A Review of the North American
Freshwater Snail Genus
Fluminicola (Hydrobiidae)
*
ROBERT HERSHLER
and
TERRENCE J. FREST
i
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ? NUMBER 583
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S M I T H S O N I A N C O N T R I B U T I O N S T O Z O O L O G Y ? N U M B E R 5 8 3
A Review of the North American
Freshwater Snail Genus
Fluminicola (Hydrobiidae)
Robert Hershler
and Terrence J. Frest
SMITHSONIAN INSTITUTION PRESS
Washington, D.C.
1996
A B S T R A C T
Hershler, Robert, and Terrence J. Frest. A Review of the North American Freshwater Snail
Genus Fluminicola (Hydrobiidae). Smithsonian Contributions to Zoology, number 583, 41
pages, 18 figures, 1996.?Species currently allocated to the northwestern North American genus
Fluminicola Carpenter, 1864, comprise a morphologically diverse group that is shown to be
paraphyletic based on a cladistic analysis of nominal species utilizing European Lithoglyphus
Hartmann, 1821, and two eastern North American genera as outgroups. Fluminicola consists of
a single, highly divergent species (F. virens) and a clade comprising all other congeners, which
is defined by synapomorphic conditions of pedal ganglia and penis. Groups of species within the
clade are united by derived modifications of shell protoconch microsculpture, operculum,
central and lateral radular teeth, ctenidium, hypobranchial gland, penis, ovary, and female
capsule gland.
A redefinition of Fluminicola in the wake of the above analysis cannot be provided until
anatomical details of the type species (which may be extinct) are obtained. Nine Recent species
currently are assigned to this group. These are Fluminicola coloradensis, distributed in the
Green River basin; F. dalli, Pyramid Lake basin; F. fuscus, Snake-Columbia River basin; F.
minutissimus, Snake River basin; F. modoci, Goose Lake basin; F. nuttallianus, Willamette
River basin; F. seminalis, Sacramento and Pit river basins; F. turbiniformis, northwestern Great
Basin; and F. virens, Willamette and lower Columbia river basins.
Amnicola hindsi Baird, 1863, Fluminicola nuttalliana columbiana Keep, 1887, and
Fluminicola columbiana Pilsbry, 1899a, are junior synonyms of F. fuscus (Haldeman, 1841);
Lithoglyphus cumingii Frauenfeld, 1863a, is a junior synonym of F. seminalis (Hinds, 1842);
and Paludina nuclea Lea, 1838, is a junior synonym of F. virens (Lea, 1838). The genus-group
taxon Heathilla Hannibal, 1912, is a synonym of Fluminicola.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is
recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral
Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging-in-Publication Data
Hershler, Robert.
A review of the North American freshwater snail genus Fluminicola (Hydrobiidae) / Robert Hershler and Terrence J.
Frest.
p. cm. ? (Smithsonian contributions to zoology ; no. 583)
Includes bibliographical references (p.37).
1. Fluminicola. I. Frest, Terrence J. II. Title. III. Series.
QL1.S54 no. 583 [QL430.5.H9] 591 s?dc20 [594'.32] 96-15188
? The paper used in this publication meets the minimum requirements of the American
National Standard for Permanence of Paper for Printed Library Materials Z39.48?1984.
Contents
Page
Introduction 1
Materials and Methods 2
Acknowledgments 2
Character Descriptions 2
Phylogenetic Reconstruction 5
Superfamily RISSOOIDEA Gray, 1847 6
Family HYDROBIIDAE Troschel, 1857 6
Subfamily LITHOGLYPHINAE Troschel, 1857 6
Genus Fluminicola Carpenter, 1864 6
Fluminicola coloradensis Morrison, 1940 8
Fluminicola dalli (Call, 1884) 9
Fluminicola fuscus (Haldeman, 1841) 10
Fluminicola minutissimus Pilsbry, 1907 13
Fluminicola modoci Hannibal, 1912 13
Fluminicola nuttallianus (Lea, 1838) 14
Fluminicola seminalis (Hinds, 1842) 15
Fluminicola turbiniformis (Tryon, 1865) 16
Fluminicola virens (Lea, 1838) 17
Appendix: Western American Fossil Species Described as or Allocated to
Fluminicola or Lithoglyphus 20
Literature Cited 37
111
A B S T R A C T
Hershler, Robert, and Terrence J. Frest. A Review of the North American Freshwater Snail
Genus Fluminicola (Hydrobiidae). Smithsonian Contributions to Zoology, number 583, 41
pages, 18 figures, 1996.?Species currently allocated to the northwestern North American genus
Fluminicola Carpenter, 1864, comprise a morphologically diverse group that is shown to be
paraphyletic based on a cladistic analysis of nominal species utilizing European Lithoglyphus
Hartmann, 1821, and two eastern North American genera as outgroups. Fluminicola consists of
a single, highly divergent species (F. virens) and a clade comprising all other congeners, which
is defined by synapomorphic conditions of pedal ganglia and penis. Groups of species within the
clade are united by derived modifications of shell protoconch microsculpture, operculum,
centra] and lateral radular teeth, ctenidium, hypobranchial gland, penis, ovary, and female
capsule gland.
A redefinition of Fluminicola in the wake of the above analysis cannot be provided until
anatomical details of the type species (which may be extinct) are obtained. Nine Recent species
currently are assigned to this group. These are Fluminicola coloradensis, distributed in the
Green River basin; F. dalli. Pyramid Lake basin; F. fuscus, Snake-Columbia River basin; F.
minutissimus, Snake River basin; F. modoci, Goose Lake basin; F. nuttallianus, Willamette
River basin; F. seminalis, Sacramento and Pit river basins; F. turbiniformis, northwestern Great
Basin; and F. virens, Willamette and lower Columbia river basins.
Amnicola hindsi Baird, 1863, Fluminicola nuttalliana columbiana Keep, 1887, and
Fluminicola columbiana Pilsbry, 1899a, are junior synonyms of F. fuscus (Haldeman, 1841);
Lithoglyphus cumingii Frauenfeld, 1863a, is a junior synonym of F. seminalis (Hinds, 1842);
and Paludina nuclea Lea, 1838, is a junior synonym of F. virens (Lea, 1838). The genus-group
taxon Heathilla Hannibal, 1912, is a synonym of Fluminicola.
OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is
recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral
Montastrea cavernosa (Linnaeus).
Library of Congress Cataloging-in-Publication Data
Hershler, Robert.
A review of the North American freshwater snail genus Fluminicola (Hydrobiidae) / Robert Hershler and Terrence J.
Frest
p. cm. ? (Smithsonian contributions to zoology ; no. 583)
Includes bibliographical references (p.37).
1. Fluminicola. I. Frest, Terrence J. II. Title. III. Series.
QL1.S54 no. 583 [QL430.5.H9] 591 s?dc20 [594'32] 96-15188
? The paper used in this publication meets the minimum requirements of the American
National Standard for Permanence of Paper for Printed Library Materials Z39.48 1984.
Contents
Page
Introduction 1
Materials and Methods 2
Acknowledgments 2
Character Descriptions 2
Phylogenetic Reconstruction 5
Superfamily RISSOOIDEA Gray, 1847 6
Family HYDROBIIDAE Troschel, 1857 6
Subfamily LrTHOGLYPHlNAE Troschel, 1857 6
Genus Fluminicola Carpenter, 1864 6
Fluminicola coloradensis Morrison, 1940 8
Fluminicola dalli (Call, 1884) 9
Fluminicola fuscus (Haldeman, 1841) 10
Fluminicola minutissimus Pilsbry, 1907 13
Fluminicola modoci Hannibal, 1912 13
Fluminicola nuttallianus (Lea, 1838) 14
Fluminicola seminalis (Hinds, 1842) 15
Fluminicola turbiniformis (Tryon, 1865) 16
Fluminicola virens (Lea, 1838) 17
Appendix: Western American Fossil Species Described as or Allocated to
Fluminicola or Lithoglyphus 20
Literature Cited 37
in

A Review of the North American
Freshwater Snail Genus
Fluminicola (Hydrobiidae)
Robert Hershler
and Terrence J. Frest
Introduction
Hydrobiid snails of the genus Fluminicola Carpenter, 1864
(subfamily Lithoglyphinae), are among the more ubiquitous
macroinvertebrates in large, lotic water bodies of northwestern
North America. Fluminicola is a relatively small genus as
measured by the number of nominal taxa now associated with
the group, but malacologists have paid scant attention to these
snails and their systematics is confused. In addition, virtually
nothing is known about the morphology and biology of these
animals.
Several species now allocated to Fluminicola were placed in
Lithoglyphus Hartmann, 1821 (an otherwise European group),
subsequent to their original description, and there is no modern
consensus as to whether the North American genus merits
distinction from the latter. Pilsbry (1935) stated that these
genera differ in their penes and therefore did not advocate
placement of North American species in Lithoglyphus. This
opinion was shared by numerous subsequent workers (includ-
ing Starobogatov, 1970; Burch, 1979, 1982; Thompson, 1984;
Vaught, 1989; Banarescu, 1990, 1992). Taylor (1966a:131),
however, found no "anatomical warrant" for the separation of
the two genera and hence treated Fluminicola as a junior
synonym of Lithoglyphus. He later reiterated this opinion
(Taylor 1966b, 1981, 1985; Taylor and Smith, 1971), which
has been accepted by at least two other workers (Clarke 1976,
1981; Beetle, 1989).
Additionally, the composition of Fluminicola has not been
well clarified in the literature, and there are lingering problems
Robert Hershler, Department of Invertebrate Zoology, NHB MRC
118, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560. Terrence J. Frest, Deixis Consultants, 2517
NE 65th Street, Seattle, WA 98115.
concerning the identity, taxonomic status, and limits of
nominal forms. For instance, modern treatments of the small
members of the genus have varied substantially: Taylor (1966a)
placed F. dalli (Call, 1884) and F. modoci Hannibal, 1912, in
synonymy with F. turbiniformis (Tryon, 1865), whereas Burch
and Tottenham (1980) accepted F. modoci and F. turbiniformis
as distinct (F. dalli is not treated in this work). In the only two
papers (Stimpson, 1865b; Thompson, 1984) purporting to treat
the morphology of the type species, F. nuttallianus (Lea,
1838), the exemplar material was not F. nuttallianus but was
misidentified as this snail. Stimpson's early (1865b) study was
based on specimens provided by Binney from an unspecified
locality. The shell illustrated (Stimpson, 1865b, fig. 15) clearly
is not F. nuttallianus, and, although we have not examined
anatomical material for this (probably extinct) species, the
figured penis (Stimpson, 1865b, fig. 16) is not similar to that of
any snail we have studied from the historic range of this species
in the Willamette River. Instead, it resembles penes of
undescribed snails from the Klamath basin. In Thompson's
(1984) recent review of the North American lithoglyphine
genera, the three lots used to characterize F. nuttallianus are of
undescribed species and F. virens (Lea, 1838).
Stabilization of the taxonomy of Fluminicola is especially
desirable at this time because of the interest now being shown
by land management agencies in the conservation of Pacific
Northwest forest ecosystems within the range of the northern
spotted owl, Strix occidentalis caurina (Merriam) (where many
of these snails occur). One species, F. fuscus (Haldeman,
1841), has been a Federal Listing candidate since 1984 (USDI,
1984; as Lithoglyphus columbianus (Pilsbry)).
The purposes of this paper are to determine which nominal
species of Fluminicola are recognizable and to assess the
distinguishing features of these taxa. Based on a preliminary
hypothesis that we propose for phylogenetic relationships
within the genus, we conclude that although western American
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
lithoglyphines (traditionally allocated to Fluminicola) are
distinct from European Lithoglyphus, they do not comprise a
monophyletic group when eastern North American taxa are
considered; instead, they represent two phylogenetically
distinct clades. We cannot determine at this time which (if any)
of these clades corresponds to Fluminicola as the anatomy of
its type species is unknown (we have been unable to collect
material for this presumably extinct animal). For the time
being, we accept this group as a composite assemblage of
western American lithoglyphines.
We hope that this paper will serve as a framework for future
descriptive studies as many novelties in this group await the
attention of systematists, and large geographic regions in the
Northwest still have not been thoroughly surveyed for these
animals. We acknowledge that our concept of nominal species
of Fluminicola will likely change somewhat as additional
anatomical material (for records based on dry shells) becomes
available for study. The three groups of taxa most in need of
scrutiny in this regard are the large snails from the coastal
drainages of Oregon and Washington (often assigned to F.
nuttallianus or F. virens in the literature or in museum
collections); the small snails from the northwest Great Basin
and adjacent regions in California and Oregon (assigned to F.
turbiniformis by Taylor, 1966a, fig. 9); and the large snails
from the northeastern Great Basin and portions of southern
Idaho and southwestern Wyoming (assigned to F. hindsii [sic]
(Baird) by Taylor, 1966a, fig. 14).
Recent nominal species of Fluminicola now assigned to
Pyrgulopsis are not treated herein (for summary of these see
Hershler, 1994). We also do not address fossil taxa associated
with this genus (beyond their listing in the Appendix), given
the difficulty in clarifying systematic relationships of hydrobiid
snails solely on the basis of empty shells.
MATERIALS AND METHODS.?Fluminicola species were
characterized based on examination of type and other dry
museum material and on anatomical study of topotypes or other
available specimens. Anatomical material used as exemplars to
characterize species is listed at the end of the "Remarks"
sections (other alcohol lots were examined more cursorily).
Acronyms for institutional repositories of examined specimens
are those of Leviton et al. (1985). Lots that include alcohol
material are indicated in the "Material Examined" sections by
asterisks. Anatomical study was of alcohol-preserved snails
that had been relaxed with menthol crystals and fixed in dilute
(4%) formalin. Methods of morphological study are those of
Hershler (1994). The phylogenetic analysis was performed
using HENNIG86 (Farris, 1988). Character-state evolution on
the tree generated was studied using CLADOS (Nixon, 1992).
Because anatomical material was not available for the type
species F. coloradensis Morrison, 1940, was used instead to
illustrate details pertaining to the genus generally (the shells of
these two species are closely similar). Type and nontype shells,
shell protoconch, operculum, radula, penis, distal female
genitalia, and geographic distribution are illustrated for each
species.
ACKNOWLEDGMENTS.?We thank the following individuals
(and their respective institutions) for lending material pertinent
to this study: G. Rosenberg and D. Bardes (ANSP), F. Naggs
and J. Freeman (BMNH), T. Gosliner and E. Kools (CAS), F.G.
Thompson (FSM), K. Boss and T. Kausch (MCZ), and K.
Edlinger and the late E. Wawra (NMW). We also thank the
following individuals for the donation of specimens pertinent
to this study and/or for assistance in the field: D. Clark
(Portland General Electric, Oregon City), M. Haase (NMW), P.
Hovingh (Salt Lake City, Utah), E. Johannes (Deixis Consult-
ants, Seattle), D. Sada (Bishop, California), S. Welty (Dubois,
Wyoming), and G. Vinyard (University of Nevada, Reno). We
thank E. Coan (CAS), A. Kabat (Invertebrate Zoology,
National Museum of Natural History, Smithsonian Institution
(NMNH)), F. Naggs, and G. Steyskal (FSM) for their input
pertaining to taxonomic, nomenclature!, or other problems in
this genus. We also thank two anonymous reviewers for their
comments on the manuscript.
Scanning electron micrographs were taken by Susanne
Braden, Scanning Electron Microscopy Laboratory (NMNH),
and prints of these were prepared by Victor Krantz and the staff
of the Office of Printing and Photographic Services (NMNH).
Victor Krantz photographed and prepared prints of shells.
Maps and illustrations were prepared by M. Ryan (Invertebrate
Zoology, NMNH) and S. Escher (Front Royal, Virginia).
Partial support for fieldwork and laboratory studies was
provided by awards to the first author from the Smithsonian
Institution Research Opportunities Fund and Scholarly Studies
Program. The second author's California fieldwork in part was
supported by the California Department of Fish and Game.
Character Descriptions
A preliminary elucidation of phylogenetic relationships
within Fluminicola was derived using a data set of 27
characters (56 character states) for the seven congeners for
which anatomical material was available, plus three outgroups.
