM3, 0.5% with 

SM19

P. limaria, differ 

by 0.5-1.2%

Spring, central 

Soldier Meadow, 

Humboldt Co., NV

Pyrgulopsis 

variegata P106A AY627937 1 0.00%

P. kolobensis, 

differ by

1.1-2.1%, 

P. sp. B, differ

by 1.1%

Spring south of 

South Patterson 

Spring, Pilot Valley, 

Box Elder Co., UT

HYDROBIIDAE
124 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis 

varneri SM8 EF119080 3 0.00%

0.0-0.3%

with SM13,

0.2%

with SM15, 

SM25, SM28

P. militaris,

differ by

2.1-2.4%

Spring, southeast 

Soldier Meadow, 

150 m downflow 

from source, 

Humboldt Co., NV

SM11 EF119081 8 0.00%

0.0-0.3%

with SM13,

0.2% with SM15, 

SM25, SM28

P. militaris,

differ by

2.1-2.4%

Spring complex 

(two springs), 

southeast Soldier 

Meadow, Humboldt 

Co., NV

SM13 EF119082-84 4 0-0.5%

0.2-0.5%

with SM15, 

SM25, SM28

P. militaris,

differ by

2.1-2.7%

Spring brook 

downflow from 

SM11, Humboldt 

Co., NV

SM15 EF119085 5 0.00%

0.2-0.5%

with SM13,

0.3% with

SM25, SM28

P. militaris,

differ by

2.0-2.4%

Spring, northern 

Soldier Meadow, 

Humboldt Co., NV

SM25 EF119086 5 0.00%

0.2-0.5% with 

SM13, 0.3%

with SM15

P. militaris,

differ by

2.3-2.6%

Satellite Spring 

complex, spring "E," 

Humboldt Co., NV

SM28 EF119087 4 0.00%

0.2-0.5%

with SM13,

0.3% with SM15

P. militaris,

differ by

2.3-2.6%

Satellite Spring 

complex, spring "G," 

Humboldt Co., NV

Pyrgulopsis 

ventricosa P87 AY426365-66 2 0.00%

P. kolobensis, 

differ by

4.5-5.5%,

P. marcida,

differ by 4.5%,

P. plicata,

differ by 4.5%

Spring, Seigler 

Canyon, Clear Lake 

Basin, Lake Co., CA

Pyrgulopsis 

villacampae P83A AY627933 1 0.00% P. lockensis,differ by 2.6%

Little Warm Spring, 

Railroad Valley, Nye 

Co., NV

Pyrgulopsis 

vinyardi SM38A EU700482 1 0.00%

P. gibba, differ

by 0.5-1.1%, 

P. plicata, differ

by 4.6%

Unnamed spring, 

Squaw Valley, Elko 

Co., NV

Pyrgulopsis

wongi AF520923 1 0.00% 7.1%with W3

P. kolobensis, 

differ by

3.4-4.6%, 

P. marcida,

differ by 3.8%,

P. nonaria,

differ by 3.9%,

P. pilsbryana, 

differ by

3.8-3.9%,

P. transversa, 

differ by 3.8%, 

P. sp. D, differ

by 3.8%, 

P. sp. H, differ

by 3.9%

Spring, lower Pine 

Creek Canyon, Inyo 

Co., CA

P185

AY627956 

(P185A,

P185C)

4 0.00% 6.7%

with W3

P. kolobensis, 

differ by

3.3-4.5%,

P. marcida,

differ by 3.7%

Spring, Birchim 

Canyon, Inyo Co.,

CA

HYDROBIIDAE
Technical Note 449 125

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

wongi

(continued)

W01 DQ251052(HI) 8 0.00% 6.5% with W3

P. kolobensis, 

differ by

3.5-4.7%, 

P. marcida,

differ by 3.9%

Spring near Conway 

Summit, Mono 

Co., CA

W02

DQ251053 

(HXXI, N=5),

DQ251054 

(HXXII, N=1),

DQ251055 

(HXXIII, N=1)

7 0-0.3% 7.1-7.3%

with W3

P. kolobensis, 

differ by

4.4-5.6%,

P. marcida, differ 

by 4.5-4.7%

Springs at Little 

Lake, Inyo Co., CA

W03

DQ251075 

(W3A, N=3), 

DQ251076 

(W3C, N=2)

5 0-0.2%

7.1-7.3%

with W2,

7.3% with W28

P. rupinicola,

differ by

4.8-5.1%

Doud Springs, 

Douglas Co., NV

W04 DQ251056

(HII) 5 0.00% 6.4% with W3

P. kolobensis, 

differ by

3.6-4.8%,

P. marcida,

differ by 4.0%

River Spring, Mono 

Co., CA

W05

DQ251057

(HIII, N=2), 

DQ251058

(HIV, N=1),

DQ251059

(HV, N=1)

4 0-0.3% 6.5-6.7%

with W3

P. kolobensis, 

differ by

3.5-4.8%,

P. marcida, differ 

by 3.9-4.0%

Spring in Clark 

Canyon, Mono Co., 

CA

W06 HVI 5 0.00% 6.7% with W3

P. kolobensis, 

differ by

3.3-4.5%,

P. marcida,

differ by 3.7%

Springs in Owens 

River gorge, Mono 

Co., CA

Haplotype 

VI is not in 

GenBank

W07 HVI 7 0.00% 6.7% with W3

P. kolobensis, 

differ by

3.3-4.5%,

P. marcida,

differ by 3.7%

Spring along Pine 

Creek, Inyo Co., CA

Haplotype 

VI is not in 

GenBank

W08 DQ251060 

(HVII) 4 0.00% 6.5% with W3

P. kolobensis, 

differ by

3.5-4.7%,

P. marcida,

differ by 3.9%

Spring along 

Clearwater Creek, 

Mono Co., CA

W09 DQ251060 

(HVII) 3 0.00% 6.5% with W3

P. kolobensis, 

differ by

3.5-4.7%,

P. marcida,

differ by 3.9%

Spring along

Rough Creek,

Mono Co., CA

W10 DQ251056

(HII) 4 0.00% 6.4% with W3

P. kolobensis, 

differ by

3.6-4.8%, 

P. marcida,

differ by 4.0%

Huntoon Spring, 

Mineral Co., NV

W11

DQ251056

(HII, N=3),

DQ251061 

(HVIII, N=1)

4 0-0.2% 6.4-6.5%

with W3

P. kolobensis, 

differ by

3.6-5.0%,

P. marcida, differ 

by 4.0-4.2%

Jacks Spring, 

Mineral Co., NV

W12

DQ251056

(HII, N=1), 

DQ251062

(HIX, N=3)

4 0-0.2% 6.4-6.5%

with W3

P. kolobensis, 

differ by

3.6-5.0%,

P. marcida, differ 

by 4.0-4.2%

Company Spring, 

Mineral Co., NV

HYDROBIIDAE
126 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

wongi

(continued)