We used as the prime outgroup (occupying basal positions on
generated trees) the genus Lithoglyphus (as exemplified by its
type species, Paludina naticoides Pfeiffer, 1828), to which
Fluminicola has been most closely allied in the recent
literature. Data for L. naticoides were obtained from the
literature (as referenced below) and from study of a single lot of
relaxed anatomical material (River March, Austria, USNM
883415). We used as additional outgroups the type species of
the eastern North American lithoglyphine genera Gillia
Stimpson, 1865a (Melania altilis Lea, 1841), and Somatogyrus
Gill, 1863 (Amnicola depressa Tryon, 1864a). Data for these
species were obtained from Thompson (1984) and from the
study of available anatomical material (G. altilis, Lake
Waccamaw, Columbus County, North Carolina, UF 27550; 5.
depressus, Cedar River, near Osage, Mitchell County, Iowa,
USNM 883757). Character states found in the prime outgroup
were treated as plesiomorphic (i.e., if also found in Flumini-
cola). Multistate characters were treated as unordered.
NUMBER 583
Characters are discussed below. Derived states are illustrated
in association with the species description section unless
otherwise stated.
SHELL.?Characters are protoconch microsculpture (charac-
ter 1) and shell umbilicus (character 2).
Protoconchs are variable within Fluminicola, with sets of
species having either of two conditions involving spiral
microsculpture. Two small species (F. dalli, F. turbiniformis)
have protoconchs sculptured with weakly developed spiral
striae (state 1), whereas Gillia, Somatogyrus, and other
Fluminicola have strong spiral lines (state 2). Lithoglyphus, by
contrast, has a completely smooth protoconch (Falniowski,
1990, figs. 121, 122). In this analysis, we treat both of the
conditions seen in Fluminicola as derived because the smooth
protoconch of the outgroup also is considered primitive among
hydrobiid snails generally (Falniowski, 1991). All Fluminicola
species except F. coloradensis and F. fuscus share a derived
condition in which the umbilical region often has a well-
developed axial ridge; the outgroups and latter two species
have a simple umbilical region.
OPERCULUM.?Characters are thickening of attachment-scar
margin (character 3) and rim development along outer edge of
operculum (character 4).
Fluminicola modoci and F. seminalis (Hinds, 1842) share a
derived condition in which the scar margin is strongly
thickened. The outgroups and all other species have a weak or
unthickened attachment scar. A derived condition in which the
outer edge of the operculum has a well-developed rim (state 1)
is shared by F. coloradensis and F. fuscus (plus Gillia and
Somatogyrus). Other species of Fluminicola and Lithoglyphus
have a weakly developed rim (state 0) (or lack one entirely).
CENTRAL RADULAR TEETH.?Characters are dorsal indenta-
tion of tooth (character 5), width of central cusp (character 6),
and number of basal cusps (character 7).
Fluminicola coloradensis and F. fuscus share a derived
condition in which the dorsal edge of the central tooth is only
slightly concave. The outgroups and other Fluminicola have a
moderate to strong dorsal indentation. These same two species
also share with F. seminalis a derived condition in which the
central cusp is very broad. The outgroups and other Flumini-
cola have a narrow to medium-wide central cusp. Fluminicola
dalli and F. turbiniformis share a derived condition in which
only a single pair of basal cusps is present on the central tooth.
The outgroups and other Fluminicola have at least two pairs of
basal cusps.
LATERAL RADULAR TEETH.?Characters are width of
central cusp (character 8) and tooth shaft length (character 9).
As with the central tooth, F. coloradensis and F. fuscus share
a derived condition in which the central cusp is especially
broad. The outgroups and other Fluminicola have a narrow to
medium-wide central cusp. Five species (F. dalli, F. modoci, F.
seminalis, F. turbiniformis, F. virens) plus Gillia and Somato-
gyrus have a derived condition in which the length of the lateral
tooth shaft is greater than the height of the tooth face (state 1).
Lithoglyphus and other Fluminicola have a shaft that is shorter
than the tooth face (state 0).
MARGINAL RADULAR TEETH (character 10).?Three species
(F. dalli, F. modoci, F. turbiniformis) have a derived condition
of numerous cusps (>20) on the inner marginal teeth, whereas
the outgroups and other Fluminicola have no more than 17
cusps on this tooth.
CEPHALIC TENTACLES.?Characters are length of tentacles
relative to snout (character 11) and dorsal pigmentation of
tentacles (character 12).
Short, almost stubby tentacles (Figure lA-D) represent a
synapomorphy for the group consisting of Fluminicola, Gillia,
and Somatogyrus. Lithoglyphus, by contrast, has tentacles that
are much longer than the snout (Krause, 1949, fig. 4). All
Fluminicola (plus Somatogyrus) except F. virens have a
derived condition in which dorsal tentacle pigmentation is
diffuse or uniform (not illustrated). Lithoglyphus, Gillia, and F.
virens have a completely different pigment pattern consisting
of an elongate, longitudinal band on each tentacle.
CTENIDIUM.?Characters are posterior extent of ctenidium
relative to pericardium (character 13) and condition of faces of
ctenidial filaments (character 14).
Fluminicola dalli and F. turbiniformis have a derived
condition in which the ctenidium ends well anterior to the
pericardium (see Ponder et al., 1993, fig. 2a for illustration of
this state). In the outgroups and in other Fluminicola, the
ctenidium overlaps the pericardium posteriorly (Figure 2).
Three species (F. dalli, F. seminalis, F. turbiniformis) have a
derived condition in which the faces of the ctenidial filaments
are smooth (not illustrated). The outgroups and other Flumini-
cola have prominent pleats on the faces.
HYPOBRANCHIAL GLAND (character 15). Fluminicola dalli
and F. turbiniformis have a derived condition in which the
posterior portion of the hypobranchial gland is thin (not
illustrated). The outgroups and other Fluminicola have a
pronounced posterior swelling of the hypobranchial gland.
CEPHALO-PEDAL GANGLIA.?Characters are length of pedal
commissure (character 16) and pigmentation of ganglia
(character 17).
All Fluminicola except F. virens have derived conditions in
which the pedal commissure is elongate relative to the width of
the pedal ganglia (i.e., more than 50% of ganglion width), and
in which all cephalo-pedal ganglia are pigmented (neither state
illustrated). The outgroups and F. virens have a much shorter
commissure and have unpigmented ganglia.
TESTIS (character 18).?A relatively small testis, which fills
no more than 67% of the length of the visceral coil behind the
stomach, is a derived condition shared by Fluminicola and
Somatogyrus (not illustrated). Lithoglyphus and Gillia, by
contrast, have an elongate testis that extends virtually to the
apical tip of the animal. (The very small testis diagramatically
illustrated by Krause (1949, fig. 22) for L. naticoides is an
error.)
PENIS.?Characters are penis shape (character 19) and
folding of dorsal surface (character 20).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 1.?Critical-point-dried tentacles and penes of Fluminicola species. A-D, tentacles, F. coloradensis,
USNM 883493: A, left tentacle, dorsal surface (bar = 0.33 mm); B. left tentacle, ventral surface (bar = 0.33 mm);
C. right tentacle, dorsal surface (bar = 0.33 mm); D. right tentacle, ventral surface (bar = 0.33 mm). EJ=. penis, F.
coloradensis, USNM 883493: E, distal penis (bar = 50 um); F. distal tip (bar = 15 um). Gtf, penis, F. virens,
USNM 883673: G. dorsal aspect (bar = 0.38 mm); H, distal tip (bar = 100 urn).
NUMBER 583
Cgo
Pc Ct
FIGURE 2.?Contents of pallial cavity of Fluminicola coloradensis. USNM
883493 (bar= 1.0 mm). Pallial roof has been separated from the head-foot
Note the large pallial portion of the renal organ and the ctenidium overlapping
the pericardium posteriorly. The hypobranchial gland covers the rectum and
capsule gland (except for its opening) and is swollen posteriorly (lighter
stipple). (Cgo = capsule gland opening; Ct = ctenidium, Hg = hypobranchial
gland, Os = osphradium, Pc = pericardium, Pe = pallial edge; R = renal organ;
Ro = opening of renal organ.)
Penial shape is variable within Fluminicola. All but two
species have a sickle-shaped penis that is a derived state (state
1); F. dalli has a uniquely elongate penis that also is derived
(state 2). The outgroups (Bole, 1982, fig. 4; Thompson, 1984,
fig. 51; Falniowski, 1987, fig. 31) and F. virens have a stout,
broadly triangular penis that is nearly straight. A very weakly
folded or smooth dorsal surface is a derived condition shared
by most species of Fluminicola. The outgroups and three of the
larger species (F. coloradensis, F. fuscus, F. virens) have a
strongly folded surface.
OVARY (charater 21).?A very small ovary, filling less than
25% of the length of the visceral coil behind the stomach, is a
derived condition shared by F. dalli and F. turbiniformis (not
illustrated). The outgroups and other Fluminicola have a much
larger ovary that fills 33%-50% of the visceral coil behind the
stomach.
BURSAL DUCT.?Characters are insertion of duct into bursa
copulatrix (character 22) and length of duct (character 23).
Three species (F. modoci, F. seminalis, F. virens) have a
derived condition in which the bursal duct inserts into or near
the middle of the long axis of the bursa copulatrix. In the
outgroups and in other Fluminicola, the duct inserts into the tip
of the long axis of the bursa copulatrix. All Fluminicola (plus
Gillia) except F. coloradensis and F. virens have a derived
condition consisting of a short bursal duct (relative to length of
bursa copulatrix). The latter two species as well as Litho-
glyphus and Somatogyrus have a relatively elongate bursal
duct.
SEMINAL RECEPTACLE (character 24).?Three species (F.
modoci, F. seminalis, F. turbiniformis) share a derived
condition in which the seminal receptacle overlaps the bursa
copulatrix. In the outgroups and in other Fluminicola, the
seminal receptacle is anterior to the bursa copulatrix.
GLANDULAR OVIDUCT.?Characters are development of
rectal furrow (character 25), shape of genital opening (character
26), and development of anterior vestibule (character 27).
Three species (F. coloradensis, F. fuscus, F. seminalis) share
a derived condition consisting of a deep rectal furrow along the
length of the glandular oviduct. In the outgroups and in other
Fluminicola, the furrow is either very shallow or absent.
Fluminicola coloradensis and F. fuscus share a slit-like genital
opening and a well-developed anterior vestibule that are
derived character states. The outgroups and other Fluminicola
have a simple, pore-like genital opening and have a weakly
developed or absent anterior vestibule.
Phylogenetic Reconstruction
The small size of the data set permitted use of an exact
method of calculating trees (the "ie" option of HENNIG86),
which yielded two fully resolved and maximally parsimonius
trees of 41 steps that have a consistency index of 0.70 and a
retention index of 0.75. One of these trees is shown in Figure 3.
The other tree was identical in topology except that the
near-basal branches that terminate in F. virens and Gillia altilis
were swapped.
Homoplasy and/or reversals are associated with 11 of the 27
characters. For eight characters (2, 6, 10, 14, 22-25), a single
parallelism is involved; for one character (9), a single reversal
is involved. Character 4 exhibits two reversals on the illustrated
tree, but it has only a single reversal on the second tree.
Character 18 is a synapomorphy on the illustrated tree, but it is
reversed on the second tree.
The cladistic analysis reveals that the North American
lithoglyphines that we studied are distinguished from European
Lithoglyphus by two nonhomoplastic synapomorphies (proto-
conch with spiral microsculpture, short cephalic tentacles).
Among the North American lithoglyphines, western American
species (allocated to Fluminicola) do not comprise a mono-
phyletic group. Note, however, that with the exception of F.
virens, other members of Fluminicola do comprise a clade
united by two nonhomoplastic synapomorphies (elongate pedal
commissure, sickle-shaped or elongate penis).
Within this group of six species, F. coloradensis and F.
fuscus comprise a basal clade united by four nonhomoplastic
synapomorphies (weak dorsal indentation of central radular
teeth, broad central cusp of lateral radular teeth, slit-like genital
opening, well-developed anterior vestibule of capsule gland).
The other four species are united by a single nonhomoplastic
synapomorphy (a weakly folded or smooth dorsal penis).
Within this group, F. dalli and F. turbiniformis comprise a
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
23
1 4 9 11
HHHHH
2 1 1 1
2 4 22
1 0 1
12 17
+
1 1
16 19
++
1 1
5 6 8 9 25 26 27
4Hh-HHh+-H
1 1 1 0 1 1 1
naticoides
altilis
virens
depressus
coloradensis
fuscus
2 4 20 23
HHHHH
1 0 1 1
seminalis
FIGURE 3.?Cladogram for Fluminicola species plus three outgroups {Uthoglyphus naticoides, Gillia altilis, and
Somatogyrus depressus). White bars indicate character state transformations supporting nodes, with character
number given above and state indicated below. Black bars indicate unreversed synapomorphies.
well-differentiated clade united by five nonhomoplastic
synapomorphies (weak spiral lines on shell protoconch, single
pair of basal cusps on central radular teeth, short ctenidium,
thin posterior portion of hypobranchial gland, small ovary).
Fluminicola modoci and F. seminalis comprise a second clade
supported by a single nonhomoplastic synapomorphy (strongly
thickened opercular attachment-scar margin).
Super-family RiSSOOIDEA Gray, 1847
Family HYDROBIIDAE Troschel, 1857
Subfamily LITHOGLYPHINAE Troschel, 1857
Genus Fluminicola Carpenter, 1864
Fluminicola Carpenter, 1864:676 [type species: Paludina nuttalliana Lea,
1838; original designation.]
Heathilla Hannibal, 1912:186 [type species: Paludina seminalis Hinds, 1842;
original designation.]
Fluminicola is a morphologically diverse assemblage of
northwestern North American lithoglyphine snails that vary
widely in size. This group, as currently constituted, is probably
paraphyletic (see "Phylogenetic Reconstruction," above) and is
not readily distinguishable from several eastern North Ameri-
can lithoglyphine genera, hence a diagnosis is not provided
herein.
DESCRIPTION.?Shell usually globose to trochoid, rarely
ovate- or narrow-conic; height 2.5-12.0 mm; whorls 3.5-5.0.
Protoconch of 1.25-1.50 whorls; diameter 0.6-1.0 mm;
microsculpture of numerous spiral striae. Tfeleoconch whorls
convex or (rarely) nearly flat; whorls shouldered, sometimes
with broad sutural shelf. Microsculpture of collabral growth
lines and occasional weak spiral striae or scratches. Perios-
tracum brown, green, tan, or red. Shell opaque-translucent,
clear white, grey, or pink. Aperture usually large, ovate-lunate,
rounded below, narrowed above. Outer lip prosocline, straight
or weakly sinuate, usually thin. Parietal lip usually complete,
variably thickened, adnate or slightly detached. Columellar lip
usually thick; columellar swelling usually broad, often cover-
ing umbilical region. Umbilical region sometimes excavated,
with adaxial ridge. Shell anomphalous or with small umbilicus.
Operculum ovate-ellipsoidal, amber, of thin to medium
thickness, nearly flat, dorsal surface frilled, nucleus eccentric.
Outer margin of operculum sometimes with obvious rim.
Attachment-scar margin variably thickened. Callus nearly
absent to strongly developed.
Radula with about 60-90 rows of teeth; ribbon elongate;
central tooth width 30-100 urn. Central tooth trapezoidal;
dorsal edge weakly to strongly indented; lateral cusps 2-5;
median cusp dagger or U-shaped, enlarged relative to laterals;
basal cusps 1-3, originating from lateral angles or tooth face,
NUMBER 583
innermost cusp largest; basal tongue well developed, basal
sockets usually well excavated; lateral angles thickened,
inclined at about 45?-50?,not extending beyond base of tooth.
Lateral tooth with flexed shaft; shaft as long as or longer than
height of tooth face; dorsal edge of tooth convex to concave;
cutting edge comprising 35%-60% of tooth width; tooth face
usually taller than wide; shaft inclined about 30?-40?; basal
tongue knob-like; central cusp U-shaped, lateral cusps 2-5.
Inner marginal teeth with 7-27 cusps; outer marginal teeth
with 6-29 cusps.
Epithelial body pigmentation usually well developed. Tenta-
cles usually with pale to light patch surrounding eyes. Pallial
roof and visceral coil usually black.
Snout squat; distal lobation pronounced. Tentacles short,
parallel sided, and well ciliated (more so on ventral surfaces)
but without well-defined tracts; eyelobes weakly developed.
Foot narrowly ovate, anterior end convex, lateral wings well
developed, posterior end rounded; anterior mucous gland of
numerous similar-sized glands.
Ctenidium extending almost entire length of pallial cavity,
often overlapping pericardium posteriorly; filaments about
16-40, well developed, broadly to narrowly triangular, usually
pleated, apices on right, free edge usually concave. Osphradium
usually narrow, about 30%-50% of ctenidium length, centered
posterior to middle of ctenidial axis, anterior end simple or
weakly hooked. Hypobranchial gland thin on pallial roof,
thicker on rectum and genital ducts, often swollen proximally.
Rectum nearly straight, broadly overlapping genital ducts, fecal
pellets usually longitudinal; anus near pallial edge, slightly
anterior to anterior end of capsule gland. Renal organ with
40%-60% of length in pallial roof, sometimes greatly swollen;
renal gland horizontal, on left side of renal organ; renal opening
small, usually white. Pericardium slightly bulging into pallial
cavity. Buccal mass filling length of snout. Salivary glands
simple, narrow, usually overlapping cerebral ganglia. Stomach
about as long as style sac; posterior caecum absent. Cerebral
and pedal commissures usually elongate (more than 50% of
ganglia width); right pleural and supraoesophageal ganglia
abutting. Ganglia usually pigmented. Body spaces with
relatively little connective tissue.
Testis more than 1.0 whorl, of multiple branches connected
by narrow vas efferens and bearing grape-like clusters of lobes,
usually overlapping posterior stomach chamber and filling
about 50% of visceral coil behind stomach. Seminal vesicle of
numerous coils, abutting stomach beneath anterior 25%-50%
of testis. Prostate gland with about 33% of length in pallial
roof, bean-shaped, oval in section, of complexly folded cells;
vas deferens opening just in front of pallial wall near anterior
end of gland. Vas deferens straight or slightly undulating,
rarely coiled in pallial roof and neck. Penis of variable size,
narrowly elongate to sickle-shaped, weakly curved to tightly
coiled, unlobed, surface variably folded, usually gently
tapering along basal and medial portions; distal portion either
simply tapering or with subterminal constriction followed by
pointed tip or papilla. Dorsal penis usually moderately ciliated
(rarely unciliated); cilia often dense near distal tip. Penial duct
near center, muscular, of narrow to medium width, often
undulating along most of length. Dorsal penis lightly to darkly
pigmented along most of length; subepithelial pigment usually
well developed along length of penial duct.
Females oviparous; egg capsules large, hemispherical
(Stimpson's (1865b, fig. 17) early illustration of compound egg
capsules was obviously based on pulmonate material). Ovary
of compound lobes, 0.25-1.0 whorl, positioned behind
posterior edge of stomach and filling up to 50% of visceral coil
behind stomach. Coiled oviduct usually having inverted
U-shape, positioned slightly behind pallial wall and anterior to
bursa copulatrix; usually unpigmented, bound in connective
tissue, surface of irregular appearance; posterior arm of primary
coil often swollen with sperm; proximal portion usually with
1-2 small secondary twists or coils; distal portion short,
straight. Coiled oviduct and bursal duct join slightly to well
behind pallial wall; oviduct opening to albumen gland
ventrally. Bursa copulatrix> about 33%-67% of albumen gland
length, ovate-pyriform, rarely globular, positioned on right side
of albumen gland, part or most of length posterior to gland,
long axis perpendicular to gland. Bursal duct short to elongate,
narrow, deeply imbedded in albumen gland, issuing from
ventral tip to near midpoint of long axis of organ. Seminal
receptacle small, pouch-like, usually unpigmented; positioned
on left side of albumen gland (although edge often partly or
entirely overlapped by gland) near ventral edge, usually at or
near posterior edge, anterior to or slightly overlapping bursa
copulatrix; duct very short. Pallial gonoduct of complexly
folded cells and with shallow to deep rectal furrow. About
17%-50% of albumen gland lying in front of posterior pallial
wall. Capsule gland of 2 tissue sections (anterior section much
shorter), usually narrower than albumen gland. Ventral channel
sometimes with short anterior vestibule. Genital opening a pore
or short slit, subterminal to terminal.
DISTRIBUTION AND ECOLOGY.?Northwestern North Amer-
ica, including portions of the northern Great Basin, Snake-
Columbia River system, Sacramento River system, and Pacific
coastal drainages (British Columbia, California, Idaho, Ne-
vada, Oregon, Utah, Washington, Wyoming). Early Yellow-
stone River and South Dakota records (Henderson, 1935,
1936a) require confirmation.
Fluminicola are usually found in clear, cold waters with high
dissolved oxygen content. Large species are typically found in
streams, whereas smaller species are commonly found in either
spring or stream environments. Many taxa are apparently
lithophiles and graze on periphyton. Often, species in this
genus appear to be community dominants, comprising most of
the invertebrate biomass. Fluminicola are fairly intolerant of
impounded waters and soft substrates as well as of nutrient-
enhanced or lacustrine habitats. Although the free-flowing,
oligotrophic waters where these animals thrive were once
ubiquitous throughout the historic range of the genus, the genus
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
is apparently now extirpated from large areas (e.g., much of the
Columbia and Snake river drainages and the Olympic
Peninsula) owing to human-related activities. Note that only
relatively small reaches of the Columbia River system remain
free flowing.
FOSSIL RECORD.?Based on fossil species that may belong
to this genus (Appendix), a Miocene (or earlier) origin is
inferred. Taylor (1966b: 160) observed that several Recent
species are known from the Pliocene.
SPECIES OF UNCERTAIN STATUS.?Taylor (1975a) allocated
Bithynia perfecta Frauenfeld, 1862 (Recent, Columbia, North
America; see Frauenfeld, 1865, pi. IX: unnumbered fig.), and
its subspecies dubitata (see Frauenfeld, 1865, pi. IX: unnum-
bered fig.) to Lithoglyphus, presumably on the assumption that
the type localities referred to British Columbia. Type material
for these taxa (MNW uncat.) do not closely resemble any North
American material for Fluminicola that we have seen and,
instead, they are more similar to South American Potamolithus
Pilsbry in Pilsbry and Rush, 1896. We reserve further judgment
of these taxa pending collection and study of anatomical
material.
REMARKS.?Carpenter (1864:676) introduced Fluminicola
in reference to an unpublished work of Stimpson {"Fluminicola
[Stimpson, MS] nuttallii") and did not include a description.
Nevertheless, his genus was validly described as he provided an
indication by using the name in combination with an available
species-group name (International Commission on Zoological
Nomenclature (ICZN), Article 12b[5]). Authorship of this
genus name has invariably been attributed in the literature to
Stimpson (1865a:52-53), who provided a full description.
Carpenter based the genus on the same type species as did
Stimpson, and the concept of Fluminicola is not altered as a
result of this authorship change.
Hannibal (1912) distinguished his subgenus Heathilla by its
more globose shell, shorter spire, more rapid teleoconch
expansion rate, and shallower sutures; he placed five species in
this group (F. columbiana, F.fusca, F. modoci, F. seminalis, F.
virens). This assemblage actually encompasses a much broader
range of shell morphologies than implied above, and there is no
justification for such a subgeneric grouping based on our
phylogenetic hypothesis.
Yen (1944) distinguished fossil Pilsbryus (type species
Lithasia antiqua Gabb, 1866; original designation) from
Fluminicola (and Lithoglyphus) by its smaller, more depressed
spire, retreating outer lip, thicker parietal and columellar lips,
naticoid appearance, and thicker shell. Although Taylor
(1966b) later placed Pilsbryus as a junior synonym of
Lithoglyphus (including Fluminicola), we concur with Yen's
assessment and accept the former as distinct
Stimpson (1865b) placed Fluminicola in his broadly
conceived subfamily Hydrobiinae. Tryon (1866), in a review of
Stimpson's paper, instead advocated placement of the genus in
the Lithoglyphinae, which has been accepted by most
subsequent authors (including Fischer, 1883; Walker, 1918;
Taylor, 1966b; Burch 1978, 1979; Thompson, 1984; Vaught,
1989). Clessin (1880) erected the Fluminicolinae to accomo-
date Fluminicola and similar-shelled taxa (including Li-
thoglyphus). This taxon has been little used in the literature
(but see Starobogatov, 1970) and is now considered
equivalent to the Lithoglyphinae (Ponder and Ware"n, 1988).
Thiele (1928, 1929) instead placed Fluminicola in the
Amnicolinae, which was accepted by Wenz (1939).
Thompson (1984) reviewed the North American litho-
glyphine genera and postulated a close relationship between
Fluminicola and Gillia based on a phenetic analysis (Th-
ompson, 1984, fig. 58, table 2). Characters linking these two
genera were large size, narrowly umbilicate shell, penial
papilla, large central cusps of central and lateral radular teeth,
and vertical shaft of the lateral radular tooth. Note, however,
that these features are not characteristic of most species now
assigned to Fluminicola: e.g., only F. virens has a (eversible)
penial papilla, and the shaft of the lateral tooth is flexed (not
vertical) in all of these species.
Fluminicola coloradensis Morrison, 1940
Fluminicola fusca.?Binney, 1865:92 [in part].?Ingersoll, 1875:127 [in
part].?Pilsbry, 1899a:123 [in part].?Orcutt 19Ola:35 [in part].?
Henderson, 1924:192 [in part].?Chamberlin and Jones, 1929:180 [in
part].?Henderson, 1935:107 [in part]; 1936a:139 [in part].?Beetle, 1961:5.
Fluminicola coloradoense Morrison, 1940:125 [in part; holotype: USNM
526631; type locality: Green River, Wyoming].?Taylor, 1966a:34, fig. 14
[as synonym of L. hindsi].?Rosewater, 1984:3.?Beetle, 1989:639 [as
synonym of L. hindsi].
Lithoglyphus hindsii [sic].?Taylor, 1966a:34, fig. 14 [in part].
Lithoglyphus coloradoensis.?Taylor, 1975a:60.
Fluminicola hindsi.?Burch and Tottenham, 1980:102 [in part].
Lithoglyphus hindsi.?Taylor, 1985:306 [in part].?Beetle, 1989:639 [in part].
DIAGNOSIS.?Large with subglobose to broadly conical
shell; penis of medium size, sickle-shaped.
A pigmented seminal receptacle (and occasionally a pig-
mented coiled oviduct) is unique to this species among
members of the genus. Fluminicola coloradensis is further
distinguished from sister taxon F.fuscus by the grey white shell
color, absence of either pronounced subsutural angulation or
keel on body whorl, weaker operculum callus, frequent
presence of third pair of basal cusps on central radular teeth,
bulbous distal penis, longer bursal duct, and greater overlap of
seminal receptacle by albumen gland.
DESCRIPTION.?Shell (Figures 4A, 5A-D) subglobose to
broadly conical; height 6.5-10.8 mm; whorls 3.5-4.5. Proto-
conch (Figure 6A) of 1.5 whorls; diameter about 1.0 mm;
microsculpture of numerous, sometimes anastomosing, strong
spiral striae. Teleoconch whorls convex, shouldered, often with
broad sutural shelf. Microsculpture of collabral growth lines
and occasional weak spiral striae or scratches. Periostracum
tan. Shell opaque, grey white. Aperture large, broadly lunate,
narrowed above. Outer lip usually thick. Parietal lip complete,
usually thick, adnate or rarely slightly detached. Columellar lip
NUMBER 583
thick, columellar swelling broad, often covering umbilical
region. Shell anomphalous, cryptomphalous, or with small
umbilicus.
Operculum (Figure 7A) of medium thickness; outer margin
with obvious rim. Outer edge of attachment-scar margin
slightly thickened, inner edge unthickened. Callus weak or
absent.
Radula (Figure 8A-C) with about 65 rows of teeth; ribbon
length 3.0 mm, ribbon width 0.3 mm; central tooth width 95
um. Central tooth with slight dorsal indentation; lateral cusps
2-3; median cusp broadly U-shaped, slightly broader and
longer than laterals; basal cusps 2-3, narrow, originating near
dorsal edge of lateral angles; basal tongue medium wide, basal
sockets moderately excavated; lateral angles thickened. Lateral
tooth with convex dorsal edge, very slightly indented centrally;
dorsal edge about 50% of tooth width; lateral shaft shorter than
height of tooth face; tooth face taller than wide; central cusp
broadly U-shaped, lateral cusps 3 (sometimes 4 on outer side).
Inner marginal teeth with 9-11 cusps; outer marginal teeth
with 10-11 cusps.
Brown black pigment well developed on snout and tentacles
(except for light patch around and slightly posterior to eyes).
Head light grey, foot grey black and often dark along anterior
and posterior edges. Pallial roof and visceral coil black.
Contents of pallial cavity shown in Figure 2. Ctenidium
overlapping pericardium; filaments about 38, tall, pleated.
Osphradium about 40% of ctenidium length. Hypobranchial
gland swollen on proximal genital duct. Fecal pellets oblique,
almost perpendicular. Renal organ greatly swollen; renal
opening white. Salivary glands terminating anterior to cerebral
ganglia. Stomach about as long as style sac. Ganglia
pigmented.
Testis 1.5 whorls, overlapping posterior stomach chamber,
filling more than 50%-67% of visceral coil behind stomach.
Prostate gland with 33% of length in pallial roof. Vas deferens
nearly straight in pallial roof and in neck. Penis (Figure 9A) of
medium size, sickle-shaped, usually very tightly coiled, with
strong folds along proximal 67% of length; base slightly
narrowed; medial section gently tapering; bulbous distal
section narrowing considerably to short, pointed tip (Figure
lE.F). Penial duct near center, of medium width,
strongly undulating in basal and medial sections. Penis usually
with dark epithelial pigment on proximal half of dorsal surface,
rarely pale; black subepithelial pigment scattered along length
of penial duct.
Ovary 1.0 whorl, abutting posterior edge of stomach, filling
about 50% of visceral coil behind stomach. Distal female
genitalia shown in Figure 10A. Coiled oviduct narrow,
sometimes weakly pigmented; proximal portion usually with
1-2 small horizontal twists or coils; distal portion of primary
coil sometimes containing sperm. Coiled oviduct and bursal
duct join just behind pallial wall. Bursa copulatrix about 50%
of albumen gland length, pyriform, about as long as wide, with
central section decidedly narrowed, positioned largely posterior
to gland. Bursal duct about 67% of bursa length, originating
slightly lateral to tip of organ. Seminal receptacle about 33% of
bursa copulatrix length, lightly pigmented, positioned anterior
to bursa copulatrix near posterior edge of albumen gland, partly
or entirely overlapped by gland. Pallial gonoduct with deep
rectal furrow. About 17% of albumen gland lying in front of
posterior pallial wall. Capsule gland about as long and as wide
as albumen gland. Ventral channel with anterior vestibule.
Genital opening a subterminal slit fringed by small papilla.
DISTRIBUTION.?Upper Green River drainage, Wyoming
(Figure 11). We have been unable to confirm the Emery
County, Utah, record, which is well downflow from other
Green River sites for the species and may represent drift
material (shells in this lot are worn and were collected empty).
REMARKS.?Morrison (1940) described this species to
accommodate snails not only from the Green River drainage
but also from the Great Basin of Utah. Although shells of these
two groups of populations are generally similar in form, a
preliminary survey of available alcohol material revealed
significant differences in penial morphology between them,
suggesting that the Great Basin populations comprise one or
more undescribed species. Pending further study of this
problem, we have decided to restrict F. coloradensis to
populations from the Green River. Anatomical data are from
USNM 883493.
MATERIAL EXAMINED.?UTAH. Green River, UF 78511.
Head of Green River, USNM 28103, USNM 120462. ?Emery
County: Green River, Hideout, FMNH 178501.
WYOMING. ANSP 88598. Wyoming Territory, USNM
28538, USNM 170789. Green River, USNM 526631 (ho-
lotype), USNM 526576 (paratypes), USNM 159396, USNM
160394. Green River, Blacks Fork, Millersville, ANSP 27769.
Lincoln County: Hams Fork, Green River, Bells Fish Cliff,
USNM 526754. Green River, Smiths Fork, ANSP 27773.