W13 DQ251063

(HX) 4 0.00% 6.5%with W3

P. kolobensis, 

differ by

3.5-4.7%,

P. marcida,

differ by 3.9%

Spring east of 

McNett Ranch, 

Esmeralda Co., NV

W14 DQ251064 

(HXI) 4 0.00% 6.4%with W3

P. kolobensis, 

differ by

3.3-4.4%, 

P. marcida,

differ by 3.7%

Cave Spring, 

Esmeralda Co., NV

W15 DQ251064 

(HXI) 8 0.00% 6.4%with W3

P. kolobensis, 

differ by

3.3-4.4%, 

P. marcida,

differ by 3.7%

Corral Springs, Inyo 

Co., CA

W16 DQ251065 

(HXII) 6 0.00% 6.5%with W3

P. kolobensis, 

differ by

3.5-4.5%, 

P. marcida,

differ by 3.9%

Antelope Springs, 

Inyo Co., CA

W17

DQ251056

(HII, N=5), 

DQ251063

(HX, N=1)

6 0-0.5% 6.4-6.5%

with W3

P. kolobensis, 

differ by

3.5-4.8%,

P. marcida, differ 

by 3.9-4.0%

Springs at Pizona, 

Mono Co., CA

W18

DQ251066 

(HXIII, N=3), 

DQ251067 

(HXIV, N=1)

4 0-0.2% 6.5%

with W3

P. kolobensis, 

differ by

3.5-4.8%,

P. marcida, differ 

by 3.9-4.0%

Layton Springs, 

Mono Co., CA

W19

DQ251060 

(HVII, N=2), 

DQ251068 

(HXV, N=1),

DQ251069 

(HXVI, N=1)

4 0-0.3% 6.5-6.7%

with W3

P. kolobensis, 

differ by

3.5-4.8%,

P. marcida, differ 

by 3.9-4.0%

Springs along 

northeast side of 

Blind Spring Hill, 

Mono Co., CA

W20 DQ251060 

(HVII) 6 0.00% 6.5%with W3

P. kolobensis, 

differ by

3.5-4.7%, 

P. marcida,

differ by 3.9%

Spring in West 

Queen Canyon, 

Mineral Co., NV

W21 DQ251070 

(HXVII) 6 0.00% 6.4%with W3

P. kolobensis, 

differ by

3.3-4.4%, 

P. marcida,

differ by 3.7%

Spring south of 

Warren Lake, Inyo 

Co., CA

W22

DQ251071 

(HXVIII, N=5), 

DQ251072 

(HXIX, N=1)

6 0-0.2% 6.5-6.7%

with W3

P. kolobensis, 

differ by

3.2-4.7%, 

P. marcida, differ 

by 3.6-3.7%

Spring along

Lubken Creek,

Inyo Co., CA

W23 HVI 4 0.00% 6.7%

with W3

P. kolobensis, 

differ by

3.3-4.5%,

P. marcida,

differ by 3.7%

Spring in Marble 

Canyon, Inyo Co.,

CA

Haplotype 

VI is not in 

GenBank

W24 DQ251053 

(HXXI) 5 0.00% 7.1%with W3

P. kolobensis, 

differ by

4.4-5.5%,

P. marcida,

differ by 4.5%

Spring south of 

Summit Creek, Inyo 

Co., CA

HYDROBIIDAE
Technical Note 449 127

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

wongi

(continued)

W25 DQ251064 

(HXI) 5 0.00% 6.4% with W3

P. kolobensis, 

differ by

3.3-4.4%,

P. marcida,

differ by 3.7%

Boron Springs, Inyo 

Co., CA

W26

DQ251064 

(HXI, N=3), 

DQ251073 

(HXX, N=2)

5 0-0.2% 6.2-6.4%

with W3

P. kolobensis, 

differ by

3.2-4.4%,

P. marcida, differ 

by 3.6-3.7%

French Spring, Inyo 

Co., CA

W27 DQ251064 

(HXI) 5 0.00% 6.4% with W3

P. kolobensis, 

differ by

3.3-4.4%,

P. marcida,

differ by 3.7%

Barrel Springs, Inyo 

Co., CA

W28 DQ251074 

(HXXIV) 5 0.00% 7.3% with W3

P. kolobensis, 

differ by

4.6-5.6%,

P. marcida,

differ by 4.7%

Spring north of 

Johnson Canyon, 

Inyo Co., CA

Pyrgulopsis

sp. A BS/D23A

AY197587

(BS/D23A), 

AY426367 

(D23C)

2 1.20% 1.0-1.2% with 

KL_b

P. archimedis, 

differ by

2.6-3.6%,

P. rupinicola,

differ by

2.4-3.4%

Big Springs at 

Bonanza, Klamath 

basin, Klamath 

Co., OR

AY197587/

BS/D23A 

listed as P. 

sp. HPL-2003 

isolate BS 

in GenBank, 

AY426367/

D23C listed 

as P. sp. 

USNM

1016099 

isolate D23C 

in GenBank

KL_b AY197588 1 0.00% 1.0-1.2% with 

BS/D23

P. archimedis, 

differ by

2.7-3.2%, 

P. rupinicola,

differ by

3.1-3.2%

Seventh Link River 

spring, Klamath 

basin, Klamath 

Co., OR

AY197588KLb 

listed as P. 

sp. HPL-2003 

isolate KLb in 

GenBank

SP/D25

AY197589

(SP/D25A), 

AY426368 

(D25B)

2 0.00%

0.5-0.7% with 

D23, 0.5% with 

KL_b

P. archimedis, 

differ by

2.4-3.0%,

P. rupinicola,

differ by

2.6-2.8%

Sprague River north 

of Beatty, Klamath 

basin, Lake Co., OR

AY197589/

SP/D25A 

listed as P. 

sp. HPL-2003 

isolate SP 

in GenBank, 

AY426368/

D25B listed 

as P. sp. 

USNM

1016100 

isolate D25B 

in GenBank

Pyrgulopsis

sp. B P161 AY379446-47 2 0.00%

P. variegata,

differ by 1.1%, 

P. sp. I, differ

by 0.2%

Teton River, Buxton 

Bridge crossing, 

Teton Co., WY

P. sp. 

B-HPL-2003 

isolate 

P161A/

P161B in 

GenBank

HYDROBIIDAE
128 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

sp. C P139

EU700472 

(P139A), 

AY426369 

(P139B),

AY426970 

(P139C)

3 0.00%

P. kolobensis, 

differ by

0.2-3.0%,

P. pilsbryana, 

differ by

0-0.2%,

P. transversa, 

differ by 1.1%,

P. sp. D,

differ by 0.2%, 

P. sp. H,

differ by 0%

Mud Creek, Birch 

Creek Valley, Lemhi 

Co., ID

P. sp. Birch 

Creek isolate 

P139A in 

GenBank, 

P. sp. 

USNM905287 

isolate P139B 

in GenBank, 

P. sp. 

USNM905287 

isolate P139C 

in GenBank, 

P. pilsbryana

clade in 

Hershler et al. 

(2008)

Prygulopsis

sp. D P186 AY426344-45 2 0.00%

P. kolobensis, 

differ by

0.0-2.9%, 

P. nonaria, differ 

by 1.1%, 

P. pilsbryana, 

differ by 0-0.2%,

P. transversa, 

differ by 1.1%,

P. sp. C,

differ by 0.2%,

P. sp. H,

differ by 0.2%

East Fork Rock 

Creek, Rockland 

Valley, Power 

Co., ID

P. sp. USNM

905313

isolate

P186A/P186D 

in GenBank, 

P. pilsbryana

clade in 

Hershler et al. 