Sublette County: Green River at bridge, HWY 359, 12.8 km
north of Place Cafe, USNM 883191*. Green River, HWY
187-189 bridge, USNM 883493*.
Fluminicola dalli (Call, 1884)
Amnicola dalli Call, 1884:21,45-47, figs. 2. 3; pi. VI: figs. 4-6 [lectotypc:
MCZ 2087; type locality: small tributary to Pyramid Lake, near the north
end, at Symons Ranch].?Call, 1886:2-4.?Orcutt 1901 b:47.?Taylor,
1966a:24, fig. 9 [as synonym of L. turbiniformis].?Johnson, 1975:142
[lectotype selection].
Fluminicola seminalis dalli.?Pilsbry, 1899a: 123.?Walker. 1918:142.
Lithoglyphus dalli.?Taylor, 1975a:71.
DIAGNOSIS.?Small with broadly conical shell; penis of
medium size, nearly vermiform.
The broad osphradium, nearly vermiform penis, and absence
of penial folds are unique to this species within the genus.
Fluminicola dalli is further separable from sister taxon F.
turbiniformis by the weaker operculum callus, darker body
pigmentation, larger penis, simply tapering distal penis, smaller
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
bursa copulatrix, greater overlap of albumen gland onto bursa
copulatrix, more anteriorly positioned seminal receptacle, and
greater overlap of albumen gland onto seminal receptacle.
DESCRIPTION.?Shell (Figures 4B, 5E-G) broadly conical,
with low spire; height 2.1-3.4 mm; whorls 3.5-4.0. Proto-
conch (Figure 6B) of 1.25 whorls, diameter about 0.6 mm;
microsculpture of weak, anastomosing spiral striae. Teleo-
conch whorls of medium convexity, sometimes with very weak
peripheral angulation; shoulders pronounced but narrow.
Microsculpture of collabral growth lines and weak spiral striae.
Periostracum light brown or green. Shell translucent, clear
white. Aperture large, ovate, weakly angled above. Outer lip
thin. Parietal lip complete, thick, broadly adnate to slightly
detached. Columellar lip thick, columellar swelling broad.
Shell shallowly perforate to broadly rimate; umbilical region
sometimes narrowly excavated, with adaxial ridge.
Operculum (Figure 7B) thin; rim along outer margin very
weak or absent. Attachment-scar margin slightly thickened
along inner edge to nucleus; outer edge thickening very weak
or absent. Callus moderately developed.
Radula (Figure 8D-F) with about 81 rows of teeth; ribbon
length 1.2 mm, ribbon width 0.15 mm; central tooth width 38
|im. Central tooth with pronounced dorsal indentation; lateral
cusps 3-5; median cusp narrowly U-shaped, slightly broader
and considerably longer than laterals; basal cusps 1 (rarely with
faint suggestion of second cusp), broadly triangular, arising
from tooth face near intersection with lateral angle; basal
tongue medium wide, basal sockets moderately to strongly
excavated; lateral angles thickened. Lateral tooth with slightly
concave dorsal edge; dorsal edge about 35% of tooth width;
lateral shaft considerably longer than height of tooth face; tooth
face taller than wide; central cusp narrowly U-shaped, lateral
cusps 2 or 3 (inner side) to 3 or 4 (outer side). Inner marginal
teeth with 24-25 cusps; outer marginal teeth with 22 cusps.
Snout and head nearly colorless to medium grey. Tentacle
pigment as for snout, with pale patch surrounding eyes. Foot
pigment as for snout, darkest along anterior edge. Pallial roof
and visceral coil black.
Ctenidium not overlapping pericardium; filaments 16,
narrow, without pleats. Osphradium broad, about 30% of
ctenidium length. Hypobranchial gland uniformly thin. Fecal
pellets longitudinal. Renal organ not swollen; renal opening
opaque. Salivary glands overlapping cerebral ganglia. Stomach
slightly longer than style sac. Ganglia pigmented.
Testis 1.0 whorl, abutting posterior edge of stomach, filling
slightly less than 50% of visceral coil behind stomach. Prostate
gland small, narrow, with 30% of length in pallial roof. Vas
deferens undulating in pallial roof, straight in neck. Penis
(Figure 9B) of medium size, nearly vermiform, slightly
curved, without folds; penis tapering gently throughout to
simple distal tip. Penial duct slightly to right of midline,
narrow, gently undulating in proximal 67%. Penis without
epithelial pigment; subepithelial pigment restricted to distal
50%.
Ovary 0.25 whorl, positioned behind stomach, filling about
20% of visceral coil behind stomach. Distal female genitalia
shown in Figure 10B. Coiled oviduct narrow to broad,
unpigmented; proximal portion with vertical loop or small
kink; distal portion of primary coil sometimes containing
sperm. Coiled oviduct and bursal duct join well behind pallial
wall. Bursa copulatrix about 36% of albumen gland length,
ovate, wider than long, largely overlapped by gland. Bursal
duct short (25% of bursa length), originating near tip of organ.
Seminal receptacle about 33% of bursa copulatrix length,
unpigmented, positioned slightly anterior to bursa copulatrix
near posterior edge of albumen gland, entirely overlapped by
gland. Pallial gonoduct with weak rectal furrow. About
22%-30% of albumen gland lying in front of posterior pallial
wall. Capsule gland slightly shorter, narrower than albumen
gland. Ventral channel without or (less commonly) with very
short anterior vestibule. Genital opening a terminal pore.
DISTRIBUTION.?Thus far known only from type locality
area, northern Pyramid Lake basin (Great Basin), Nevada
(Figure 11).
REMARKS.?The type locality is in Washoe County. Inas-
much as Call distributed type material to several institutions
and did not explicitly state that his figured specimen was a type,
we treat MCZ 2087 as a lectotype selected by Johnson (1975),
rather than as the identified holotype. Anatomical data are from
USNM 874899 (near topotypes).
MATERIAL EXAMINED.?NEVADA. Washoe County: Small
tributary to Pyramid Lake, near the north end, at Symons
Ranch, MCZ 2087 (lectotype). Near north end of Pyramid
Lake, MCZ 2088 (paralectotypes). Stream flowing into
Pyramid Lake, USNM 31269. Stream, north end of Pyramid
Lake, ANSP 27783, ANSP 92081, USNM 526835. Spring, 3.2
km west of Thunderbolt Bay (Pyramid Lake), USNM 874899*,
USNM 883467*.
Fluminicola fuscus (Haldeman, 1841)
Anculosa fusca Haldeman, 1841 [cover; type not located; type locality:
Oregon].
Leptoxis fusca.?Haldeman, 1847:4, pi. 3: figs. 83, 84.?Adams and Adams,
1854:307.?Binney, 1859:11; 1860:10.?Brot, 1862:24.
Anculotus fuscus.?Reeve, 1860-1861:11 [as synonym of A. nuttalii}.
Amnicola hindsi Baird, 1863:67 [lectotype: BMNH 1863.2.4.17A, selected
herein; type locality: River Kootanie].?Carpenter, 1864:604.?Binney,
1865:90 [as synonym of F. nuttalliana].?Pilsbry, 1899a: 123 [as synonym
of F. nuttalliana].?Dall, 1905:119 [as synonym of F. nuttalliana].?
Hannibal, 1912:187 [as synonym of F. virens].?Henderson, 1929:168 [as
synonym of F. nuttalliana].
Paludina hindsi.?Carpenter, 1864:676 [as synonym of F. nuttallii].?
Frauenfeld, 1864:613.
Fluminicola fusca.?Binney, 1865:92 [in part].?J.G. Cooper, 1867:30 ?
Tryon, 1870:66, pi. 17: fig. 19 [in part].?Call, 1884:21 [in part].?J.G.
Cooper, 1888:240 [in part].?Pilsbry, 1899a: 123 [in part].?Orcutt
1901a:35 [in part]; 19027:61 [in part].?Hannibal, 1912:187 [in part].?
Henderson, 1929:167 [as synonym of F. columbiana].?Morrison,
1940:125.?Baker, 1964:172.?Burch and Tottenham, 1980:102, fig. 141.?
Turgeon et al., 1988:60.?Beetle, 1989:639 [as synonym of L. hindsi].
NUMBER 583 11
Fluminicola hindsii [sic].?Stimpson, 1865a:53; 1865b:54.?Hemphill,
1881:12.
Fluminicola hindsi.?J.G. Cooper, 1867:30.?Orcutt 19Ola:35; 19027:61.?
Morrison, 1940:125.?Burch and Tottenham, 1980:102 [in part].?Turgeon
etal., 1988:60.
Fluminicola nuttaliana var. columbiana Hemphill, 1881:12 [nomen nudem].
Amnicola fusca.?Grasset, 1884:91.
Fluminicola nuttaliana columbiana Keep, 1887:63 [ex Hemphill MS; type not
located (see Coan, 1985); type locality: rivers of Oregon and Washington].?
Coan, 1985:212.?Coan and Roth, 1987:327.
Pomatiopsis fusca.?Paetel, 1888:471.
Fluminicola columbiana Pilsbry, 1899a: 125 [lectotype: ANSP 27767; type
locality: Columbia River, near Wallula].?Hannibal, 1912:188.?Walker,
1918:141.?Morrison, 1940:125.?Baker, 1964:172 [lectotype selection].?
Henderson, 1924:191; 1929:167-168 [in part]; 1936b:277? Burch and
Tottenham, 1980:102, fig. 145.?Turgeon et al., 1988:60.?Neitzel and
Frest, 1989:7, fig. 1.?USDI, 1991:58819.?Neitzel and Frest, 1993:9,
fig. 1.
Fluminicola nuttalliana hindsi.?La Rocque, 1953:270.
Lithoglyphus columbianus.?Taylor, 1966b: 160; 1975a:60.?Clarke,
1976:18.?USDI, 1984:21673.
Lithoglyphus hindsii [sic].?Taylor, 1966a:34, fig. 14 [in part]; 1975a:96.
Lithoglyphus fuscus.?Taylor, 1975a:86.
Lithoglyphus hindsi.?Clarke, 1981:70 [as synonym of L. virens].
DIAGNOSIS.?Large with subglobular to trochoid shell
having strong subsutural angulation or keel on body whorl;
penis of medium size, sickle-shaped.
In addition to body whorl shape, the frequently incomplete
parietal lip, pigmented salivary glands, and position of seminal
receptacle entirely lateral to albumen gland are unique to F.
fuscus within the genus.
DESCRIPTION.?Shell (Figures 4C,D, 12) subglobular to
trochoid, apex often eroded; height 7.0-11.2 mm; whorls
4.0-4.5. Protoconch (Figure 6c) of 1.5 whorls; diameter about
1.0 mm; microsculpture of numerous strong spiral striae.
Teleoconch whorls slightly convex, shouldered. Body whorl
with subsutural angulation to well-developed, thick ledge;
periphery well below middle of whorl, usually angulate,
sometimes strongly so. Microsculpture of strong collabral
growth lines. Periostracum tan, brown, or red. Shell opaque,
usually pink. Aperture large, ovate, narrowly angled to
pyriform above. Outer lip sometimes weakly sinuate (adapi-
cally advanced), usually thin. Parietal lip thin, often glaze-like,
sometimes incomplete across body whorl, rarely thick and
slightly detached. Columellar lip thick, columellar swelling
broad, often covering umbilical region, sometimes slightly
flared basally. Shell usually anomphalous, rarely with small
umbilicus.
Operculum (Figure 7c) of medium thickness; outer margin
with obvious rim. Outer edge of attachment-scar margin
slightly thickened, inner edge slightly to moderately thickened.
Callus well developed, crescent-like.
Radula (Figure 8G,I) with about 85 rows of teeth; ribbon
length 3.2 mm, ribbon width 0.3 mm; central tooth width 81
|im. Central tooth with slight dorsal indentation; lateral cusps
2-4; median cusp broadly U-shaped, slightly broader and
longer than laterals; basal cusps 1-2, narrow, originating near
dorsal edge of lateral angles; basal tongue narrow, basal sockets
weakly to moderately excavated; lateral angles broad, thick-
ened. Lateral tooth with convex dorsal edge; dorsal edge about
50% of tooth width; lateral shaft slightly shorter than height of
tooth face; tooth face taller than wide; central cusp U-shaped,
lateral cusps 3 (sometimes 2 on inner side). Inner marginal
teeth with 10-13 cusps; outer marginal teeth with 10-11
cusps.
Snout, including distal lips, dark purple brown; tentacles
similarly pigmented except for pale circle around eyes. Head
moderately pigmented; foot moderately to darkly pigmented.
Pallial roof and visceral coil almost uniformly black; pallial
edge black.
Ctenidium overlapping pericardium; filaments about 36, tall,
strongly pleated. Osphradium about 45% of ctenidium length.
Hypobranchial gland swollen on proximal genital ducts. Fecal
pellets oblique. Renal organ rarely swollen; renal opening
white. Salivary glands pigmented, overlapping cerebral gan-
glia. Stomach about as long as style sac. Ganglia pigmented.
Testis 2.0 whorls, overlapping posterior stomach chamber,
filling 50%-67% of visceral coil behind stomach. Prostate
gland with 33% of length in pallial roof. Vas deferens nearly
straight in pallial roof and in neck. Penis (Figure 9c) of
medium size, sickle-shaped, coiled, with strong folds along
proximal 50%-67% of length; base slightly narrowed; medial
section gently tapering; distal section narrowing slightly to
medium-long, pointed tip. Penial duct near center, medium
wide, strongly undulating in basal and medial sections. Penis
usually with very dark epithelial pigment on proximal 50% of
dorsal surface; black subepithelial pigment scattered along
length of penial duct.
Ovary 0.75 whorl, abutting posterior edge of stomach, filling
about 50% of visceral coil behind stomach. Distal female
genitalia shown in Figure 10c. Coiled oviduct narrow;
proximal portion with 1-2 small, horizontal to oblique twists
or coils; distal portion of primary coil sometimes containing
sperm. Coiled oviduct and bursal duct join just behind pallial
wall. Bursa copulatrix 53%-63% of albumen gland length,
pyriform, longer than wide, positioned largely posterior to
gland. Bursal duct short (15%-25% of bursa length), originat-
ing slightly lateral to tip of organ. Seminal receptacle about
25% of length of bursa copulatrix, positioned just anterior to
bursa copulatrix near posterior edge of albumen gland. Pallial
gonoduct with deep rectal furrow. About 33% of albumen
gland lying in front of posterior pallial wall. Capsule gland
about as long and as wide as albumen gland. Ventral channel
with anterior vestibule. Genital opening a very short, sub-
terminal slit.
DISTRIBUTION.?Lower Snake River, Columbia River, and
large tributaries (Figure 11).
REMARKS.?Haldeman described F. fuscus based on speci-
mens from Thomas Nuttall, who presumably collected the snail
when he journeyed along the lower Snake-Columbia River
system in 1834 (Pennell, 1936). The two lots listed by Baker
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
(1964:172) as possible type material (ANSP 27772, ANSP
27774) are whitened, worn shells devoid of periostracum,
which he admitted do not conform to the species as originally
described and illustrated; there is no convincing evidence from
associated labels that either lot represents type material (only
one of these lots is from Oregon; the other lot is from Utah
Lake). Despite the fact that Haldeman's types have not been
found, his original description and subsequent illustrations
(Haldeman, 1847) clearly identify his species as the large form
with angulate body whorl that is (or at least was historically)
widespread in the lower Snake-Columbia River system.
Baird (1863:67) gave the type locality for A. hindsi as "River
Kootanie, and stream at the foot of the Rocky Mountains." A
single lot (seven specimens) identified as this species in the
British Museum (BMNH 1863.2.4.17, labelled "syntypes,"
Kootanie River) probably is from the original type series that
Baird gave to that institution. As Baird did not figure any
specimens along with his description, we herein select the only
fully adult specimen (Figure 4D) as the lectotype. Baird's
description mentioned the carinate nature of the body whorl,
and both his types and shells from the nearby Wigwam River
locality (the second locality that he listed for this species)
closely conform to F. fuscus; hence, we treat A. hindsi as a
subjective junior synonym of the former, despite its apparently
disjunct occurrence in British Columbia.
Keep's (1887:63) minimal description of F. nuttalliana
columbiana as being "more rounded [than F. nuttalliana], with
a shorter spire" is relatively uninformative; however Pilsbry's
(1899a: 125) types and description of having "dark
color , . . . narrow but prominent shoulder of the last
whorl,... absence of callous on the inner lip" readily permits
identification as F. fuscus. Pilsbry (1899a: 125) gave the type
locality for his species as "Columbia River, Washington, near
Wallula and near mouth of Snake River, Snake River, near
Weiser, Idaho," but original labels associated with his type
series restrict the locality to the first mentioned site. Note that
F. columbiana Pilsbry, 1899, is a junior homonym of F.
nuttaliana columbiana Keep, 1887.
Fluminicola fuscus (or its synonyms) has been attributed by
many authors to the Great Basin of Utah, middle or upper
Snake River drainage, and other waters to the east (Tryon,
1870; Call, 1884; Pilsbry, 1899a; Orcutt, 1901a; Hannibal,
1912; Henderson and Daniels, 1917; Henderson, 1924, 1931,
1936a; Chamberlin and Jones, 1929; Jones, 1935, 1940;
Woolstenhulme, 1942; Berry, 1948; Baily and Baily, 1951;
Beetle, 1957; Rubin and Taylor, 1963; Branson, Sisk, and
McCoy, 1966; Taylor 1966a, 1985; Taylor and Bright, 1987;
Neitzel and Frest, 1989; Bowler and Frest, 1992; Frest and
Bowler, 1993; Neitzel and Frest, 1993). As mentioned above,
we believe that these populations comprise one or more
undescribed species. Although we allocated lower Salmon
River material to F. fuscus, populations found above the River
of No Return do not closely conform to this animal and may be
conspecific with undescribed snails living in the headwaters of
the Snake River. Anatomical data are from USNM 883182.
MATERIAL EXAMINED.?Columbia River, USNM 27901.
BRITISH COLUMBIA. Kootanie [sic; Kootenai] River,
BMNH 1863.2.4.17A (lectotype, A. hindsi Baird), BMNH
1863.2.4.17 (paralectotypes, A. hindsi Baird). Wigwam River,
west slope of Rocky Mountains, BMNH 1863.2.4.16.
IDAHO. Snake River, near Weiser City, ANSP 62341.
Salmon River, USNM 762527. Idaho County: Salmon River,
3-5 mi (4.8-8.0 km) east of Riggins, UF 47432 (mixed with
Fluminicola sp.). Salmon River, 10.5 mi (16.9 km) east of
Riggins, UF 47436 (mixed with Fluminicola sp.). Salmon
River, 18 mi (28.9 km) north of Riggins, UF 47427 (mixed
with Fluminicola sp.). Skookumchuck Creek, access across
from US 95, Salmon River, river mi 57.9-58.3 (km 93.2-
93.8), USNM 883520*. Salmon River, river mi 83.6-83.8 (km
134.5-134.8), north of Riggins, USNM 883516*. Spring Bar,
Salmon River, river mi 97.0 (km 156.1), east of Riggins,
USNM 883510*. Salmon River, River of No Return, river mi
107.6 (km 173.1), USNM 883519*. Valley County: Salmon
River, 25 mi (40.2 km) east of Riggins, UF 47438 (mixed with
Fluminicola sp.), UF 47439* (mixed with Fluminicola sp.).
Washington County: Snake River, Weiser, CAS 93885.
OREGON. USNM 47875. Columbia River, ANSP 122135.
Portland, USNM 473990. Willamette River, UF 88656 (mixed
with Fluminicola sp.), USNM 99303 (mixed with F. virens),
USNM 854693. Multnomah County: CAS 93905. Wallowa
County: Grande Ronde River, 0.5 mi (0.8 km) northeast of
Troy, UF 46002*, UF 46003 (mixed with Fluminicola sp.).
Grande Ronde River, river mi 52.7 (km 84.8), USNM 883495*.
Mouth of Imnaha River, Hells Canyon, USNM 883515*.
Wasco County: Columbia River, The Dalles, CAS 93904, CAS
93786 (mixed with Fluminicola sp.).
WASHINGTON. ANSP 27763, CAS 93900. Columbia River,
near mouth of Snake River, ANSP 62926. Snake River, ANSP
122133, ANSP 122136, CAS 93902. Spokane River, ANSP
122137. Asotin County: Grande Ronde River, 13.2 mi (21.2
km) northeast of Troy, UF 46007. Grande Ronde River, river
mi 0.5 (km 0.8), near junction with Snake River, Hells Canyon,
USNM 883499*. Grande Ronde River, river mi 30.2 (km 48.6),
USNM 883497*. Grande Ronde River, river mi 36.4 (km 58.6),
USNM 883509*. Grande Ronde River, river mi 41.1 (km 66.1),
USNM 883504*. Chelan County: Mouth of Wenatchee River,
USNM 130527. Okanogan County: Methow River, first bridge
north of Pateros, USNM 883182*. Methow River, WA 50,
USNM 883637*, USNM 883642*. Okanogan River, river mi
14.0-14.3 (km 22.5-23.0), USNM 883639*. Okanogan River,
Rattlesnake Point, river mi 15.0-15.3 (km 24.1-24.6), USNM
883640*. Spokane County: Spokane Falls, ANSP 73837, CAS
93884, CAS 93893, USNM 511023, USNM 511051. Spokane,
ANSP 75779, USNM 128665. Little Spokane River, north of
Spokane, USNM 380804. Walla Walla County: Columbia
River, near Wallula, ANSP 27767 (lectotype, F. columbiana
Pilsbry), ANSP 398349 (paralectotypes, F. columbiana
Pilsbry), CAS 58880, CAS 58881. Columbia River, near mouth
of Snake River, Wallula, CAS 93908. Snake River, near mouth,
USNM 30588.
NUMBER 583 13
Fluminicola minutissimus Piisbry, 1907
Fluminicola minutissima Piisbry, 1907:76 [lectotype: ANSP 94273; type
locality: Price Valley, Weiser Canyon, Washington County, Idaho].?
Piisbry, 1908:133, pi. XI: fig. 4.?Hannibal, 1912:188.?Walker,
1918:142.?Henderson, 1924:192.?Baker, 1964:174 [lectotype selec-
tion].?Burch and Tottenham, 1980:102, fig. 147.?Turgeon et al., 1988:60.
"Fluminicola" minutissima.?Taylor, 1975a: 124.
DIAGNOSIS.?Small with globose shell; animal unknown.
Fluminicola minutissimus is distinguished from other small
congeners by its minute size, globose shell, and blunt spire.
DESCRIPTION.?Shell (Figure 4E) globose, with blunt spire;
height about 1.5 mm; whorls 3.0. Protoconch (Figure 6D) of 1.5
whorls; diameter about 0.56 mm; microsculpture of numerous
weak spiral striae. Teleoconch whorls convex, sometimes with
narrow sutural shelf; whorls broadest below midpoint. Mi-
croscupture of medium growth lines. Periostracum green to
light brown. Shell transparent, clear white. Aperture large,
nearly circular, slightly angled above. Outer lip prosocline,
weakly sinuate, of medium thickness. Parietal lip complete, of
medium thickness, broadly adnate. Columellar lip of medium
thickness, columellar swelling moderate. Shell perforate;
umbilical region narrowly excavated, with adaxial ridge.
DISTRIBUTION.?Thus far known only from the type locality,
in the Snake River drainage (Figure 11).
REMARKS.?Pilsbry's type material, collected by Ashmun in
1904, comprises the only known specimens for this species. We
have visited the type locality area on several occasions, and we
were unable to locate this animal.
MATERIAL EXAMINED.?IDAHO. Washington County:
Price Valley, Weiser Canyon, ANSP 94273 (lectotype), ANSP
398350 (paralectotypes).
Fluminicola modoci Hannibal, 1912
Amnicola micrococcus.?Hannibal in Keep, 1911:314, pi. Ill: fig. 13.
Fluminicola modoci Hannibal, 1912:187, pi. 8: fig. 30 [in part; lectotype: CAS
60798; type locality: Fletchers Spring, south end, Goose Lake, California].?
Walker, 1918:142.?Taylor, 1966a:24, fig. 9 [as synonym of L. turbini-
formis].?Taylor and Smith, 1971:312, figs. 16,21 [lectotype designation; as
synonym of L. turbiniformis].?Burch and Tottenham, 1980:102.?Taylor,
1981:153 [as synonym of L. turbiniformis].?Turgeonet al., 1988:60.
Lithoglyphus modoci.?Taylor, 1975a: 125.
DIAGNOSIS.?Small with broadly conical shell; penis large,
sickle-shaped.
A pigmented renal opening is unique to F. modoci within the
genus. Flumincola modoci is further separable from the closely
related F. seminalis by narrow central cusps on central and
lateral teeth, more numerous cusps on marginal teeth, and
pleated ctenidial filaments.
DESCRIPTION.?Shell (Figures 4F, 5H-J) broadly conical,
usually with eroded spire; height 2.7-4.1 mm; whorls
3.5-3.75. Protoconch (Figure 6E) of 1.5 whorls, diameter
about 0.65 mm; microsculpture of numerous, strong, anasto-
mosing spiral striae. Teleoconch whorls convex; shoulders well
developed but narrow. Microsculpture of strong collabral
growth lines. Periostracum yellow green. Shell translucent,
clear white. Aperture large, broadly lunate, strongly angled
above. Outer lip prosocline, weakly sinuate, thin. Parietal lip
complete, thin, broadly adnate. Columellar lip thin, columellar
swelling medium wide, sometimes covering umbilical region.
Shell usually anomphalous, sometimes narrowly rimate;
umbilical region sometimes narrowly excavated, with adaxial
ridge.
Operculum (Figure 7D) of medium thickness; outer margin
without obvious rim. Attachment-scar margin strongly thick-
ened all around, although somewhat less so along outer edge.
Callus well developed.
Radula (Figure 8J-L) with about 73 rows of teeth; ribbon
length 1.1 mm, ribbon width 0.14 mm; central tooth width 42
|im. Central tooth with pronounced dorsal indentation; lateral
cusps 4-5; median cusp narrowly U-shaped, considerably
broader and slightly longer than laterals; basal cusps 3, broadly
triangular, originating from lateral angles, outermost some-
times weakly developed; basal tongue medium wide, basal
sockets moderately excavated; lateral angles weakly thickened.
Lateral tooth with flat or very slightly concave dorsal edge;
dorsal edge about 40% of tooth width; lateral shaft slightly
longer than height of tooth face; tooth face taller than wide;
central cusp narrowly U-shaped, lateral cusps 3-4 (inner side)
to 3-5 (outer side). Inner marginal teeth with 23-26 cusps;
outer marginal teeth with 26-29 cusps.
Snout black; distal lips grey black, pale near edge. Head pale
grey. Tentacles dark grey black, with pale to light grey patch
around eyes. Foot dark grey black. Pallial roof and visceral coil
black.
Ctenidium overlapping pericardium; filaments about 20,
short, pleated. Osphradium about 32% of ctenidium length.
Hypobranchial gland swollen on proximal genital duct Fecal
pellets longitudinal. Renal organ not swollen; renal opening
pigmented. Salivary glands overlapping cerebral ganglia.
Stomach longer than style sac. Ganglia pigmented.
Testis 1.0 whorl, overlapping posterior and portion of
anterior stomach chambers, filling about 67% of visceral coil
behind stomach chamber. Prostate gland with about 40% of
length in pallial roof, fat. Vas deferens straight in pallial roof,
with a few bends in neck. Penis (Figure 9D) large, medium
wide, sickle-shaped, curved to tightly coiled, with a few weak
folds along proximal inner curvature; penis tapering gently
along entire length to pointed distal tip. Penial duct near center,
narrow, straight or very weakly undulating. Penis pale or with
light epithelial pigment on distal half; subepithelial pigment
moderate along entire length of penis.
Ovary 0.5 whorl, positioned just behind posterior edge of
stomach, filling about 33% of visceral coil behind stomach.
Distal female genitalia shown in Figure 10D. Coiled oviduct
narrow, unpigmented; proximal portion with small vertical
coil; distal portion of primary coil with prominent bulge
containing sperm. Coiled oviduct and bursal duct join well
behind pallial wall. Bursa copulatrix about 55% of albumen
gland length, ovate, wider than long, partly overlapped by
albumen gland. Bursal duct very short (<16% of bursa length),
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
originating near middle of long axis of organ. Seminal
receptacle small, 25%-33% of bursa copulatrix length,
unpigmented, partly overlapping anterior bursa copulatrix near
posterior edge of albumen gland, partly imbedded in gland.
Pallial gonoduct with weak rectal furrow. About 33% of
albumen gland lying in front of posterior pallial wall. Capsule
gland shorter, narrower than albumen gland. Ventral channel
with or without very short anterior vestibule. Genital opening a
terminal pore.
DISTRIBUTION.?Thus far known only from the type locality
area, along the southwest corner of Goose Lake (California)
(Figure 11).
REMARKS.?As implied by Taylor and Smith (1971:312),
Hannibal (1912:187) mixed material of two species when he
described F. modoci ("elevated-conic, spire decidedly ele-
vated"), as his figured "type" is a narrow, high-spired form
{Pyrgulopsis?) that conflicts with the broader, typically
decollate Fluminicola shells comprising the available type
series (readily identifiable by original labels associated with
Hannibal material transferred from Stanford University to the
CAS; Taylor and Smith, 1971). Also note that in his
description, Hannibal refers to an earlier illustration of F.
modoci (in Keep, 1911, pi. Ill: fig. 13) that also conforms to its
identity as a Fluminicola. It is evident that Hannibal used the
wrong photograph as an illustration of F. modoci: note that he
used this same photograph earlier (in Keep, 1911, pi. Ill: fig.
11) to depict Pyrgulopsis longinqua. We interpret Hannibal's
error as a lapsus calami, and thus accept his description of F.
modoci as a Fluminicola (as opposed to treating F. modoci as
an inadvertently described Pyrgulopsis, thus requiring a new
name for the Fluminicola). The specimen identified as the
holotype off. modoci by Taylor and Smith (1971, figs. 16,21)
is not the shell Hannibal illustrated (although it appears to be
that which he earlier illustrated as Amnicola micrococcus (in
Keep, 1911)); hence, it cannot serve as the holotype (ICZN,
Article 73a[iv]). In addition, the original labels associated with
this specimen contain several different handwriting styles, and
it is not clear whether Hannibal himself indicated that this
specimen is the holotype. In our view, Taylor and Smith
selected a lectotype for F. modoci (instead of identifying the
holotype). We have not seen Hannibal's material from the other
two sites he originally listed, but one of us (Hershler) has
visited these localities, which occur in the Great Basin
southeast of Goose Lake basin, and found only Pyrgulopsis;
hence, we are restricting F. modoci to the type locality area.
This situation is complicated yet further in that several paratype
lots are mixed with a second, smaller (undescribed) Flumini-
cola species having a squatter shell.
The type locality probably refers to unnamed springs at
Fletcher Place along the southwest corner of Goose Lake,
township (T) 45N, range (R) 13E, section (sec.) 18. We could
not access this site owing to problems related to private
ownership of this land, and for our anatomical study we instead
used material (USNM 883468) from nearby springs (no more
than 10 km distant) that also are along the southwest margin of
Goose Lake.
MATERIAL EXAMINED.?CALIFORNIA. Modoc County:
Fletchers Spring, south end, Goose Lake, CAS 60798
(lectotype, paralectotypes), CAS 60799 (paratypes), CAS
66545 (paratypes). Unnamed spring, Three Springs Ranch,
Goose Lake basin, T46N, R 13E, sec. 16,21, USNM 883468*.
USNM 883469*, USNM 883867*, USNM 883868*.
Fluminicola nuttallianus (Lea, 1838)
Paludina nuttalliana Lea, 1838:101, pi. XXIII: fig. 89 [lectotype: USNM
121467, selected herein; type locality: Wahlamat (sic) (River), near its
junction with the Columbia River].?Frauenfeld, 1864:630.?Hannibal,
1912:187 [as synonym of F. virens].
Lepioxis nutlaliana [sic].?Haldeman, 1847:6, pi. 5: fig. 156 [in part].?Brot,
1862:25 [in part].
Lepioxis nuttalliana.?Binney, 1859:12; 1860:11.
Amnicola nuttalliana ? W. Cooper, 1860:374.?Frauenfeld, 1863b:1029.?
Grasset, 1884:92.