(2008)

Pyrgulopsis

sp. E P78B EU700465 1 0.00%

P. kolobensis, 

differ by

7.5-9.6%,

P. pilsbryana, 

differ by

7.6-8.0%,

P. sp. H,

differ by 7.7%

Indian Springs, 

Cold Creek drainage, 

Power Co., ID

P. sp. Indian 

Springs

isolate P78B

in GenBank

Pyrgulopsis

sp. F P158C EU700474 1 0.00% P. variegata,differ by 1.7%

McClenden Spring, 

Raft River drainage, 

Cassia Co., ID

P. sp. 

McClenden 

isolate P158C 

in GenBank

Pyrgulopsis

sp. G P160A EU700476 1 0.00%

P. kolobensis, 

differ by

3.3-4.6%,

P. marcida,

differ by 3.8%,

P. nonaria,

differ by 3.7%,

P. variegata,

differ by 3.8%

Upper Rock Spring, 

Bannock Creek 

drainage, Power 

Co., ID

P. sp. Upper 

Rock Spring 

isolate P160A 

in GenBank

HYDROBIIDAE
Technical Note 449 129

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Pyrgulopsis

sp. H P163A EU700477 1 0.00%

P. kolobensis, 

differ by

0.2-2.7%, 

P. nonaria,

differ by 1.2%,

P. pilsbryana, 

differ by 0-0.2%,

P. transversa, 

differ by 1.1%,

P. sp. C, differ

by 0.0%,

P. sp. D, differ

by 0.2%

Kaufman Cabin 

Springs, Birch Creek 

Valley, Lemhi Co., ID

P. sp.

Kaufman 

Cabin isolate 

P163A in 

Genbank, 

P. pilsbryana

clade in 

Hershler et al. 

(2008)

Pyrgulopsis

sp. I P214A EU700481 1 0.00%

P. variegata,

differ by 1.3%,

P. sp. B, differ

by 0.2%

Spring, at Porcupine 

Ranger Station, 

Henrys Fork 

drainage, Fremont 

Co., ID

P. sp. 

Porcupine 

isolate P214A 

in GenBank

Fluminicola

Fluminicola 

ahjumawi sem93C AY962894 1 0% 0.9% with SP476B

F. lunsfordensis, 

differ by 2.2%

Lost Creek (middle 

site), Shasta Co., CA

F10_116B AY962895 1 0% 1.1% with 

SP476B

F. lunsfordensis, 

differ by 2.0%

Hat Creek at Bridge 

Picnic Area, Shasta 

Co., CA

F10_164A AY962896 1 0% 1.1% with 

SP476B

F. lunsfordensis, 

differ by 2.0%

Three springs on 

point opposite large 

island in Pit River, 

Shasta Co., CA

F10_195C AY962897 1 0% 1.1% with 

SP476B

F. lunsfordensis, 

differ by 1.9%

Spring run near 

Pit River hatchery, 

Shasta Co., CA

sem340A AY962898 1 0% 1.2% with 

SP476B

F. lunsfordensis, 

differ by 1.9%

Honn Creek, Shasta 

Co., CA

F10_344B AY962899 1 0%

0.9% with 

SP476B, 0.9% 

with Fsem535A

F. lunsfordensis, 

differ by 1.9%

Spring run north of 

Sam Wolfin Spring, 

Shasta Co., CA

FSP345E AY962900 1 0% 1.1% with 

SP476B

F. lunsfordensis, 

differ by 1.9%

Upper Sucker 

Springs Creek, 

Shasta Co., CA

sem347B AY962901 1 0% 1.1% with 

SP476B

F. lunsfordensis, 

differ by 2.0%

Spring west of 

Thousand Spring 

run, Shasta Co., CA

sem349A AY962902 1 0% 1.1% with 

SP476B

F. lunsfordensis, 

differ by 2.0%

West spring source 

of Mallard Creek, 

Shasta Co., CA

F10_405A AY962903 1 0%

0.9% with 

SP476B, 0.9% 

with Fsem535A

F. lunsfordensis, 

differ by 1.9%

Beaver Creek, 

Lassen Co., CA

sem408C AY962904 1 0% 0.9% with 

SP476B

F. lunsfordensis, 

differ by 2.5%

Burney Creek, 

upstream of

Burney Falls,

Shasta Co., CA

SP476B AY962905 1 0% 1.2% with 

sem340A

F. lunsfordensis, 

differ by 2.7%

Jimmerson Spring, 

Modoc Co., CA

HYDROBIIDAE LITHOGLYPHIDAE
130 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

ahjumawi

(continued)

Fsem535A AY962906 1 0%

0.9% with 

F10_195C, 

sem340A, 

F10_344B, 

F10_405A, 

SP476B

F. lunsfordensis, 

differ by 2.7%

Lost Creek near 

source spring, 

Shasta Co., CA

Fsem536A AY962907 1 0%

0.8% with 

F10_195C, 

sem340A, 

F10_344B, 

F10_405A, 

SP476B

F. lunsfordensis, 

differ by 2.5%

Lost Creek 

(uppermost site), 

Shasta Co., CA

Fluminicola 

anserinus F6_40B AY962984 1 0%

2.7% with 

F15_270B, 

SP395A

F. multifarius, 

differ by

2.9-3.8%

Spring near Chalk 

Mountain, Shasta 

Co., CA

F15_270B AY962908 1 0% 2.7% with 

F6_40B

F. multifarius, 

differ by

2.7-3.6%

Spring, Goose 

Valley, Shasta Co., 

CA

F6_321A AY962985 1 0%

2.4% with 

F15_270B, 

SP395A

F. multifarius, 

differ by

2.6-3.5%

Blackberry Creek, 

Shasta Co., CA

SP395A AY962909 1 0% 2.7% with 

F6_40B

F. multifarius, 

differ by

2.7-3.6%

Rim of the Lake 

Spring, Shasta 

Co., CA

Fluminicola 

caballensis SP400A AY962910 1 0% 0.8% with FSP401A

F. erosus, differ 

by 1.7-2.0%

Bob Creek, Lassen 

Co., CA

FSP401A AY962911 1 0% 0.8% with 

SP400A

F. erosus, differ 

by 1.5-1.8%

Davis Creek, Lassen 

Co., CA

SP402A AY962912 1 0% 0.6% with 

SP400A

F. erosus, differ 

by 1.4-1.7%

Russell Dairy 

Spring, Lassen 

Co., CA

SP403B AY962913 1 0% 0.6% with 

SP400A

F. erosus, differ 

by 1.4-1.7%

Spring run west of 

Russell Dairy 

Spring, Lassen 

Co., CA

SP404A AY962914 1 0% 0.6% with 

SP400A

F. erosus, differ 

by 1.4-1.7%

Second spring west 

of Russell Dairy 

Spring, Lassen 

Co., CA

Fluminicola 

coloradoense F01 JQ996156 6 0% 0.6-0.8% with F13, F60

F. fuscus, differ 

by 3.9-4.3%

Portneuf River, 

upper access area, 

Bannock Co., ID

F02

JQ996157 

(n=1), 

JQ996158 

(n=1),

JQ996159 

(n=1),

JQ996160 

(n=2)

5 0-0.6% 0.9-1.4% with 

F60

F. fuscus, differ 

by 4.3-4.7%

Snake River, Shelley, 

Bingham Co., ID

F04 JQ996161 3 0% 0.8-0.9% with 

F13, F60

F. fuscus, differ 

by 4.1-4.4%

Blackfoot River, The 

Narrows, ca. 1.5 km 

above weir, Caribou 

Co., ID

F05 JQ996156 5 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Raft River, The 

Narrows bridge, 

Cassia Co., ID

LITHOGLYPHIDAE
Technical Note 449 131

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

coloradoense

(continued)

F07 JQ996156 5 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Summit Creek, BLM 

camp, Custer Co., ID

F08

JQ996156 

(n=5), 

JQ996162 

(n=1)

6 0-0.2% 0.6-0.9% with 

F13, F60

F. fuscus, differ 

by 3.9-4.4%

Henrys Fork, Saint 

Anthony, Fremont 

Co., ID

F09 JQ996163 5 0% 0.8-0.9% with 

F13, F60

F. fuscus, differ 

by 4.1-4.4%

Birch Creek, Mud 

Creek, Lemhi Co., ID

F10

JQ996164 

(n=4), 

JQ996165 

(n=1)

5 0-0.2% 0.8-1.1% with 

F60

F. fuscus, differ 

by 3.8-4.3%

Little Wood River, 

Jim Brown bridge, 

Lincoln Co., ID

F12 JQ996165 5 0% 0.9-1.1% with 

F60

F. fuscus, differ 

by 3.9-4.3%

Snake River, above 

Eagle Rock, Power 

Co., ID

F13

JQ996166 

(n=4), 

JQ996167 

(n=1)

5 0-0.2% 1.3-1.6% with 

F60

F. fuscus, differ 

by 4.4-5.0%

Teton River, Buxton 

bridge, Teton Co., ID

F14

JQ996168 

(n=2), 

JQ996156 

(n=1),

JQ996169 

(n=1),

JQ996170 

(n=1)

5 0-0.5% 0.6-1.1% with 

F60

F. fuscus, differ 

by 3.9-4.6%

Snake River, above 

Murtaugh, Twin 

Falls Co., ID

F15 JQ996160 5 0% 0.9-1.1% with 

F60

F. fuscus, differ 

by 4.3-4.6%

Salt River, Freedom, 

Lincoln Co., WY 

F16

JQ996160 

(n=7), 

JQ996163 

(n=1)

8 0-0.5% 0.8-1.1% with 

F60

F. fuscus, differ 

by 4.1-4.6%

Bear River, Black 

Canyon, above 

Grace Power Plant, 

Caribou Co., ID

F17

JQ996156 

(n=1), 

JQ996163 

(n=2)

3 0-0.2% 0.6-0.9% with 

F13, F60

F. fuscus, differ 

by 3.9-4.4%

Big Malad Spring, 

Oneida Co., ID

F18 JQ996156 3 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Hams Fork, Taylor 

Creek, Lincoln Co., 

WY

F19

JQ996156 

(n=4), 

JQ996171 

(n=1)

5 0-0.2% 0.6-0.9% with 

F13, F60

F. fuscus, differ 

by 3.9-4.4%

New Fork and 

East Fork Rivers, 

southwest of New 

Fork, Sublette Co., 

WY

F20 JQ996156 3 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Beaver Creek, 

below Thorn Creek 

confluence, Summit 

Co, UT

F21

JQ996156 

(n=3), 

JQ996172 

(n=1)

4 0-0.2% 0.6-0.9% with 

F13, F60

F. fuscus, differ 

by 3.9-4.4%

East Canyon Creek, 

above East Canyon 

Reservoir, Morgan 

Co., UT

F22 JQ996156 3 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Strawberry Creek, 

above Weber River 

confluence, Morgan 

Co., UT

F23

JQ996165 

(n=3), 

JQ996173 

(n=1)

4 0-0.9% 0.9-1.4% with 

F60

F. fuscus,

differ by

3.9-4.7%

Murray Spring, 

below source,

Cache Co., UT

LITHOGLYPHIDAE
132 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

coloradoense

(continued)

F24

JQ996165 

(n=3), 

JQ996174 

(n=1)

4 0-0.9% 0.9-1.4% with 

F60

F. fuscus, differ 

by 3.9-4.9%

Blacksmith Fork, 

below Big Hollow, 

Cache Co., UT

F25

JQ996156 

(n=2), 

JQ996175 

(n=2)

4 0-0.2% 0.6-0.9% with 

F13, F60

F. fuscus, differ 

by 3.9-4.4%

South Fork Smiths 

Fork, Cokeville, 

Lincoln Co., WY

F26

JQ996156 

(n=2), 

JQ996176 

(n=2)

4 0-0.3% 0.6-1.1% with 

F13, F60

F. fuscus, differ 

by 3.9-4.6%

Green River, above 

Telephone Island, 

Sweetwater Co., 

WY

F27 JQ996176 3 0% 0.9-1.1% with 

F13, F60

F. fuscus, differ 

by 4.3-4.6%

La Barge Creek, 

west-southwest of 

La Barge, Lincoln 

Co., WY

F28 JQ996156 5 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

New Fork River, 

Hwy 351 bridge, 

Sublette Co., WY 

F29

JQ996156 

(n=3), 

JQ996161 

(n=1),

JQ996177 

(n=1)

4 0-0.3% 0.6-0.9% with 

F13, F60

F. fuscus, differ 

by 3.9-4.4%

Snake River, Grand 

Canyon, Lincoln 

Co., WY

F30

JQ996178 

(n=4), 

JQ996179 

(n=1)

5 0-0.3% 1.1-1.3% with 

F60

F. fuscus, differ 

by 4.4-4.7%

Snake River, 

Byington Park, 

Jefferson Co., ID

F31 JQ996156 3 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Falls River, northnortheast of 

Grainville, Fremont 

Co., ID

F32

JQ996156 

(n=5), 

JQ996180 

(n=1)

6 0-0.2% 0.6-0.8% with 

F13, F42, F60

F. fuscus, differ 

by 3.9-4.4%

Rock Springs,

Arbon Valley,

Power Co., ID

F33 JQ996156 3 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Little Malad Spring, 

Oneida Co., ID

F35 JQ996181 3 0% 1.1-1.3% with 

F60

F. fuscus, differ 

by 4.4-4.7%

Chubb Springs, 

north-northeast of 

Blackfoot Reservoir, 

Caribou Co., ID

F36 JQ996161 3 0% 0.8-0.9% with 

F13, F60

F. fuscus, differ 

by 4.1-4.4%

Blackfoot River, 

mouth of The 

Narrows, Caribou 

Co., ID 

F38 JQ996182 3 0% 1.1-1.3% with 

F60

F. fuscus, differ 

by 4.4-4.7%

North Creek, Liberty, 

Bear Lake Co., ID 

F39 AY962915 1 0% 0.9-1.1% with 

F13, F60

F. fuscus, differ 

by 4.1-4.6%

Green River, Warren 

Bridge, Sublette 

Co., WY

F40 AF520931/

JQ996183 1 0% 1.0-1.4% with F60

F. fuscus, differ 

by 4.0-4.3%

Provo River, 

above Deer Creek 

Reservoir, Wasatch 

Co., UT 

F41

JQ996156 

(n=1), 

JQ996184 

(n=1)

2 0-0.6% 0.6-1.4% with 

F13, F60

F. fuscus, differ 

by 3.9-4.9%

Snake River, ca.