Anculotus nuttalii [sic].?Reeve. 1860-1861 :[11; species 46] [in part].
Fluminicola nuttallii [sic].?Carpenter. 1864:676 [in part].
Fluminicola nuttalliana?Carpenter, 1864:598.?Binney, 1865:89-91 [in
part].?Stimpson. 1865a:53; 1865b:54.?J.G. Cooper. 1867:30.?Tryon,
1870:65. pi. 17: fig. 17 [in part].?Paetel. 1883:71.?Rftmer. 1891:66.?
Pilsbry, 1899a:123 [in part].?Orcutt 1901a:35 [in part]; 19027:61 [in
part].?Dall, 1905:119 [in part].?Walker, 1918:32.?Henderson. 1929:168
[in part].?Morrison, 1940:125.?La Rocque, 1953:270 [in part].?Burch
and Tottenham, 1980:102, fig. 142.?Turgeon et al., 1988:61.
Paludina nutalliana [sic].?Anonymous, 1866:272.
Fluminicola nuttalii [sic].?Schmeltz, 1874:103.?Hemphill, 1881:8.
Fluminicola nuttaliana [sic].?Wenz, 1939:576.
Lithoglyphus nuttallianus.?Taylor, 1975a: 132.
DIAGNOSIS.?Large with broadly conical to globose shell;
animal unknown.
The shell of this apparently extinct species closely resembles
that of larger specimens of F. coloradensis, but it is thicker and
has a narrower columellar swelling and more consistently
reduced umbilicus.
DESCRIPTION.?Shell (Figure 13A) broadly conical to
globose, with very short spire; height 8.5-10.0 mm; whorls
4.25-4.5. Protoconch (Figure 6F) of 1.5 whorls; diameter
about 0.92 mm; microsculpture of numerous, strong, anasto-
mosing spiral striae. Teleoconch whorls highly convex, with
very broad, flattened sutural shelf; whorls broadest below
midpoint. Microsculpture of collabral growth lines. Perios-
tracum tan. Shell opaque, grey white. Aperture large, broadly
lunate, rounded below, angled above. Outer lip strongly
prosocline, thin. Parietal lip complete, thin, adnate. Columellar
lip of medium thickness, columellar swelling narrow to
moderate. Shell anomphalous or with small, chink-like
umbilicus.
DISTRIBUTION.?Known with certainty only from the type
locality area, near the mouth of the Willamette River (Figure
14). We have been unable to locate the Hartford, Oregon, site,
although it may refer to Hanford, Washington, which is well
upstream from the mouth of the Willamette, on the Columbia
River.
REMARKS.?This species was incorrectly referred to as Lea
NUMBER 583 15
(1839) by Kabat and Hershler (1993). Identification of the three
species of Fluminicola that Lea named from the mouth of the
Willamette River historically has posed difficulties. This is due
to the brief and sometimes enigmatic original descriptions,
uncertain status of type material, virtual destruction of
Fluminicola habitat in the type locality and vicinity as a result
of the growth and development of the Portland metropolitan
area in the past 150 years, and historic complexity of the
Fluminicola fauna in this river, which includes not only Lea's
taxa andF.fiiscus, but also two undescribed forms (a neritiform
species rare in historical collections and probably extinct, and
a large, convex-whorled form still common in the river, which
has been confused with F. nuttallianus). Lea's description and
illustration defined F. nuttallianus as a large, nearly globose
form (the shell resembling that of F. coloradensis), which is
rarely represented in historic collections from the region and is
probably now extinct. A single lot of three shells (now labeled
"Wahlamette [sic], Oregon") from the Lea collection in the
National Museum of Natural History conforms to Lea's
description and probably represents part of his type series
(original labels are absent). All three specimens in this lot are
somewhat smaller than the size he indicated for the species. Of
these, one is juvenile and another is extremely worn; hence, we
have selected the third specimen, an adult, as the lectotype
(Figure 1 3 A).
We recognize F. nuttallianus as distinct from a larger, more
elongate, undescribed form from the Willamette River and
distinct from one or more large, subglobose, round-whorled
forms found elsewhere in western Washington and Oregon.
Both of these forms have been frequently confused with F.
nuttallianus. We similarly view the disjunct population in the
Clearwater River (Idaho), referred to as F. nuttallianus by
Taylor (1966a), as both undescribed and distinct from our
concept of F. nuttallianus.
Valid records of F. nuttallianus are from a limited portion of
the lower Willamette River, namely from the vicinity of
Oregon City (at approximately river mi 20 (km 32.2)) to the
historic mouth. This reach extends from Willamette Falls, at
Oregon City, along the present Multnomah Channel to its
junction with the Columbia River near St. Helens, Oregon. The
Multnomah Channel was abandoned as the primary Willamette
channel when the Willamette River shifted to approximately its
present position after a major flood in 1890. On several
occasions between 1991 and 1994 we searched for Fluminicola
in the vicinity of Willamette Falls and at both the Multnomah
channel and the present course of the Willamette River
downflow from Oregon City. This area is now highly
industrialized. A lumber mill, locks, and a power plant have
long been present at Willamette Falls. Sites surveyed above the
falls have yielded F. virens and an undescribed species in this
genus but have not yielded F. nuttallianus. All of the adjoining
lower Columbia River is impounded, generally has soft
substrates, and has few surviving colonies of Fluminicola, none
of which resembes F. nuttallianus. We consider it likely that
this species is now extinct.
MATERIAL EXAMINED.?West Coast, ANSP 365331.
OREGON. UF 88670. Hartford (?), USNM 47874 (mixed
with Fluminicola sp.). Willamette (River), ANSP 27771.
Wahlamette [sic], Oregon, USNM 121467 (lectotype), USNM
860648 (paralectotypes).
Fluminicola seminalis (Hinds, 1842)
Paludina seminalis Hinds, 1842:83-84 [type not located; type locality: Rio
Sacramento, California].?Hinds, 1845:59, pi. XVI: fig. 22.?Haldeman,
1847:6 [as synonym of L. nuttaliana].?Carpenter, 1857:206, 211.?Brot,
1862:25 [as synonym of L. nuttaliana].?Carpenter, 1864:676 [as synonym
of F. nuttallii].?Frauenfeld, 1864:645 [as synonym of L. nuttalliana].?
Binney, 1865:89 [as synonym of F. nuttalliana].
Melania seminalis.?Lea, 1856:81.
Bithinia seminalis.?Carpenter, 1857:326.?Binney, 1859:13.
Amnicola seminalis.?W. Cooper, 1860:374.?Grasset, 1884:91.
Lithoglyphus cumingii Frauenfeld, 1863a: 195 [lectotype: BMNH 1933047,
selected herein; type locality: California]; 1865:530, pi. 11: unnumbered
fig.?Walker, 1918:142 [as synonym of F. seminalis].?Taylor, 1975a:69.
Fluminicola seminalis.?Carpenter, 1864:598.?Stimpson, 1865a:53;
1865b:54.?Pilsbry, 1899a:123 [in part]; 1899b:66.?Hannibal, 1912:187
[in part].?Walker, 1918:142 [in part].?Henderson, 1929:169 [in part].?
Morrison, 1940:125.?Burch and Tottenham, 1980:102 [in part].?Turgeon
etal., 1988:61.
Melania (Paludina) seminalis.?Carpenter, 1864:634.
Paludina cumingii.?Frauenfeld, 1864:595.
Lithoglyphus cumingi [sic].?Pilsbry, 1899a:123 [as synonym of F. semi-
nalis].?Taylor, 1981:153.
Fluminicola seminis [sic].?Wenz, 1939:576.
Fluminicola seminaris [sic].?Wenz, 1939:576.
Lithoglyphus seminalis.?Taylor, 1966*48; 1975a:172; 1981:153.
DIAGNOSIS.?Medium to large, with globose to broadly
conical shell; penis large, sickle-shaped.
DESCRIPTION.?Shell (Figures 13B,C, 15A-F) globose to
broadly conical, spire often eroded; height 5.8-8.0 mm; whorls
4.0-4.5. Protoconch (Figure 6G) of 1.5 whorls; diameter about
0.8 mm; microsculpture of numerous strong spiral striae.
Teleoconch whorls convex, often highly so, weakly to strongly
shouldered. Microsculpture of collabral growth lines, some-
times with fine spiral striae. Periostracum tan, brown, or light
green. Shell opaque, clear white. Aperture large, lunate, slightly
angled above. Outer lip thin. Parietal lip complete, thin, usually
adnate, rarely slightly detached. Columellar lip thin, columellar
swelling medium wide to broad. Shell usually anomphalous,
sometimes with slit-like umbilicus; umbilical region some-
times narrowly excavated, with adaxial ridge.
Operculum (Figure 16A) of medium thickness; outer margin
with weak rim. Outer edge of attachment-scar thickened,
sometimes strongly so; inner edge slightly thickened. Callus
weak to moderately thickened, small.
Radula (Figure 17A-C) with about 71 rows of teeth; ribbon
length 2.2 mm, ribbon width 0.25 mm; central tooth width 70
Hm. Central tooth moderately indented dorsally; lateral cusps
4-5; median cusp broadly U-shaped, considerably broader and
slightly longer than laterals; basal cusps 2-3, narrow,
innermost arising from tooth face; basal tongue medium wide,
basal sockets moderately excavated; lateral angles broad.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
thickened. Lateral tooth with slightly concave dorsal edge;
dorsal edge about 50% of tooth width; lateral shaft slightly
longer than height of tooth face; face tooth face about as tall as
wide; central cusp broadly U-shaped, lateral cusps 3-4. Inner
marginal teeth with 15-17 cusps; outer marginal teeth with
18-21 cusps.
Snout and foot black; head pale grey; tentacles black except
for pale circle surrounding eyes. Pallial roof and visceral coil
black.
Ctenidium overlapping pericardium; filaments about 35, tall,
without pleats. Osphradium about 40% of ctenidium length.
Hypobranchial gland swollen over proximal genital ducts.
Fecal pellets longitudinal. Renal organ swollen; renal opening
opaque. Salivary glands overlapping cerebral ganglia. Stomach
slightly longer than style sac. Ganglia pigmented.
Testis 1.5 whorls, overlapping posterior stomach chamber,
filling 67% of visceral coil behind stomach. Prostate gland with
33% of length in pallial roof, elongate. Vas deferens slightly
undulating in pallial roof, straight in neck. Penis (Figure 9E)
large, curved, sickle-shaped, with weak folds along proximal
half; penis gently tapering along entire length to pointed tip.
Penial duct near outer edge, narrow, coiled along proximal 67%
of length. Penis with grey black epithelial pigment on proximal
50%; black subepithelial pigment concentrated distally.
Ovary 0.5 whorl, abutting posterior edge of stomach, filling
about 50% of visceral coil behind stomach. Distal female
genitalia shown in Figure 18A. Coiled oviduct narrow,
positioned about midway between pallial wall and anterior
edge of bursa copulatrix, unpigmented; proximal portion with
2 small oblique-horizontal loops; distal section of primary coil
overlapped by albumen gland, sometimes containing sperm.
Coiled oviduct and bursal duct join well behind pallial wall.
Bursa copulatrix about 50% of albumen gland length, pyriform,
wider than long, positioned about 33% posterior to gland.
Bursal duct extremely short, originating near middle of long
axis of organ. Seminal receptacle about 25% of bursa
copulatrix length, unpigmented, abutting anterior edge or
slightly overlapping bursa copulatrix near posterior edge of
albumen gland, partly imbedded in gland. Pallial gonoduct with
deep rectal furrow. About 35% of albumen gland lying in front
of posterior pallial wall. Capsule gland about as long as and
slightly narrower than albumen gland. Ventral gland without
vestibule. Genital opening a pore mounted on small terminal
papilla.
DISTRIBUTION.?Sacramento River basin from near its
mouth (historically), upstream into the Pit River, including
large, spring-fed tributaries (Figure 14; modified from Taylor,
1981). The species is apparently now extinct in the main-stem
Sacramento River.
REMARKS.?Hinds described P. seminalis based on speci-
mens he collected during his voyage on the H.M.S. Sulphur by
means of the consort ship Starling, which coursed up the
Sacramento River in fall, 1837. The precise type locality is
unknown, but it surely was from the lower portion of the
Sacramento River as the ship did not penetrate more than 160
km upstream from the mouth (from original voyage accounts
summarized in Pierce and Winslow, 1979). (Note that Taylor
(1981) gave the type locality as Sacramento River below mouth
of American River.) Hinds' type series has not been located,
although specimens in the British Museum identified as this
species (Sacramento River, uncataloged, H. Cuming coll.) may
comprise part of such, as much of his collection was donated to
that institution, either by himself or by private collectors,
including Cuming (Keen, 1966). Two lots of single specimens
identified as syntypes of L. cumingii (labeled "California")
were located in the BMNH and NMW: the former specimen
more closely resembles Frauenfeld's (1865) illustration of the
species and thus we select this as the lectotype (Figure 13c).
Frauenfeld's description and type material (i.e., medium-sized,
globose or broadly conical shell with thin parietal lip) allow
recognition of L. cumingii as a junior subjective synonym ofF.
seminalis as previously suggested in the literature.
Material from the San Joaquin River, Napa County, and
Klamath basin has been assigned to this species either in
museum collections or in the literature (Pilsbry, 1899a; Hanna,
1924; Henderson, 1929; Clench, 1940), but based on differ-
ences in shell features, we prefer to exclude these from F.
seminalis for the time being. Anatomical data are from USNM
883465.
MATERIAL EXAMINED.?CALIFORNIA. BMNH 1933047
(lectotype, L. cumingii Frauenfeld), NMW uncat. Sacramento
River, ANSP 27776, ANSP 27778, BMNH 1994109, BMNH
1994110, CAS 93865, UF 227630, USNM 11860, USNM
121465. Shasta County: ANSP 341358. Battle Creek, Battle
Creek County Park (on Tehama County line), USNM 883465*.
McCloud River, USNM 883188*. Lava Creek, Lava Springs
Ranch, USNM 883180*. Pit River, above Pit #1 foot bridge,
USNM 883739*. Big Lake, Ahjumawi Lava Springs State Park,
USNM 883742*. Rising River, bridge crossing on Redding-
Alturas HWY, CAS 93863. Burney Falls (Bumey Creek), CAS
93777, CAS 93862. Fall River, near Glenbum, CAS 93860.
Tehama County: Sacramento River, near Red Bluffs, ANSP
122139.
Fluminicola turbiniformis (Tryon, 1865)
Amnicola turbiniformis Tryon, 1865:219, pi. 22: fig. 5 [in part; lectotype:
ANSP 27779; type locality: west side of Steens Mountains, Harney County,
Oregon].?J.G. Cooper, 1867:30?Tryon, 1870:56, pi. 17: fig. 6 [in
part].?Grasset, 1884:91.?Pilsbry, 1899a: 123 [as synonym of F.
seminalis].?Orcutt, 1901b:48 [as synonym of Amnicola protea
Gould, 1855).?Hannibal, 1912:187 [as synonym of F. seminalis].?Walker,
1918:142 [as synonym of F. seminalis].?Henderson, 1929:169 [as synonym
of F. seminalis].
Amnicola turtoniformis [sic].?Paetel, 1888:444.
Fluminicola turbiniformis.?Baker, 1964:177 [lectotype selection].?Burch
and Tottenham, 1980:102, fig. 152 [in part].?Turgeon et al., 1988:61.
Lithoglyphus turbiniformis.?Taylor, 1966a:24, fig. 9 [in part].?Taylor and
Smith, 1971:312 [in part]; Taylor, 1975a:197; 1981:153 [in part]; 1985:309
[in part].
DIAGNOSIS.?Small with ovate- to narrow-conic shell; penis
small, sickle-shaped.
The extremely small size of the penis (relative to head) in
this species is unique among Fluminicola species.
NUMBER 583 17
DESCRIPTION.?Shell (Figures 5K,L, 13D) ovate- to narrow-
conic; height 3.1-4.3 mm; whorls 3.5-4.0. Protoconch (Figure
6H) of 1.5 whorls, diameter about 0.7 mm; microsculpture of
weak, anastomosing spiral striae. Teleoconch whorls highly
convex; shoulders weak or absent. Microsculpture of collabral
growth lines. Periostracum light green or brown. Shell
transparent, clear white. Aperture of medium size, ovate,
slightly angled above. Outer lip weakly sinuate, thin. Parietal
lip complete, thick, usually slightly detached. Columellar lip
thick, columellar swelling broad. Shell narrowly umbilicate;
umbilical region often narrowly excavated with adaxial ridge.