2.4 km below mouth 

of Malad River, 

Gooding Co., ID

LITHOGLYPHIDAE
Technical Note 449 133

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

coloradoense

(continued)

F42

JQ996185 

(n=1), 

JQ996186 

(n=1)

2 1.30% 0.6-1.4% with 

F13

F. fuscus, differ 

by 4.1-4.6%

Snake River, 

Thousand Springs, 

Minnie Miller 

Springs, Gooding 

Co., ID

F43 JQ996156 1 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Snake River, below 

Bliss Dam, Gooding 

Co., ID 

F45

JQ996156 

(n=3), 

JQ996187 

(n=1),

JQ996188 

(n=1)

5 0-0.8% 0.6-1.4% with 

F13, F60

F. fuscus, differ 

by 3.9-4.7%

Deep Creek, above 

Stone Canal, Oneida 

Co., ID

F47 JQ996160 5 0% 0.9-1.1% with 

F60

F. fuscus, differ 

by 4.3-4.6%

Snake River, Black 

Canyon, Bonneville 

Co., ID

F48 JQ996156 5 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Porcupine Station 

Springs, Fremont 

Co., ID

F49 JQ996157 3 0% 0.9-1.1% with 

F60

F. fuscus, differ 

by 4.3-4.6%

Willow Creek,

Kepps Crossing, 

Bonneville Co., ID

F50

JQ996156 

(n=1), 

JQ996160 

(n=2)

3 0-0.3% 0.6-1.1% with 

F60

F. fuscus, differ 

by 3.9-4.6%

Snake River, Clear 

Lake bridge,

Gooding Co., ID

F51

JQ996164 

(n=2), 

JQ996189 

(n=1)

3 0-0.3% 0.8-1.3% with 

F13, F60

F. fuscus, differ 

by 3.5-4.3%

Willow Creek, 

Timmerman rest 

stop, Blaine Co., ID

F52

JQ996164 

(n=1), 

JQ996190 

(n=1),

JQ996191 

(n=1)

3 0.2-0.8% 0.8-1.4% with 

F13

F. fuscus, differ 

by 3.8-4.6%

Big Wood River, 

Malad Gorge, 

Gooding Co., ID

F53 JQ996156 4 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Crooked Creek,

State Wayside, 

Malheur Co., OR

F56

JQ996156 

(n=1), 

JQ996192 

(n=2)

3 0-0.2% 0.6-0.9% with 

F13, F60

F. fuscus, differ 

by 3.9-4.4%

Pahsimeroi River, 

Goldburg Creek, 

Custer Co., ID

F57 JQ996156 3 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Salmon River, 

Kilpatrick, Lemhi 

Co., ID

F58 JQ996156 3 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Salmon River, 

Mackay Bar, Idaho 

Co., ID

F59 JQ996156 3 0% 0.6-0.8% with 

F13, F60

F. fuscus, differ 

by 3.9-4.3%

Salmon River, 

Gasper Creek, Idaho 

Co., ID

F60

JQ996193 

(n=2), 

JQ996194 

(n=1)

3 0-0.8% 1.3-1.6% with 

F13

F. fuscus, differ 

by 4.1-4.7%

Len Lewis spring, 

Hagerman National 

Fish Hatchery, 

Gooding Co., ID

LITHOGLYPHIDAE
134 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

coloradoense

(continued)

F61

JQ996186 

(n=2), 

JQ996195 

(n=1)

3 0-0.5% 1.1-1.4% with 

F60

F. fuscus, differ 

by 4.3-4.7%

Thousand Springs, 

power plant outflow 

just above bridge, 

Gooding Co., ID

F63

JQ996156 

(n=2), 

JQ996196 

(n=1)

3 0-0.2% 0.6-0.9% with 

F13, F60

F. fuscus, differ 

by 3.9-4.4%

Snake River, King 

Hill SA, Elmore 

Co., ID

Fluminicola

dalli D43A AY962916 1 0%

F. anserinus,

differ by

3.8-4.1%,

F. fremonti,

differ by 3.8%,

F. multifarius, 

differ by

3.2-4.6%,

F. umbilicatus, 

differ by

3.8-4.0%

Spring west of 

Thunderbolt Bay, 

Washoe Co., NV

Fluminicola 

erosus

F11_200B, 

F12_200E

AY962918

AY962920 2 0.20% 0.3-0.5% with F11_202A

F. caballensis, 

differ by

1.4-1.8%,

F. favillaceus, 

differ by

0.9-1.7%

SmokeyCharley 

Spring, Modoc Co., 

CA

F10_202E, 

F11_202A,

F12_202A

AY962917

AY962919

AY962921

3 0.3-0.6% 0.2-0.5% with 

F11_200B

F. caballensis, 

differ by

1.4-2.0%,

F. favillaceus, 

differ by

0.9-1.8%

Spring southeast 

of Smokey Charley 

Spring, Modoc Co., 

CA

Fluminicola 

favillaceus SP364B AY962927 1 0% 0.5% with SP510A

F. erosus, differ 

by 1.1-1.4%

Ash Creek at north 

culvert, Lassen 

Co., CA

SP487A AY962928 1 0% 0.6% with 

SP510A

F. erosus, differ 

by 0.9-1.2%

Ash Creek above 

FS39N50 bridge, 

Lassen Co., CA

SP509A AY962929 1 0% 0.5% with 

SP487A

F. erosus, differ 

by 1.4-1.7%

Chisolm Spring, 

Lassen Co., CA

SP510A AY962930 1 0% 0.6% with 

SP487A

F. erosus, differ 

by 1.5-1.8%

Ash Creek, south 

culvert at Ash 

Valley Road 

crossing, Lassen 

Co., CA

Fluminicola 

fremonti

D45aA, 

D45aB

AY962931

AY962932 2 0%

F. anserinus,

differ by

3.5-3.8%,

F. dalli, differ

by 3.8%

Hunters Spring, 

Lake Co., OR

Fluminicola 

fuscus F03 JQ996197 8 0% 0.3-1.6% with F06

F. coloradoense, 

differ by

3.6-4.9%

Boise River, Willow 

Creek, Boise Co., ID

F06

JQ996198 

(n=4), 

JQ996199 

(n=1)

5 0-1.9%

0.5-1.7% with 

F44, F55, 0.2-

1.9% with F46

F. coloradoense, 

differ by

3.6-4.9%

Owyhee River,

below Whistling

Bird Rapids, 

Malheur Co., OR

LITHOGLYPHIDAE
Technical Note 449 135

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

fuscus

(continued)

F11

JQ996197 

(n=4), 

JQ996200 

(n=1)

5 0-0.2% 0.3-1.7% with 

F06

F. coloradoense, 

differ by

3.6-5.0%

Bruneau River, BLM 

low site, Owyhee 

Co., ID

F44 DQ372901/

JQ996201 1 0% 0.5-1.7% with F06

F. coloradoense, 

differ by

3.8-4.7%

Grande Ronde

River, 0.2 km above 

mouth, Asotin Co., 

WA

F46

JQ996202 

(n=4), 

JQ996203 

(n=1)

5 0-0.2% 0.2-1.9% with 

F06

F. coloradoense, 

differ by

3.5-4.9%

Methow River north 

of Squaw Creek 

mouth, Okanogon 

Co., WA

F54 JQ996197 4 0% 0.3-1.6% with 

F06

F. coloradoense, 

differ by

3.6-4.9%

Payette River, 

Banks Camp 

[Campground],

Boise Co., ID

F55 JQ996204 3 0% 0.5-1.7% with 

F06

F. coloradoense, 

differ by

3.8-5.0%

Imnaha River, Bare 

Creek, Wallowa 

Co., OR

F62 JQ996197 3 0% 0.3-1.6% with 

F06

F. coloradoense, 

differ by

3.6-4.9%

Snake River, Grand 

View SA, Owyhee 

Co., ID

F64 JQ996198 3 0% 0-1.9% with

F06

F. coloradoense, 

differ by

3.6-4.9%

Owyhee River,

above Beaver 

Charlie

Cabin, Malheur 

Co., OR

F65 JQ996197 4 0% 0.3-1.6% with 

F06

F. coloradoense, 

differ by

3.6-4.9%

Boise River, Willow 

Creek, Boise Co., ID

F66 JQ996197 4 0% 0.3-1.6% with 

F06

F. coloradoense, 

differ by

3.6-4.9%

Boise River, 

Badger Camp 

[Campground],

Boise Co., ID

Fluminicola 

gustafsoni F67

JQ731609 

(n=1), 

JQ731610 

(n=2)

3 0-0.2% 0.5-0.8% with 

F73

F. virens,

differ by

6.5-6.6%

Clearwater River, 

Jim Ford Creek, 

Clearwater Co., ID

F68 JQ731611 4 0% 0.5-0.6% with 

F73

F. virens,

differ by 6.6%

Clearwater River, 

Orofino, Clearwater 

Co., ID

F69 JQ731612 3 0% 0.3-0.5% with 

F67

F. virens,

differ by 6.3%

Salmon River, Pine 

Bar Rapids, Idaho 

Co., ID

F70 JQ731613 2 0% 0.5-0.6% with 

F73

F. virens,

differ by 6.6%

South Fork 

Clearwater River, 

Battlefield, Idaho 

Co., ID

F73

JQ731614 

(n=1), 

JQ731615 

(n=2)

3 0-0.6% 0.5-0.8% with 

F67

F. virens,

differ by

6.2-6.3%

Snake River, below 

mouth of Couse 

Creek, Asotin Co., 

WA

LITHOGLYPHIDAE
136 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

insolitus D35A AY962934 1 0%

F. coloradoense, 

differ by

8.7-9.8%,

F. fuscus, differ 

by 8.8-9.1%

Page Springs, 

Harney Co., OR

Fluminicola 

lunsfordensis

F3_59A, 

F3_59B

AY962935,

AY962936 2 0%

F. ahjumawi,

differ by

1.9-2.7%, 

Fluminicola sp.

B, differ by

2.4-2.5%

Lunsford Spring, 

Modoc Co., CA

Fluminicola 

modoci D42B AY962938 1 0% 0.9% with D40A

F. seminalis,

differ by

5.0-5.6%

Spring at Three 

Springs Ranch, 

Modoc Co., CA

D40A AY962937 1 0% 0.9% with D42B

F. seminalis,

differ by

5.6-6.1%

Link River at 

Klamath Falls 

bridge, Klamath 

Co., OR

Fluminicola 

multifarius SP8B AY962939 1 0% 2.9% with F10_99B

F. anserinus,

differ by

3.3-3.6%

Spring near Conant, 

Siskiyou Co., CA

F2_10B AY962975 1 0% 2.6% with 

F3_69A

F. anserinus,

differ by

3.0-3.5%

Ney Springs, 

Siskiyou Co., CA

F1_12B AY962976 1 0% 2.6% with 

F3_69A

F. anserinus,

differ by

3.0-3.5%

Sacramento River 

near Stink Creek, 

Siskiyou Co., CA

F6_27B AY962940 1 0% 2.9% with 

F10_99B

F. anserinus,

differ by

3.3-3.6%

Crystal Spring, 

Siskiyou Co., CA

F4_30A, 

F5_30A

AY962941,

AY962942 2 0% 2.9% with F10_99B

F. anserinus,

differ by

3.3-3.6%

Rock Spring, 

Siskiyou Co., CA

F3_65A, 

F4_65A,

F5_65A,

FSP65F

AY962943,

AY962944,

AY962945,

AY962946

4 0-0.2% 2.9% with 

F10_99B

F. anserinus,

differ by

3.3-3.6%

Southernmost of 

Shasta Springs, 

Shasta Co., CA

F4_68A AY962947 1 0% 2.9% with 

F10_99B

F. anserinus,

differ by

3.3-3.6%

Spring north of 

Mossbrae Falls, 

Shasta Co., CA

F3_69A AY962948 1 0% 3.0% with 

F10_99B

F. anserinus,

differ by

3.5-3.8%

Spring runs north 

of Mossbrae Falls, 

Shasta Co., CA

F10_99B AY962977 1 0% 3.0% with 

F3_69A

F. anserinus, 

differ by

2.6-2.9%

Big Springs 

(source), northwest 

of city of Mount 

Shasta, Shasta 

Co., CA

F1_140A AY962949 1 0% 2.9% with 

F10_99B

F. anserinus,

differ by

3.3-3.6%

Sacramento River 

at Cave Springs, 

Siskiyou Co., CA

F2_143A AY962978 1 0% 2.7% with 

F3_69A

F. anserinus,

differ by

2.6-3.0%

Spring along 

Sacramento River 

(third to the) east 

of Cantara Bend, 

Siskiyou Co., CA

LITHOGLYPHIDAE
Technical Note 449 137

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

multifarius

(continued)