Operculum (Figure 16B) thin; outer margin without obvious
rim. Attachment-scar margin narrowly thickened along inner
edge to nucleus; outer edge thickening weak or absent. Callus
strongly developed.
Radula (Figure 17D-F) with about 83 rows of teeth; ribbon
length 1.25 mm, ribbon width 17 \im; central tooth width 46
^m. Central tooth with moderate dorsal indentation; lateral
cusps 4-5; median cusp narrowly U-shaped, slightly broader,
considerably longer than laterals; basal cusps 1, elongate,
originating from junction between tooth face and lateral angle;
basal tongue medium wide, basal sockets moderately exca-
vated; lateral angles thickened. Lateral tooth with straight to
slightly concave dorsal edge; dorsal edge about 40% of tooth
width; lateral shaft considerably longer than height of tooth
face; tooth face slightly taller than wide; central cusp U-shaped,
lateral cusps 2 (inner side) to 3 or 4 (outer side). Inner marginal
teeth with 26 cusps; outer marginal teeth with 29 cusps.
Snout light medium grey; distal lips pale. Head pale to
lightly pigmented. Tentacles pale to lightly pigmented, with
small, dark internal pigment patch distal to eyes. Foot lightly to
moderately pigmented. Pallial roof and visceral coil medium
brown to black, although not uniformly pigmented.
Ctenidium not overlapping pericardium; filaments 17, short,
without pleats. Osphradium about 30% of ctenidium length.
Hypobranchial gland thin. Fecal pellets longitudinal. Renal
organ slightly swollen; renal opening opaque. Salivary glands
overlapping cerebral ganglia. Stomach slightly longer than
style sac. Ganglia pigmented.
Testis 1.0 whorls, slightly overlapping posterior stomach
chamber, filling slightly less than 50% of visceral coil behind
stomach. Prostate gland with about 33% of length in pallial
roof, fat. Vas deferens nearly straight in pallial roof and in neck.
Penis (Figure 9F) small, flattened, narrow, tightly coiled,
sickle-shaped, weakly folded along portion of proximal left
side; base slightly swollen; medial section gently tapering;
distal section with subterminal narrowing followed by elon-
gate, pointed tip. Penial duct slightly to right of center, narrow,
gently undulating in proximal 67% of length. Penis without
epithelial pigment; dark internal pigment flanking penial duct.
Ovary 0.25 whorl, positioned slightly behind stomach,
filling about 12% of visceral coil behind stomach. Distal
female genitalia shown in Figure 18B. Coiled oviduct broad,
unpigmented; proximal portion with small twist or oblique
loop; distal portion of primary coil sometimes containing
sperm. Coiled oviduct and bursal duct join well behind pallial
wall. Bursa copulatrix about 50%-70% of albumen gland
length, ovate, wider than long, positioned largely posterior to
albumen gland. Bursal duct very short, originating near tip of
organ. Seminal receptacle 25%-33% of bursa copulatrix
length, unpigmented, slightly overlapping bursa copulatrix
near posterior edge of albumen gland, partly overlapped by
gland. Rectal furrow in pallial gonoduct weak or absent. About
38%-5O% of albumen gland lying in front of posterior pallial
wall. Capsule gland slightly longer but narrower than albumen
gland. Ventral channel without anterior vestibule. Genital
opening a terminal pore.
DISTRIBUTION.?Thus far known only from the type locality
area, in the northwestern Great Basin of Oregon (Figure 14).
REMARKS.?Tryon (1865:219) gave the type locality as
"Crane Lake Valley, and Surprise Valley, N.E. California; and
W. side of Stein's [sic; Steens] Mountains, S. Oregon."
Original labels associated with the lectotype permit a more
precise determination of the type locality, which Taylor (1985)
identified as Catlow Valley. Tryon's material from the other
two sites does not closely resemble F. turbiniformis in shell
features, and we have decided to restrict the species to the type
locality area. Anatomical data are from USNM 883470 (near
topotypes).
MATERIAL EXAMINED.?OREGON. Harney County: West
side of Steens Mountains, ANSP 27779 (lectotype), ANSP
398352 (paralectotypes). Roaring Springs, Catlow Valley,
USNM 883470*.
Fluminicola virens (Lea, 1838)
Paludina virens Lea, 1838:91, pi. XXIII: fig. 93 [lectotype: USNM 121431.
selected herein; type locality: Wahlarnat (sic) (River), near its junction with
the Columbia River].?Frauenfeld, 1864:659.
Paludina nuclea Lea, 1838:91, pi. XXIII: fig. 103 [type not located; type
locality: Wahlamat (sic) (River), near its junction with the Columbia
River.?Haldeman, 1847:5 [as synonym of L. virens].?Carpenter,
1857:207, 297.?Brot, 1862:26 [as synonym of L. virens].?Frauenfeld,
1864:630 [as synonym of P. virens].?Binney, 1865:91 [as synonym of F.
virens].?Pilsbry, 1899a: 123 [as synonym off. virens].?Dall. 1905:119 [as
synonym of F. virens].?Hannibal, 1912:187 [as synonym of F. virens].?
Henderson, 1929:169 [as synonym of F. virens].
Leptoxis virens.?Haldeman, 1847:5, pi. 5: figs. 147-150.?Binney, 1859:12;
1860:11.?Brot, 1862:26.
Bithinia nuclea.?Carpenter, 1857:326.?Binney, 1859:13.
Vivipara nuclea.?Binney, 1859:12 [as synonym of L. virens].
Vivipara virens.?Binney, 1859:13.
Anculosa nuclea.?Tryon, 1864b: 104.
Anculosa virens.?Tryon, 1864b: 104.
Fluminicola nuclea.?Stimpson, 1865a:53; 1865b:54.
Fluminicola virens.?Binney, 1865:91-92.?Stimpson, 1865a:53;
1865b:54.?J.G. Cooper, 1867:30.?Tryon, 1870:65, pi. 17: fig. 18 [in
part].?Pilsbry, 1899a:122-123 [in part].?Orcutt, 1901a:35-36 [in part];
19027:61-62 [in part].?Dall, 1905:119 [in part].?Hannibal, 1912:187 [in
part].?Henderson, 1929:169 [in part]; 1936b:277 [in part].?La Rocque,
1953:271 [in part].?Burch and Tottenham, 1980:102-103, fig. 145 [in
part].?Turgeonet al., 1988:61.
Lithoglyphus virens.?Taylor, 1966b:160; 1975a:205.?Clarke, 1981:70 [in
part].
Lithoglyphus nucleus.?Taylor, 1975a: 131.
Fluminicola nuttalliana.?Thompson, 1984:140 [in part].
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
DIAGNOSIS.?Large with ovate- to narrow-conic shell; penis
large, broadly triangular.
This species is phylogentically distinct from the clade
comprising all other species allocated to Fluminicola, and it is
set apart by various unique features (within the genus),
including unpigmented ganglia, short pedal commissure, vas
deferens coiled on neck, broadly triangular penis, penis deeply
folded along almost entire length, terminal papilla on penis,
simple, posterior oblique coiled oviduct, absence of secondary
storage of sperm in coiled oviduct, subglobular bursa copula-
trix, extremely long bursal duct, and large seminal receptacle.
DESCRIPTION.?Shell (Figures 13E,F, 15G-N) ovate- to
narrow-conic, usually with eroded spire; height 5.8-12.0 mm;
whorls 4.5-5.0. Protoconch (Figure 6i) of 1.5 whorls, diameter
about 0.85 mm; microsculpture of numerous strong spiral
striae. Teleoconch whorls flat to slightly convex; shoulders
very narrow or absent. Microsculpture of fine collabral growth
lines. Periostracum light tan, yellow, or green. Shell opaque,
grey white. Aperture of medium size, ovate, strongly angled
above. Outer lip weakly sinuate, usually thick. Parietal lip
complete, usually thick, broadly adnate, sometimes slightly
expanded over body whorl. Columellar lip of medium
thickness, columellar swelling narrow to medium wide. Shell
usually anomphalous; umbilical region sometimes narrowly
excavated, with adaxial ridge.
Operculum (Figure 16c) of medium thickness; outer margin
with very weak rim. Attachment-scar margin slightly thick-
ened, stronger along inner edge. Callus weak but with
pronounced narrow thickening extending to outer side of
nucleus.
Radula (Figure 17G-I) with about 78 rows of teeth; ribbon
length 1.84 mm, ribbon width 0.2 mm; central tooth width 59
|im. Central tooth with pronounced dorsal indentation; lateral
cusps 4-5; median cusp narrowly U-shaped, slightly broader
and longer than laterals; basal cusps 2-3, elongate; basal
tongue medium wide, basal sockets strongly excavated; lateral
angles slightly thickened. Lateral tooth with slightly concave
dorsal edge; dorsal edge about 50% of tooth width; lateral shaft
considerably longer than height of tooth face; tooth face about
as tall as wide; central cusp narrowly U-shaped, lateral cusps 3
(inner side) or 4 (outer side). Inner marginal teeth with 13-14
cusps; outer marginal teeth with 15-16 cusps.
Snout grey black; distal lips pale. Tentacles with black,
central, longitudinal band extending from eyes nearly to distal
tip, otherwise pale except for internal grey pigment at base.
Head grey. Foot pale to light grey. Pallial roof grey black.
Visceral coil nearly colorless to medium grey.
Ctenidium overlapping pericardium; filaments about 39, tall,
pleated. Osphradium almost 50% of ctenidium length. Hy-
pobranchial gland swollen on proximal genital duct. Fecal
pellets longitudinal. Renal organ swollen; renal opening white.
Salivary glands overlapping cerebral ganglia. Stomach slightly
longer than style sac. Pedal commissure less than 25% of
ganglia width. Ganglia unpigmented.
Testis 1.5 whorls, overlapping posterior stomach chamber,
filling 50%-67% of visceral coil behind stomach chamber.
Prostate gland with 33% of length in pallial roof, fat. Vas
deferens nearly straight in pallial roof, coiled in a series of tight
loops in neck. Penis (Figure 9G) large, broad, flattened, slightly
curved (but uncoiled), with strong folds along entire length
except distal tip (Figure lG); base slightly narrowed; medial
section weakly tapering; distal section strongly tapering, with
rounded tip bearing eversible papilla (Figure lH). Penial duct
central, narrow, tightly coiled in basal and medial sections.
Penis without epithelial pigment; two broad, dark bands of
black subepithelial pigment flanking penial duct.
Ovary 0.5 whorl, abutting posterior edge of stomach, filling
about 50% of visceral coil behind stomach. Distal female
genitalia shown in Figure 18c. Coiled oviduct narrow, posterior
oblique, unpigmented; proximal portion rarely with small kink.
Coiled oviduct and bursal duct join just behind pallial wall.
Bursa copulatrix about 33% of albumen gland length,
subglobular, about as long as wide, almost entirely overlapped
by gland. Bursal duct as long as or slightly longer than bursa,
originating from middle of long axis of organ. Seminal
receptacle 40%-50% of bursa copulatrix length, positioned
anterior to bursa copulatrix near middle of albumen gland,
entirely overlapped by gland. Pallial gonoduct without rectal
furrow. About 25% of albumen gland lying in front of posterior
pallial wall. Capsule gland shorter, narrower than albumen
gland. Ventral channel without anterior vestibule. Genital
opening a subterminal pore.
DISTRIBUTION.?Willamette River (and large tributary
streams), from Corvallis to its mouth, and the lower Columbia
River below Portland (Figure 14).
REMARKS.?Like F. nuttallianus, both P. virens and P.
nuclea were described from material provided by Nuttall. Two
lots in the National Museum of Natural History collection
identified as P. virens (now labeled "Wahlamette [sic],
Oregon") are Nuttall material from the Lea collection and
probably represent the type series (original labels are not
associated with these lots). One of these lots includes shells
closely resembling Lea's original figure of P. virens, and we
have selected from these a lectotype (Figure 13E). Identifica-
tion of Lea's P. virens is made difficult by the fact that his
description was based on decollate shells; nevertheless, his
illustration permits recognition of this snail as the relatively
tall, slightly convex-whorled species commonly found in the
lower Willamette River.
Despite the fact that Lea compared P. nuclea to (Campe-
loma) decisa, it is clear from his figure that this is the same
species as F. virens (as concluded by many earlier workers),
differing only in its less eroded spire and in having a remnant
of black periostracum bordering the aperture. We have not been
able to conclusively identify type material for P. nuclea,
although possible syntypes conforming to Lea's description
may be a single lot from the National Museum of Natural
History (labeled "Wahlamette [sic], Oregon," I. Lea collection,
NUMBER 583 19
ex Nuttall, USNM 121432; specimens mixed with more typical
F. virens) and a single specimen lot from the ANSP
(Willamette River, ex I. Lea, ANSP 27765).
This species has been incorrectly ascribed to other drainages
of western North America (e.g., Clarke, 1981; Beetle, 1989).
Anatomical data are from USNM 883673 (near topotypes).
MATERIAL EXAMINED.?OREGON. Wahlamette [sic],
USNM 121432, USNM 860649 (paralectotypes, P. virens
Lea). Willamette River, ANSP 27755, ANSP 27756, ANSP
27765, USNM 28115, USNM 99303 (mixed with F.fuscus),
USNM 121431 (lectotype, P. virens Lea). Portland, ANSP
27761 (mixed with Fluminicola sp.), ANSP 113137, USNM
473989, USNM uncat. (Lea 1379). Corvallis, USNM 531629.
Clackamas River, CAS 93810. Clackamas County: Canal or
flume from Tualatin River to Oswego Lake, ANSP 111291
(mixed with Fluminicola sp.). Willamette River, Oregon City,
ANSP 73605, USNM 511059 (mixed with Fluminicola sp.).
Oswego, USNM 511057 (mixed with Fluminicola sp.).
Willamette River, Canby Ferry (north side), USNM 883673*.
USNM 883779*. Willamette River, Bemert Landing (on
Clackamas-Linn County line) (mixed with Fluminicola sp.),
USNM 883670*. Willamette River (south side), river mi 34.9
(km 56.2), Molalla State Park, USNM 883461. Willamette
River (south side), river mi 40.9 (km 65.8), French Prairie State
Park, USNM 883183*. USNM 883692*. Linn County: Wil-
lamette River, Albany, USNM 883676 (mixed with Flumini-
cola sp.). Multnomah County: Near Portland, CAS 93814.
WASHINGTON. Cowlitz County: Columbia River, Aber-
nathy Point, just east of mouth of Abernathy Creek, USNM
883463*, USNM 883693*.
Appendix
Western American Fossil Species Described as or Allocated
to Fluminicola or Lithoglyphus
(in alphabetical order by species name, with emendations of type localites and current generic assignments)
Paludina abavia Mayer, 1869:40, pi. X: fig. 13. Miocene, English Bay, Alaska;
Miocene-Pliocene, Kenai Formation (Taylor, 1975a). Transferred to Lithoglyphus by
Taylor (1975a).
Lithasia antiqua Gabb, 1866:13, pi. 2: fig. 22. Fresh-water Tertiary deposit on Snake
River, Idaho Territory, on the road from Fort Bois6 to the Owyhee mining country; Late
Pliocene-Early Pleistocene, Glenns Ferry Formation, Canyon or Owyhee County,
Idaho, near Walters Ferry, T IS, R 2W, sec. 17 (Taylor, 1966a). Allocated to
Lithoglyphus by Taylor (1966a; as synonym of Natica(l) occidentalis Hall, 1845).
Lithoglyphus campbelli Dall, 1924:115, pi. XXVI: fig. 1. Mouth of King Hill Creek near
Glenn's Ferry, Snake River Canyon, Idaho, United States Geological Survey (USGS)
Station 3486; Late Pliocene-Early Pleistocene, Glenns Ferry Formation, USGS
Cenozoic locality 21510: Elmore County, Idaho, T 5S, R 10E, NW'/4 sec. 14 (Taylor,
1966a). Transferred to Pliopholyx (Pliopholygidae) by Taylor (1966a).