F2_144A AY962979 1 0% 2.6% with 

F3_69A

F. anserinus,

differ by

3.0-3.5%

Spring along 

Sacramento River 

(first to the) east 

of Cantara Bend, 

Siskiyou Co., CA

SP237A AY962980 1 0%

2.4% with 

F3_69A, 

F10_322B

F. anserinus,

differ by

3.2-3.6%

Big Springs, three 

middle runs, 

Siskiyou Co., CA

SP238B AY962981 1 0%

2.4% with 

F3_69A, 

F10_322B

F. anserinus,

differ by

3.2-3.6%

Big Springs, 

westernmost run, 

Siskiyou Co., CA

F10_239B AY962982 1 0% 2.7% with 

F3_69A

F. anserinus,

differ by

2.9-3.3%

Big Springs, 

easternmost run, 

Siskiyou Co., CA

SP241A AY962983 1 0% 2.6% with 

F3_69A

F. anserinus,

differ by

3.0-3.5%

Big Springs at 

west side of park, 

Siskiyou Co., CA

F10_322B AY962950 1 0% 2.7% with 

F10_99B

F. anserinus,

differ by

3.2-3.5%

Bundoora Spring, 

Siskiyou Co., CA

F3_330A AY962951 1 0% 2.9% with 

F10_99B

F. anserinus,

differ by

3.3-3.6%

Spring north of 

Crystal Spring, 

Siskiyou Co., CA

ELKSPB AY962952 1 0% 2.6% with 

F10_99B

F. anserinus,

differ by

3.0-3.3%

Elk Spring 

(lowermost), 

Siskiyou Co., CA

Fluminicola 

neritoides F9_369B AY962953 1 0% no variation F. lunsfordensis, differ by 4.9%

Willow Creek at 

mouth of Hayden 

Canyon, Lassen 

Co., CA

F9_371A AY962954 1 0% no variation F. lunsfordensis, 

differ by 4.9%

Willow Creek at 

lower end of Lower 

McBride Springs, 

Lassen Co., CA

F9_372A AY962955 1 0% no variation F. lunsfordensis, 

differ by 4.9%

Willow Creek west

of Hayden Hill, 

Lassen Co., CA

Fluminicola 

potemicus

F2_36A, 

F2_36B

AY962956,

AY962957 2 0%

F. anserinus,

differ by

3.5-4.1%,

F. multifarius, 

differ by

3.2-3.8%,

F. umbilicatus, 

differ by

3.6-3.8%

Spring near Potem 

Creek, Shasta Co., 

CA

Fluminicola 

scopulinus F14_251B AY962958 1 0% 0.5% with F14_303B

F. anserinus,

differ by

3.8-4.0%, 

F. multifarius, 

differ by

3.5-4.3%,

Northernmost 

spring southwest 

of Popcorn Spring, 

Shasta Co., CA

LITHOGLYPHIDAE
138 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

scopulinus

(continued)

F14_303B AY962959 1 0% 0.5% with 

F14_251B

F. anserinus,

differ by

4.3-4.4%,

F. multifarius, 

differ by

3.6-4.4%

Northernmost 

spring west of 

Popcorn Spring, 

Shasta Co., CA

Fluminicola 

seminalis sem98B AY962960 1 0% 1.2% with sem329A

F. modoci,

differ by

5.5-5.9%

Big Lake outlet near 

Rat Farm, Shasta 

Co., CA

sem104B AY962961 1 0%

1.2% with 

D38, SP366B, 

sem426A, 

Fsem532A, 

Fsem552A

F. modoci,

differ by

5.2-5.8%

Baum Lake (deepwater site), Shasta 

Co., CA

sem105A AY962962 1 0% 1.2% with 

sem329A

F. modoci,

differ by

5.5-5.9%

Fall River at 

CalTrout Public 

Fishing Access 

Area, Shasta

Co., CA

sem329A AY962963 1 0%

1.4% with 

D38, SP366B, 

sem426A, 

Fsem532A, 

Fsem552A

F. modoci,

differ by

5.2-5.8%

Baum Lake

(shallow-water site), 

Shasta Co., CA

D38A AY962964 1 0% 1.4% with 

sem329A

F. modoci,

differ by

5.3-5.8%

Battle Creek east 

of Coleman Fish 

Hatchery, Shasta 

Co., CA

sem364A AY962965 1 0%

1.4% with D38, 

sem426A, 

Fsem532A, 

Fsem552A

F. modoci,

differ by

5.0-5.6%

Ash Creek at north 

culvert, Lassen 

Co., CA

SP366B AY962966 1 0% 1.4% with 

sem329A

F. modoci,

differ by

5.3-5.8%

Ash Creek 

northwest of Ash 

Creek Campground, 

Lassen Co., CA

sem411A AY962967 1 0% 1.2% with 

sem329A

F. modoci,

differ by

5.5-5.9%

Crystal Springs, 

Shasta Co., CA

sem416A AY962968 1 0% 1.2% with 

sem329A

F. modoci,

differ by

5.5-5.9%

Big Lake Springs, 

Shasta Co., CA

sem425A AY962969 1 0% 1.2% with 

sem329A

F. modoci,

differ by

5.5-5.9%

Pit River near 

confluence of Hat 

Creek, Shasta Co., 

CA

sem426A AY962970 1 0% 1.4% with 

sem329A

F. modoci,

differ by

5.6-6.1%

Spring Creek on 

south side of Spring 

Creek Road, Shasta 

Co., CA

sem487A AY962971 1 0% 1.2% with 

sem329A

F. modoci,

differ by

5.2-5.6%

Ash Creek above 

FS39N50 bridge, 

Lassen Co., CA

Fsem532A AY962972 1 0% 1.4% with 

sem329A

F. modoci,

differ by

5.6-6.1%

Lava Creek at 

boathouse on 

Hanna Estate, 

Shasta Co., CA

Fsem546A AY962973 1 0% 1.2% with 

sem329A

F. modoci,

differ by

5.5-5.9%

Spring on west side 

of Spring Creek, 

Shasta Co., CA

LITHOGLYPHIDAE
Technical Note 449 139

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

seminalis

(continued)