Fluminicola yatesiana inflata Yen, 1950:186, pi. 1: figs. 5,5a. Probably Pliocene, Truckee
Formation, western Nevada, 2 mi (3.2 km) north of the triangle marking the position of
the Desert Queen Mine, which lies in the northeastern corner of the Hot Springs
Mountains; Pliocene, Truckee Formation (Taylor, 1975a). Transferred to "Amnicola"
by Taylor (1975a).
Fluminicola junturae Taylor, 1963:38, figs. 8-10. Miocene, USGS locality 21173, Black
Butte local fauna, Malheur County, Oregon; Pliocene, Juntura Formation (Taylor,
1975a). Transferred to Lithoglyphus by Taylor (1975a).
Fluminicola kettlemanensis Pilsbry, 1934:15 (Pilsbry, 1935, pi. 19: figs. 1-3). San
Joaquin Formation outcrop; Kettleman Hills at Shell Oil Co. locality 2502, San Joaquin
Formation immediately below thcPecten coalingaensis zone (Pilsbry, 1935); Pliocene,
upper San Joaquin Formation, Kings County, California, T 22S, R 18E, near center sec.
26 (Taylor, 1966a). Transferred to Lithoglyphus by Taylor (1966a).
Fluminicola malheurensis Henderson and Rodeck, 1934:266-267, pi. 37: figs. 8a, 8b.
Pliocene, about 16 mi (25.7 km) southwest of Vale, Malheur County, Oregon; Pliocene,
Grassy Mountain Formation (Taylor, 1975a). Transferred to Lithoglyphus by Taylor
(1975a).
Natica{7) occidentalis Hall, 1845:308, pi. 3: figs. 8, 8a. Longitude 115?, latitude 43?, in a
gray siliceous limestone; Late Pliocene-Early Pleistocene, Glenns Ferry Formation,
Canyon or Owyhee County, Idaho, near Walters Ferry, T IS, R 2W, sec. 17 (Taylor,
1966a). Transferred to Lithoglyphus by Taylor (1966a).
Fluminicola percarinata Pilsbry, 1934:16 (Pilsbry, 1935, pi. 19: figs. 8,9). Lost Hills Oil
Field (California), 445-520 ft (136-158 m); Lost Hills Oil Field in Universal
Consolidated Well No. 44, at 445-520 ft (136-158 m) (Pilsbry, 1935); Pliocene, upper
part of San Joaquin Formation, Kern County, California, T 26S, R 2IE, sec. 32 (Taylor,
1966a). Transferred to Savaginius (=Pyrgulopsis; fide Hershler, 1994) by Taylor
(1966a).
Fluminicola perditicollis Pilsbry, 1934:16 (Pilsbry, 1935, pi. 19: fig. 4). Lost Hills Oil
Field (California), 455-650 ft (139-198 m); Lost Hills Oil Field in Universal
Consolidated Well No. 44, sec. 32-26-21, at 455-650 ft (139-198 m) (Pilsbry, 1935);
Pliocene, upper part of San Joaquin Formation, Kern County, California, T 26S, R 2 IE,
sec. 32 (Taylor, 1966a). Transferred to Savaginius by Taylor (1966a).
20
NUMBER 583 21
Fluminicola pilula Pilsbry, 1934:15 (Pilsbry, 1935, pi. 19: figs. 11-13). Lost Hills Oil
Field (California); San Joaquin Formation: Lost Hills Oil Field, Universal Consolidated
Well No. 44, sec. 32-26-21, at 435-445 ft (133-136 m) (Pilsbry, 1935); Pliocene,
upper part of San Joaquin Formation, Kern County, California, T 26S, R 2IE, sec. 32
(Taylor, 1966a). Transferred to Savaginius by Taylor (1966a).
Fluminicola protea Yen, 1948:41, pi. 10: fig. 12. Paleocene, Montana: Locality 17, 300 ft
(91 m) east from the southeast corner sec. 32, T 8S, R 46E, Powder River County, 100
ft (30 m) above base of Anderson clinker. Transferred to Reesidella (Bithyniidae) by
Taylor (1975b).
Fluminicola sanmateoensis Glen, 1960:1208-1209, fig. le-g. CAS locality number
36724, California, San Mateo County, T 5S, R 5W, 300 yd (274 m) north, 45? west of
the entrance (on Skyline Blvd.) to the Skyline Materials Plant No. 1; latest
Pliocene-earliest Pleistocene, Santa Clara Formation (Taylor, 1966a). Transferred to
Lithoglyphus by Taylor (1966a).
Fluminicola siegfusi Pilsbry, 1934:16 (Pilsbry, 1935, pi. 19: fig. 10). Lost Hills Oil Field
(California); San Joaquin Formation: Lost Hills Oil Field, the Universal Consolidated
Well No. 44, at 365-720 ft (111-219 m) (Pilsbry, 1935); Late Pliocene, upper part of
San Joaquin Formation, Kern County, California, T 26S, R 2IE, sec. 32 (Taylor,
1966a). Transferred to Savaginius by Taylor (1966a). Transferred to Brannerillus
(Planorbidae?) by Taylor (1975a).
Fluminicola spiralis Pilsbry, 1934:16 (Pilsbry, 1935, pi. 19: figs. 14-19). Twenty-three mi
(37 km) southeast of Hanford (California), 772-794 ft (235-242 m), and in the
McKittrick Front Oil Field; Tulare Formation, Boston Land Co. "C" well, 772-792 ft
(235-241 m), 23 mi (37 km) southeast of Hanford, sec. 27-19-18 (Pilsbry, 1935); Late
Pliocene, upper part of Tulare Formation, within or immediately above Corcoran Clay
member, Fresno County, California (Taylor, 1966a). Transferred to Savaginius by
Taylor (1966a).
Pilsbryus superbus Yen, 1944:105, fig. 8. Pliocene, Idaho Formation, Hammett, Elmore
County, Idaho; Late Pliocene-Early Pleistocene, Glenns Ferry Formation, Elmore or
Owyhee County, Idaho (Taylor, 1966a). Transferred to Lithoglyphus by Taylor
(1966a).
Pilsbryus antiquus utahensis Yen, 1947:273, pi. 43: fig. 7. Pliocene, Salt Lake Formation,
Cache/Box Elder Counties, Utah, T 13N, R 2W, sec. 15; Late Pliocene, Cache Valley
Formation, Box Edler County, Utah, USGS locality 20093 (Taylor, 1966a). Transferred
to Lithoglyphus by Taylor (1966a).
Fluminicola yatesiana utahensis Yen, 1947:273, pi. 43: fig. 6. Pliocene, Salt Lake
Formation, Cache/Box Elder Counties, Utah, T 13N, R 2W, sec. 15; Late Pliocene,
Cache Valley Formation, Box Edler County, Utah, USGS locality 20093 (Taylor,
1966a). Transferred to Savaginius by Taylor (1966a; as synonym of Paludestrina
nanna Chamberlin and Berry, 1933).
Fluminicola weaveri Yen, 1944:105, fig. 6. Pliocene, Idaho Formation, Hammett, Elmore
County, Idaho; Late Pliocene-Early Pleistocene, Glenns Ferry Formation, Elmore or
Owyhee County, Idaho (Taylor, 1966a). Transferred to Lithoglyphus by Taylor
(1966a).
Pyrgulopsis williamsi Hannibal, 1912:189-190, pi. VIII: fig. 29. Martin and Dudley's oil
well, SE !A sec. 32, T 26S, R 21E, Lost Hills; Late Pliocene, upper part of Santa Clara
Formation, Lost Hills Oil Field, Kern County, California (Taylor, 1966a). Transferred
to Savaginius by Taylor (1966a).
Amnicola yatesiana J.G. Cooper, 1894:171, pi. XIV: fig. 10. Pliocene, Santa Clara lake
beds, San Jos6 Mission, Santa Clara County; Late Pliocene, Santa Clara Formation,
Alameda County, California, ridge of gravel and alluvium at Mission San Jose,
probably in sec. 1, 2, 11, 12, T 5S, R 1W (Taylor, 1966a). Transferred to Fluminicola
by Pilsbry (1935). Transferred to Savaginius by Taylor (1966a).
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
B
FIGURE 4.?Shells of type and other historically important material of Fluminicola species. A, F. coloradensis,
holotype, USNM 526631 (shell height, 10.8 mm); B. F. dalli, lectotype, MCZ 2087 (3.1 mm); C. F. columbiana
(=F. fuscus), lectotype, ANSP 27767 (8.0 mm); D. Amnicola hindsi {=F. fuscus), lectotype, BMNH
1863.2.4.17A (8.7 mm); E, F. minutissimus, lectotype, ANSP 94273 (1.5 mm); F, F. modoci, lectotype, CAS
60798 (4.1 mm).
NUMBER 583 23
FIGURE 5.?Shells of Fluminicola species. A-D. F. coloradensis: A. USNM 160394 (shell height, 9.2 mm); B,
USNM 883493 (8.7 mm); C. USNM 526754 (6.5 mm); D. USNM 28103 (9.3 mm). E-G. F. dalli. USNM 883467
(2.3 mm, 2.4 mm, 2.1 mm, respectively); H-J, F. modoci, USNM 883468 (3.2 mm, 2.9 mm, 2.7 mm,
respectively); K.L, F. turbiniformis, USNM 883470 (3.1 mm, 3.2 mm, respectively).
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 6.?Shell protoconchs of Fluminicola species. A. F. coloradensis, USNM 526576 (bar = 0.27 mm); B. F.
dalli, USNM 526835 (bar = 231 \im); C. F.fuscus, USNM 883495 (bar = 0.33 mm); D, F. minutissimus, ANSP
398350 (bar =120 urn); E. F. modoci, USNM 883468 (bar = 200 urn); F. F. nuttallianus, USNM 47874
(bar = 0.33 mm); G. F. seminalis, USNM 883465 (bar = 231 urn); H. F. turbiniformis, USNM 883470
(bar = 250 \im); I. F. virens, USNM 883183 (bar = 0.27 mm).
NUMBER 583 25
FIGURE 7.?Opercula of Fluminicola species. A. F. coloradensis, USNM 883493 (bar = 1.2 mm); B, F. dalli,
USNM 874899 (bar = 0.38 mm); C, F.fuscus, USNM 883182 (bar= 1.2 mm); D, F. modoci. USNM 883468
(bar = 0.5 mm). Dorsal views to left, ventral views to right
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 8.?Radulae of Fluminicola species. A-C. F. coloradensis, USNM 883493 (bars = 38 Jim, 60 Jim, 136
Urn, respectively); D-F.F. da//i, USNM 874899 (bars = 17.6 Jim, 20\im, 60 urn, respectively); G-l,F. fuscus,
USNM 883182 (bars = 30 jun, 50 urn, 120 \un, respectively); J-L, F. modoci, USNM 883468 (bars = 20
\im, 30 urn, 50 Jim, respectively). Photographs show (from left to right) central teeth, lateral teeth, and general
view of portion of radula ribbon.
NUMBER 583 27
FIGURE 9.?Penes (dorsal views) of Fluminicola species. A, F. coloradensis, USNM 883493 (bar = 1 mm); B, F.
dalli. USNM 874899 (bar = 0.50 mm); C. F.fuscus, USNM 883182 (bar = 1 mm); D. F. modoci, USNM 883468
(bar = 0.5 mm); E. F. seminalis, USNM 883465 (bar = 0.5 mm); F. F. turbiniformis, USNM 883470 (bar = 0.33
mm); G. F. virens, USNM 883673 (bar = 1 mm). The penial duct is indicated by dotted lines. Epithelial pigment
is shown as dark stipple.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Ag
Cov
FIGURE 10.?Distal female genitalia (left side) of Fluminicola species. A, F. coloradensis. USNM 883493
(bar= 1 mm); B. F. dalli, USNM 874899 (bar = 0.5 mm); C. F.fuscus, USNM 883182 (bar= 1 mm); D. F.
modoci, USNM 883468 (bar = 0.5 mm). (Ag = albumen gland, Bu = bursa copulatrix, Cg = capsule gland,
Cgo = capsule gland opening, Cov = coiled oviduct, Dbu = bursal duct, Rf = rectal furrow, Sr = seminal
receptacle, Vc = ventral channel.)
NUMBER 583 29
Pa c ific
Ocean
11 I i<
FIGURE 11.?Map showing distribution of Fluminicola species ( ? = F. coloradensis, ? = F. fuscus,
modoci, ? = F. dalli, ? = F. minutissimus).
? =F.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 12.?Shells of Fluminicolafuscus. A, ANSP 62341 (shell height, 8.5 mm); B, USNM 883516 (7.3 mm);
C, USNM 473390 (9.2 mm); D. USNM 854693 (9.3 mm); E, USNM 883515 (7.8 mm); F. USNM 380804 (7.7
mm); G. USNM 883499 (8.7 mm); H. USNM 130527 (11.2 mm); I. USNM 883182 (10.0 mm); J. USNM 883640
(10.0 mm); K. USNM 511051 (8.8 mm); L. BMNH 1863.2.4.16(8.3 mm).
NUMBER 583 31
B
D
FIGURE 13.?Shells of type and other historically important material of Fluminicola species. A. F. nuttallianus,
lectotype, USNM 121467 (shell height, 8.8 mm); B, F. seminalis, possible syntype, BMNH 1994109 (7.7 mm);
C. LJthoglyphus cumingii (=F. seminalis), lectotype, BMNH 1933047 (5.8 mm); D. F. turbiniformis, lectotype,
ANSP 27779 (3.3 mm); E. F. virens, lectotype, USNM 121431 (7.5 mm); F, Paludina nuclea (=F. virens),
possible syntype, ANSP 27765 (8.3 mm).
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Pa c ific
Ocean
FIGURE 14.?Map showing distribution of Fluminicola species ( ? = F. nuttallianus,
? = F. seminalis, ? = F. virens).
= F. turbiniformis,
NUMBER 583 33
K
FIGURE 15.?Shells of Fluminicola species. A-F. F. seminalis: A, USNM 121465 (shell height, 8.0 mm); B,
USNM 883465 (6.0 mm); c. USNM 883188 (6.0 mm); D. USNM 883180 (6.2 mm); E. USNM 883742 (6.5 mm);
F. CAS 93862 (6.0 mm). G-N. F. virens: G. USNM 473989 (9.5 mm); H, CAS 93810 (5.8 mm); I. USNM 511057
(11.5 mm); J. USNM 511059 (10.3 mm); K. USNM 883673 (7.5 mm); L, USNM 883670 (10.3 mm); M, USNM
883676 (8.7 mm); N. USNM 883693 (7.7 mm).
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 16.?Opercula of Fluminicola species. A. F. seminalis, USNM 883465 (bars = 0.86 mm (left), 1 mm
(right)); B. F. turbiniformis, USNM 883470 (bar = 0.43 mm); C. F. virens, USNM 883673 (bars = 1 mm (left),
0.86 mm (right)). Dorsal views to left, ventral views to right.
NUMBER 583 35
FIGURE 17.?Radulae of Fluminicola species. A-C, F. seminalis, USNM 883465 (bars = 27 \im, 50 Jim, 100
Urn, respectively); D-F. F. turbiniformis, USNM 883470 (bars = 20 \un, 27 urn, 50 urn, respectively); G-I. F.
virens, USNM 883673 (bars = 25 ^m, 60 |im, 86 Jim, respectively). Photographs show (from left to right)
central teeth, lateral teeth, and general view of portion of radula ribbon.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
Ag
FIGURE 18.?Distal female genitalia (left side) of Fluminicola species. A. F. seminalis, USNM 883465 (bar = 1
mm); B. F. twbiniformis, USNM 883470 (bar = 0.5 mm); c, F. virens, USNM 883673 (bar=l mm).
(Ag = albumen gland, Bu = bursa copulatrix, Cg = capsule gland, Cgo = capsule gland opening, Cov = coiled
oviduct, Dbu = bursal duct, Sr = seminal receptacle, Vc = ventral channel.)
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Jones."
Arrange and paginate sequentially every sheet of manuscript
in the following order: (1) title page, (2) abstract, (3) contents, (4)
foreword and/or preface, (5) text, (6) appendices, (7) notes section,
(8) glossary, (9) bibliography, (10) legends, (11) tables. Index copy
may be submitted at page proof stage, but plans for an index should
be indicated when the manuscript is submitted.
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