Fsem552A AY962974 1 0% 1.4% with 

sem329A

F. modoci,

differ by

5.6-6.1%

Lava Creek source 

spring pool, Shasta 

Co., CA

Fluminicola 

turbiniformis D34B AY962986 1 0%

F. warnerensis, 

differ by

2.3-2.6%

Roaring Springs, 

Harney Co., OR

Fluminicola 

umbilicatus F8_92A AY962987 1 0% 0.3% with F7_338A

F. anserinus,

differ by

3.5-3.8%,

F. multifarius, 

differ by

3.5-4.3%,

F. potemicus, 

differ by 3.8%

Lost Creek 

(lowermost site), 

Shasta Co., CA

F8_93B AY962988 1 0%

0.2% with 

F7_338A, 

F8_536A,

F8_92A

F. anserinus,

differ by

3.3-3.6%,

F. multifarius, 

differ by

3.3-4.1%,

F. potemicus, 

differ by 3.6%

Lost Creek

(middle site),

Shasta Co., CA

F7_99A AY962989 1 0%

0.2% with 

F7_338A, 

F8_536A,

F8_92A

F. anserinus,

differ by

3.3-3.6%,

F. multifarius, 

differ by

3.3-4.1%,

F. potemicus, 

differ by 3.6%

Big Spring,

tributary of Hat 

Creek, Shasta Co., 

CA

F7_338A AY962990 1 0%

0.3% with 

F8_536A,

F8_92A

F. anserinus,

differ by

3.5-3.8%,

F. multifarius, 

differ by

3.5-4.3%,

F. potemicus, 

differ by 3.8%

Hat Creek at Hat 

Creek Resort, 

Shasta Co., CA

F8_536A AY962991 1 0% 0.3% with 

F7_338A

F. anserinus,

differ by

3.5-3.8%,

F. multifarius, 

differ by

3.5-4.3%,

F. potemicus, 

differ by 3.8%

Lost Creek 

(uppermost site), 

Shasta Co., CA

Fluminicola

virens D41A AY962992 1 0%

F. gustafsoni, 

differ by

6.2-6.6%

Willamette River 

at Canby Ferry, 

Clackamas Co., OR

Fluminicola 

virginius D36B AY962993 1 0%

F. dalli, differ

by 5.8%, 

F. potemicus, 

differ by 5.5%,

F. scopulinus, 

differ by

5.5-5.9%

Hardscrabble Creek, 

Washoe Co., NV

LITHOGLYPHIDAE
140 Technical Note 449

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Fluminicola 

warnerensis F13_210B AY962995 1 0% 0.6% with F13_351B

F. turbiniformis, 

differ by 2.3%

Spring creek east of 

Blue Lake, Modoc 

Co., CA

F13_217A AY962996 1 0%

0.5% with 

SP360A, 

FSP387F, 

SP388A, 

F10_482A, 

F13_210B

F. turbiniformis, 

differ by 2.4%

Parsnip Spring, 

Lassen Co., CA

F13_351B AY962997 1 0%

0.6% with 

SP360A, 

FSP387F, 

SP388A, 

F10_482A, 

F13_210B

F. turbiniformis, 

differ by 2.6%

Soup Spring, 

Modoc Co., CA

SP360A AY962999 1 0% 0.6% with 

F13_351B

F. turbiniformis, 

differ by 2.6%

Springs southeast 

of Hilton, Modoc 

Co., CA

FSP387F AY962998 1 0% 0.6% with 

F13_351B

F. turbiniformis, 

differ by 2.6%

Springs east of 

Miller Gulch, Modoc 

Co., CA

SP388A AY963000 1 0% 0.6% with 

F13_351B

F. turbiniformis, 

differ by 2.6%

Miller Spring run, 

Modoc Co., CA

F10_482A AY962994 1 0% 0.6% with 

F13_351B

F. turbiniformis, 

differ by 2.6%

Rush Creek 

(source), Modoc 

Co., CA

Fluminicola

sp. A SP113A AY962922 1 0% 0.2% with SP373B

F. lunsfordensis, 

differ by 4.6%

Springs west of 

Canby, Modoc Co., 

CA

SP373B, 

F2SP373A

AY962924,

AY962925 2 0.2% 0.2% with SP113A

F. lunsfordensis, 

differ by

4.4-4.6%

Spring at west 

end of Upper Rush 

Creek Campground, 

Modoc Co., CA

Fluminicola

sp. B

SP373A, 

F2SP373E

AY962923,

AY962926 2 0.2%

F. lunsfordensis, 

differ by

2.4-2.5%

Spring at west end 

of Upper Rush

Creek Campground, 

Modoc Co., CA

Pristinicola

Pristinicola 

hemphilli AF520940 1 0.00%

Springs, 1.8 km

east of Lower 

Kalama Hatchery, 

Cowlitz Co., WA

Taylorconcha

Taylorconcha 

insperata OWS DQ75986-DQ75990 5 0-0.46% 

0-0.76%

with OWD,

0.15-0.76%

with HCH

T. serpenticola, 

differ by

0.91-1.52%

Owyhee River, 

upstream from 

South Cross 

Canyon, Malheur 

Co., OR

OWD DQ76011-

DQ76014 4 0-0.30% 0-0.76%with OWS

T. serpenticola, 

differ by

1.06-1.82%

Owyhee River, 

at Lower Deary 

Pasture, Malheur 

Co., OR 

HCH DQ75991-

DQ75995 5 0-0.15% 0.15-0.76%with OWS

T. serpenticola, 

differ by

1.06-1.52%

Snake River, above 

High Bar Rapids, 

Wallowa Co., OR 

HCD DQ76023-

DQ76027 5 0% 0-0.61%with OWS

T. serpenticola, 

differ by

1.22-1.52%

Snake River, just 

below Davis Creek 

Rapids, Wallowa 

Co., OR

LITHOGLYPHIDAE UNCERTAIN
Technical Note 449 141

Species

Published 

Sample 

Codes

Accession 

Numbers

No. of 

Specimens (N)

Sequence 

Variation 

(% pairwise 

distance)

Most Distant 

Conspecific 

Sequences (%)

Closest 

Interspecific 

Sequences (%)

Collection Locality Comments

Taylorconcha 

serpenticola NI DQ75952-DQ75956 5 0-0.30% 0.15-0.76%with TSM

T. insperata,

differ by

1.06-1.82%

Niagara Springs, 

outflow at base 

of falls, Gooding 

Co., ID 

BA DQ76019-

DQ76022 4 0% 0.15-0.46%with TSN, NI

T. insperata,

differ by

1.22-1.67%

Banbury Springs 

outlets, Gooding 

Co., ID

SA DQ75961-

DQ75965 5 0% 0.15-0.46%with TSM

T. insperata,

differ by

1.06-1.52%

Sand Spring, ca. 

30 m below source, 

Gooding Co., ID

TSN DQ75996-

DQ776000 5 0-0.61% 0.15-0.76%with TSM

T. insperata,

differ by

0.91-1.52%

Spring pool of 

Thousand Springs 

north outlet,

Gooding Co., ID 

TSM DQ76015-

DQ76018 4 0-0.61% 0.15-0.76%with NI, TSN

T. insperata,

differ by

1.06-1.67%

Thousand Springs, 

springs just south 

of The Nature 

Conservancy's 

water pipeline, 

south of north inlet, 

Gooding Co., ID

TSS DQ76001-

DQ76005 5 0% 0.15-0.46%with TSM

T. insperata,

differ by

1.06-1.52%

Thousand Springs, 

southernmost and 

largest outflow of 

the Minnie Miller 

Springs complex, 

Gooding Co., ID

BI DQ75957-

DQ75960 4 0% 0.15-0.46%with TSM

T. insperata,

differ by

1.06-1.52%

Billingsley Creek, 

at spring source, 

Gooding Co., ID 

CO DQ75966-

DQ75970 5 0-0.30% 0.15-0.61%with TSM

T. insperata,

differ by

0.91-1.67%

Cove Creek, just 

above diversion 

to Malad River, 

Gooding Co., ID

MA DQ76006-

DQ76010 5 0% 0.15-0.46%with TSM

T. insperata,

differ by

1.06-1.52%

Snake River, just 

below Malad 

Power Plant outfall, 

Gooding Co., ID

ZI DQ75971-

DQ75975 5 0-0.15% 0.15-0.61%with TSM

T. insperata,

differ by

0.91-1.52%

Snake River, above 

Bliss Reservoir, 

Gooding Co., ID 

BC DQ75981-

DQ75985 5 0-0.15% 0.15-0.61%with TSM

T. insperata,

differ by

0.91-1.67%

Snake River, just 

below Bancroft 

Springs, Elmore 

Co., ID 

CL DQ75976-

DQ75980 5 0-0.15% 0.15-0.61%with TSM

T. insperata,

differ by

0.91-1.52%

Snake River, just 

above Clover Creek 

confluence, Elmore 

Co., ID 

UNCERTAIN
Photos by Karolyn Darrow.